​

Les ressources des vidéos de la Séquence 3 - Décrire

Sujet 1 & 2

Schémas
L'organisation générale d'une feuille est ici schématisée. Mais la morphologie des feuilles
varie d'une espèce à une autre. Découvrez et apprenez à nommer les différentes formes
que les feuilles peuvent prendre :
 par Hervé Sauquet

Les termes "pennée" et "palmée" peuvent désigner la nervation de la feuille mais aussi sa
forme, à ​voir ici​.

Les fleurs se présentent différemment en fonction de leur symétrie et de la position de

l'ovaire par rapport à l'insertion des pétales :

par Hervé Sauquet

Les plantes ont une vie sexuelle bien plus complexe que la nôtre et les fleurs d'une plante
peuvent s'organiser de bien des manières :

par Hervé Sauquet

La tomate n'est pas un légume ? La différence entre fruit et légume prend un tout autre sens
en botanique. Voici quelques exemples concrets :

([Link])

Site Internet
"Fruit ou légume ? Fin du débat !" sur ​[Link]

Vidéo
Découvrez la structure d'une inflorescence bien particulière : celle de l'Arum :
Ep.11 : L'Arum, prison dorée​ dans La Minute Nature de la Salamandre sur YouTube

Sujet 3

Vidéo
Comme l'a expliqué Thibaut, les plantes recherchent la lumière. Voici une vidéo (timelapse =
photos montées en film) où nous voyons bien ce phénomène que l'on appelle "tropisme" :
[Link]

Schéma
Le méristème est un tissu de cellules indifférenciées, c'est-à-dire non spécialisées, où la
division cellulaire est très fortement active. On retrouve principalement les méristèmes dans
les bourgeons de feuilles et les boutons de fleurs. Des tissus méristématiques
peuvent également se retrouver en bout de racine comme le présente ce schéma :

([Link])

Article
● Définition du "Méristème​" sur​ ​[Link]é[Link]

Sujet 4

Article
Rencontre avec la chercheuse Deborah Goffner, à l’origine de l’ambitieux projet « Future
Sahel » : ce projet, débuté par une traversée d’ouest en est du Sénégal, s’inscrit dans le
cadre de la Grande Muraille verte, qui a pour vocation de lutter contre la désertification de la
zone saharo-sahélienne :
● Louis Lise,​ ​Un futur plus vert pour le Sahel​, CNRS Le Journal, mai 2016.

Vidéo
Visionnez cet extrait qui vous montre le travail des scientifiques français et sénégalais pour
la régénération forestière intitulée «Grande Muraille Verte» :
● La Science et la grande muraille verte (extrait)​ sur Webcast
Film complet sur​ ​[Link]/doc=4668
 ​

Bibliothèque de la Séquence 3 - Décrire

Articles Encyclopedia Universalis

● Robert GORENFLOT, «​ ​FEUILLE​ ​», Encyclopædia Universalis [en ligne],
juillet 2016.
● Jean-Luc REGNARD, Roger ULRICH, Universalis, « ​FRUITS », ​Encyclopædia
​ Universalis [en ligne], juillet 2016.
● Georges DUCREUX, Hervé LE GUYADER, Jean-Claude ROLAND, «
DÉVELOPPEMENT (biologie) - Le développement végétal », ​Encyclopædia
​ Universalis [en ligne], juillet 2016.

Articles
● Claude Edelin, Daniel Barthelemy, Pierre Raimbault, "​Modèle architectural​" sur
[Link]
● Paul Fabre, "Observer et décrire une plante : l’appareil végétatif" (pdf)
● Paul Fabre, "Observer et décrire une plante : l'appareil reproducteur" (pdf)

● Valéry Malécot, "​Inflorescence​" sur​ ​[Link]

● Amadou TIDIANE BA, Kandioura NOBA, "​Flore et biodiversité végétale au Sénégal​",
Sciences et changements planétaires / Sécheresse, volume 12, numéro 3,
Septembre 2001.
● Rapport du colloque Patrimonialisation des ressources végétales au Sénégal​, Juillet
2012.

Sites Internet
● "​L'auxine, une phytohormone pas comme les autres​" sur​ ​[Link]
● "​Hormones végétales ou phytohormone​" sur le​ ​site de l'Université de Liège
● "​Les différents types de fruits​" sur​ ​[Link]
● Visitez le Conservatoire botanique Michel Adanson comme si vous y étiez :
[Link]/VV-endamadsahel/

Schémas
Chaque fleur a son diagramme et sa formule en fonction de sa composition florale. Voici
deux exemples de diagrammes floraux :
 réalisés par Hervé Sauquet

Cette méthode de rédaction du diagramme floral se base sur un standard international

formalisé dans le papier de Prenner et al. (pdf) et suivi par de nombreuses ressources tels
que l'ouvrage de Louis Ronse de Craene (Flora diagrams) et l'ouvrage de Simpson (Plant
Systematics).
Découvrez également un bref historique de Valéry Malécot qui vous permettra de
comprendre quelles ont été les étapes pour arriver à la rédaction des formules florales
actuelles ; méthode utilisée pour décrire les 10 familles botaniques de ce MOOC : Valéry
Malécot, "À propos des formules florales" (pdf), MOOC Botanique, Octobre 2016.

Livre
Disponibles en librairie ou sur Internet :
● Gilles BOETSCH, La grande muraille verte : Des arbres contre le désert,
Editions PRIVAT, mai 2013.
 Nombre de pièces florales
Verticilles de :

Image extraite du site : [Link]

a) 3 : fleur trimère
b) 4 : fleur tétramère,
c) 5 : fleur pentamère
d) n : fleur polymère

Soudure des pièces florales

Pour le périanthe
Les sépales peuvent être libres ou soudés entre eux :
-> calice dialysépale ou gamosépale

Les pétales peuvent être libres ou soudés entre eux :

-> corolle dialypétale ou gamopétale
…
dialystémone, gamostémone , dialycarpellé, gamocarpellé
Observer et décrire une plante
29/01/2015 Paul Fabre
 Symétrie de la fleur

Asphodèle fistuleux Ophrys marbré

Illustration tirée de laa présentation du TP de C. Heinz du 14/03/2013

Fleur actinomorphe Fleur zygomorphe ou
ou régulière irrégulière
Plusieurs plans de symétrie Un seul plan de symétrie

Observer et décrire une plante

29/01/2015 Paul Fabre
Position de l’ovaire vis-à-vis des autres pièces
florales

Illustration tirée de laa présentation du TP de C. Heinz du 14/03/2013

Partiellement enfoncé et/ou soudé dans réceptacle

Observer et décrire une plante

29/01/2015 Paul Fabre
 Inflorescence
= regroupement de fleurs sur un axe

Le racème ou grappe : croissance indéfinie, sens de floraison

centripète
pédicelle

bractée

pédoncule

Image extraite du site : [Link]

La cyme : croissance définie, sens de floraison centrifuge
Observer et décrire une plante
29/01/2015 Paul Fabre
Exemples de racèmes

Image extraite du site : [Link]

Observer et décrire une plante
29/01/2015 Paul Fabre
Inflorescence des Astéracées

Image extraite du site : [Link]

Observer et décrire une plante
29/01/2015 Paul Fabre
 Exemples de cymes

Cyme bipare

Image extraite du site : [Link]

Cyme unipare hélicoïde

Cyme unipare scorpioïde

Observer et décrire une plante

29/01/2015 Paul Fabre
 Adapté de : Stratégies végétales, B. Garrone et al., 2011
La feuille
• Définition : (en général)
-Organe plan, chlorophyllien qui pousse
latéralement sur la tige.
-Elle a un bourgeon à son aisselle (côté
supérieur de son point d’attache sur la tige)
- Ella a une croissance limitée.

• Fonctions :
- acquisition de l’énergie (photosynthèse)
- échanges gazeux (respiration : rejet C02 et
absorption d’O2, transpiration : rejet d’H2O …)

Observer et décrire une plante 15/01/2015 Paul Fabre

 La feuille
simple composée
marge
limbe

nervure

pétiole
stipule

nervation pennée

nervation palmée

crénelée dentée lobée

Observer et décrire une plante 15/01/2015 Paul Fabre
 Bourgeon apical

nœud
Entrenœud

Bourgeons axillaires

Observer et décrire une plante 15/01/2015 Paul Fabre

L’arrangement des feuilles : la phyllotaxie

Observer et décrire une plante 15/01/2015 Paul Fabre

A propos des formules florales

Valéry Malécot
Agrocampus Ouest, campus d’Angers
Octobre 2016

Le MOOC Botanique – initiation a pris le parti d’utiliser le format de rédaction

de formules florales proposé par Prenner et al. (2010). Cette rédaction pourra
sembler sensiblement distincte de celles qui sont utilisées lors de travaux
pratiques d’université ou de classes préparatoires en France. En guise d’élément
historique, diverses initiatives de rédaction de formules décrivant l’organisation
de la fleur ont eu lieu au cours du 19 ème siècle (Cassel 1820, Martius 1828,
Seringe & Guillard 1836, Morren 1852, Grisebach 1854, Alefeld 1862) mais c’est
l’introduction, en 1868, d’une forme de rédaction dans la première édition du
Lehrbuch der Botanik de Julius Sachs qui est la source principale de leur usage
puis de l’apparition de variantes. En effet, Sachs va développer cette partie dans
la seconde (1871) puis troisième édition (1873) de son ouvrage, et cette
troisième édition sera traduite en français par van Tieghem (Sachs 1874) et en
anglais par Bennet & Dyer (Sachs 1875). C’est lors de ses traductions que des
variantes de rédaction des formules florales ont été diffusées (au moins dans le
cas de van Tieghem cette variante dérive d’usages peu documentés mais issus
des initiatives antérieures). Ainsi, si Sachs écrit « K3 C3 A3+3 G3 » comme
formule florale pour les Liliaceae (c’est-à-dire en utilisant l’initiale du cycle
[K=Kelch, C=Corolle , A=Androeceum, G=Gynaeceum] devant le nombre de
pièces du cycle), van Tieghem indiquera pour des genres de la même famille
« 3S + 3P + 3E + 3E' + [3C] » (utilisant en cela l’initiale du nom de la pièce florale
[S=Sépale, P=Pétales, E=Etamine, C=Carpelle] précédée du nombre de pièces
dans le cycle), quant à Bennet & Dyer ils écrivaient « S3 P3 St3+3 C3 » comme
formule florale pour ces mêmes Liliaceae (utilisant les initiales des noms des
pièces florales [S=Sepals, P=Petals, St=Stamens et C=Carpels] mais devant le
nombre de pièces du cycle, ce dernier en indice). Je passe sur les raffinements
d’indication des soudures (parenthèses, voire crochets, au-dessus ou de part et
d’autre des pièces fusionnées) ou de l’indication d’une disposition spiralée et
autres détails. Les indications de la symétrie florale
(actinomorphe/zygomorphe) apparaitront plus tard, de même que celle de la
position de l’ovaire (infère/supère) ainsi que d’autres variantes de rédaction. Le
système retenu par Prenner et al. (2010) s’inspire du système initial de Sachs,
avec quantité de précisions qui rappellent certains de ces systèmes plus ou
moins anciens.

Et pour finir l’histoire, pour cette même famille des Liliaceae l’utilisation du
format de rédaction de Martius (1828) donne « 3ca 3co 3+3stam », celle de

Seringe & Guillard donne celle de Morren (1852)

donnerait « Cx 3 + Ca 3 + S 6 + P 3 », celle de Grisebach (1854) aboutit à
« 3, 3, 6, 3 » ; enfin celle de Alefeld (1862) donnerait, approximativement,
« 3•3•6•3‡ », tout ceci sans compter les variantes postérieures à 1875.

Bibliographie
Alefeld F., 1862. Ueber Formeln der Blüthentheile. Bonplandia 10 (18) : 275-
277.
Cassel F.P., 1820. Morphonomia botanica, sive Observationes circa
proportionem et evolutionem partium plantarum. DuMont-Schauberg,
Coloniae Agrippinae (Köhln).
Grisebach A.H.R., 1854. Grundriss der systematischen Botanik für akademische
Vorlesungen. Verlag der Dieterischen Buchhandlung, Göttingen.
Martius C.F.., 1828. Über die Architectonik der Blüthen. Isis von Oken 21 : 522-
529.
Morren C., 1852. Synanthies. L’institut, journal universel des sciences, 1 ère
section Sciences mathématiques, physiques et naturelles 20 (972) : 265-
268.
Prenner G., Bateman R.M., Rudall P.J., 2010. Floral formulae updated for routine
inclusion in formal taxonomic descriptions. Taxon 59 (1) : 241-250.
Sachs J., 1868. Lehrbuch der Botanik nach dem gegenwärtigen Stand der
Wissenschaft. Verlag von Wilhelm Engelmann, Leipzig.
Sachs J., 1870. Lehrbuch der Botanik nach dem gegenwärtigen Stand der
Wissenschaft. Zweite Auflage. Verlag von Wilhelm Engelmann, Leipzig.
Sachs J., 1873. Lehrbuch der Botanik nach dem gegenwärtigen Stand der
Wissenschaft. Dritte Auflage. Verlag von Wilhelm Engelmann, Leipzig.
Sachs J., 1874. Traité de Botanique conforme à l’état présent de la science,
traduit de l’allemand sur la 3ème édition et annoté par Ph. Van Tieghem.
Librairie F. Savy, Paris.
Sachs J., 1875. Text-book of botany, morphological and physiological, translated
and annotated by Alfred W. Bennett assisted by W.T. Thiselton Dyer.
Clarendon Press, Oxford.
Seringe N.C., Guillard A., 1836. Essai de formules botaniques représentant les
caractères des plantes par des signes analytiques qui remplacent des
phrases descriptives. J. Albert Mercklein, Paris.
TAXON 59 (1) • February 2010: 241–250 Prenner & al. • Floral formulae in taxonomic descriptions

Floral formulae updated for routine inclusion in formal taxonomic

descriptions
Gerhard Prenner,1 Richard M. Bateman1,2 & Paula J. Rudall1

1 Jodrell Laboratory, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3DS, U.K.
2 Department of Geography, Environmental and Earth Sciences, University of Birmingham, Edgbaston, Birmingham, B15 2TT, U.K.
Author for correspondence: Gerhard Prenner, [Link]@[Link]

Abstract Building on two centuries of history, we advocate an expanded and maximally informative format for floral formulae
to describe angiosperm flowers in formal taxonomic descriptions. Using standard typeface and Unicode character codes, the
format can summarise a wide range of features, including: acropetal sequence of organ initiation, number and symmetry of each
whorl of floral organs (bracts, sepals, petals [or tepals], androecium, gynoecium, ovules), position of the organs relative to each
other, partial and/or complete fusion of organs, resupination, organ loss and suppression, and deviations from standard bisexual-
ity. We use several complex flowers to demonstrate our view that all known flower morphs can be accurately represented by the
updated format, and show how use of floral formulae can outline some unanswered questions regarding the floral organisation of
the model organism Arabidopsis. Possible uses of floral formulae extend beyond static description of mature morphology into the
comparative realm. When combined with phylogenies, they help to elucidate generalised groundplans, plesiomorphic architecture,
and the location and polarity of particular character-state transitions. They can usefully be applied within as well as between
species, to compare wildtype versus mutant morphs and different ontogenetic stages. Perhaps their greatest strengths are that
they (1) require only a universally available typescript, and (2) bring a comprehensive uniformity to the description of flowers,
acting as a checklist of features to be examined. We recommend that floral formulae become a routine component of diagnoses
in protologues and other formal taxonomic (re)descriptions, functioning as a logical phenotypic counterpart to the DNA barcode.

Keywords Antirrhinum; Arabidopsis; Caesalpinioideae; floral diagram; floral formula; floral morphology; flower;
Lecythidaceae; Orchidaceae; protologue; taxonomic description

INTRODUCTION extremely useful, cannot be inserted into standard text. We

suggest that floral formulae represent a simple and user-
Two contrasting but complementary methods were devel- friendly morphological counterpart to the DNA barcode.
oped during the nineteenth century to represent the fundamen- Our pragmatic goals in preparing this advocacy were (1)
tal architectures of angiosperm flowers. The pictorial approach to stimulate discussion in order to achieve a consensus format
was termed the floral diagram, whereas the textual approach among the botanical community for the floral formula, and
was termed the floral formula (examples are given in Fig. 1). (2) to encourage the architects of the International code of bo-
Both approaches were developed during the nineteenth cen- tanical nomenclature to specify minimum levels of character-
tury when they achieved a reasonable level of usage in main- based information (including floral formulae) required in any
stream literature (cf. Grisebach, 1854; Sachs, 1873; Eichler, formal taxonomic description.
1875, 1878; Goebel, 1887). However, use of floral diagrams
declined during the twentieth century, and floral formulae
travelled even further into obscurity, despite the fact that they HISTORICAL DEVELOPMENT OF FLORAL
convey a great deal of information in a highly compact form FORMULAE
using only standard typeface.
Here, we update the floral formula for a wide range of Although the first attempt to describe floral structure via
twenty-first century uses, and argue that the floral formula floral formulae was made by Cassel (1820), the first formulae
should be routinely included in formal (re)descriptions of an- that resemble those used today were developed by Martius
giosperm taxa. We broadly follow traditional iconography, (1828). Grisebach (1854) made especially extensive use of flo-
but also propose some modifications based on symbols that ral formulae. For many plant families and genera he annotated
have become widely available. A logical conclusion of the a much-reduced formula in which he gave the number for each
current drive to establish DNA barcodes to provide species floral whorl from the periphery to the centre of the flower
identifications and accelerate species discovery (e.g., papers (e.g., he annotated for Ranunculaceae “5, 5, ∞, ∞”, indicating
in Savolainen & al. 2005) should be inclusion of such barcodes 5 sepals, 5 petals, many stamens and many carpels). Sachs
in the protologue describing new species. However, even the (1873) also made extensive use of both floral diagrams and
most avid supporters of barcoding agree that this approach floral formulae in his influential Lehrbuch der Botanik (for a
will not work efficiently for all plants. Floral diagrams, while translation of Sachs’ book see Goebel, 1887). He highlighted

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advantages of the formula over the diagram, noting that the A STANDARDISED FLORAL FORMULA FOR
former can be printed in “gewöhnlichen Typen” (i.e., ordinary THE TWENTY-FIRST CENTURY
typescript: Sachs, 1873: 519) and – perhaps more importantly
– is capable of broader generalisation, because the figures can Building on the principles established by Sachs (1873), and
be replaced by letters as numerical coefficients. after considerable discussion, we have developed an agreed set
Sachs (1873: 520) also developed a convention for gener- of guidelines and a resulting standardised format for present-
alised formulae. For example, the formula for Monocotyle- ing floral formulae. We believe that this protocol will serve
donae, Kn Cn An + n Gn (+ n), indicates that their flowers are the needs of the many diverse uses to which floral formulae
typically composed of five alternating whorls, each with the could (and should) be applied. We now briefly consider the suc-
same number of organs, and that two whorls are expressed as cessive elements of the suggested common formula (Table 1):
perianth whorls, two as staminal whorls and one usually as Floral symmetry.  — Following the terminology of floral
a carpellary whorl; the parenthetic (+ n) at the end of the for- symmetry laid down by Endress (1999, 2001), the following
mula indicates that there is occasionally a second carpel whorl symbols are available to denote the main categories of floral
present (note that the parentheses here are not used to annotate symmetry: “ * ” for polysymmetry ( = radial symmetry; with
fusion, but rather to indicate an alternative condition). Most > 2 symmetry planes), “↓” for median monosymmetry ( = zy-
petaloid monocots would have a value of 3 for “n”. gomorphy = dorsiventral symmetry; with a single symmetry
Eichler’s (1875, 1878) Blüthendiagramme set an admirable plane in median direction), “→” for transversal monosym-
standard in comparative floral morphology by making exten- metry (single symmetry plane in transverse direction), Ø”
sive use of floral diagrams. In contrast, he was surprisingly for oblique monosymmetry (single symmetry plane oblique),
cautious in employing floral formulae, using them only to “ ┼ ” for disymmetry (with two symmetry planes; note that we
summarise the “Typus” of families with relatively simple flow- prefer the symbol “ ┼ ” over the frequently used “ + ” because
ers. His approach emphasises the fact that floral formulae have the latter is used here to separate different whorls of similar or-
never been universally accepted by the botanical community. gans), and “∂  ” for true asymmetry (lacking symmetry planes;
Sattler (1973) described the ontogeny of 50 plant species, here we propose the symbol for a partial differential, because it
annotating these taxa with floral diagrams and floral formulae. is readily available on PCs and therefore more accessible than
Interestingly – and typical of his philosophical viewpoint – other previously suggested symbols).
Sattler (1973: xvii) noted that “the floral formula is interpre- Rudall & Bateman (2002: 435) argued that floral formulae
tative, based on the conception that the flower is a modified can convey a more precise explanation of floral symmetry by
monaxial shoot without axillary buds. Since this idea of the reporting the symmetry of each successive organ whorl in the
flower is the predominant one today and usually the only one flower, rather than the entire flower, using a superscript that
dealt with in textbooks, the inclusion of the floral formula may represents the number of observed symmetry planes. Accept-
facilitate understanding of the organogenetic description for ing their objective but modifying their suggested method, we
those who have received classical botanical training. Those here propose to add the above-mentioned symbols for each
who do not wish to be biased by the interpretative nature of organ category directly after the whorl that it qualifies. This
the floral formula … should disregard the floral formula.” follows the logic of the analytical method, in which the organ
Most of the recent uses of floral formulae have provided whorl is first identified, then quantified and finally qualified.
relatively brief summaries that served merely to convey the For example, the cypripedioid orchid shown in Fig. 1A has a
typical number of organs present in each of the three or four monosymmetric sepal whorl, a monosymmetric petal whorl
categories of whorl. Ellstrand (1983: 119) mentioned only that vertically bisecting the labellum, a monosymmetric andr-
the “5-5-5-3” floral formula (5 sepals, 5 petals, 5 stamens, oecium vertically bisecting the column, and a polysymmetric
3 carpels) of Ipomopsis aggregata is both remarkably con- inferior ovary. The “overall symmetry” of the mature (anthetic)
stant throughout its genus and also almost invariant in its flower, as it appears to floral visitors, can if required be an-
family, Polemoniaceae. Tian & al. (2007: 262) summarised notated using the appropriate signature at the very beginning
the flowers of the tepallate Circaeaster agrestis (Circaeast- of the floral formula.
eraceae) by presenting “the number of the floral organs by Organ abbreviation and sequence.  — We have adopted
using floral formulas Pn A nGn”, where the subscripts represent as the basis of our format the four fundamental categories of
organ number. Judd & al. (2007) made extensive use of flo- whorl that characterise eudicots. These four categories formed
ral formulae in their popular textbook on plant systematics, the basis of the typically quadripartite ‘KCAG’ (or, in appro-
though their formulae are highly simplified (without use of priate cases, the tripartite ‘PAG’) floral formulae advocated
capitals for different organs) and are less accessible via nor- by Sachs (1873). They are listed centripetally (acropetally)
mal typescript. Méndez & Gómez (2006: 225) generalised from the periphery to the centre (base to apex) of the flower
that “floral structure in angiosperms is usually represented along the floral axis: K calyx (sepal number), C corolla (petal
by means of a floral formula, which summarises the number number), A androecium (stamen number), G gynoecium (car-
and arrangement of floral organs (sepals, petals, stamens and pel number). We indicate stamen fascicles using superscripts.
carpels).” In contrast, we are proposing the consistent use of For example, the three fascicles, each consisting of many sta-
more comprehensive floral formulae with a far wider range mens, of Hypericum perforatum are given as A3∞ (for discus-
of applications. sion of the morphology and homologies of stamen fascicles see

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Table 1. Symbols used in updated floral formulae (symbol description plus Unicode character code in parentheses; see text for a
detailed discussion of each symbol).
Symmetry:
↓ median monosymmetry (downwards arrow; Unicode: 2193)
→ transverse monosymmetry (sidewards arrow; Unicode: 2192)
Ø oblique monosymmetry (Latin capital letter O with stroke; Unicode: 00D8)
┼ disymmetry (box drawings light vertical and horizontal; Unicode 253C)
* polysymmetry (asterisk)
∂ asymmetry (partial differential; Unicode 2202)

Organs:
B bracteate (flower in the axil of subtending bract)
Bt bracteolate (flower preceded by bracteole/s)
K calyx (number of sepals) (superscript = sepaloid organ)
C corolla (number of petals) (superscript = petaloid organ)
P perigon (number of tepals)
A androecium (number of stamen)
∞
AX stamen fascicles (X = number of fascicles, highlighted with superscript infinity symbol)
AX↔X obdiplostemonous androecium (X = number of stamens per whorl, connected by “left right arrow”; Unicode 2194)
AX↔ obhaplostemonous androecium (X = number of stamens followed by “left right arrow”; Unicode 2194)
G gynoecium (number of carpels)
G superior ovary
-G- half-inferior ovary
Ĝ inferior ovary (capital G with circumflex; Unicode 011C)
V number of oVules per ovary
Va apical placentation
Vb basal placentation
Vc free central placentation
Vm marginal placentation
Vp parietal placentation
Vx axile placentation
® resupination (registered sign; Unicode 00AE)

Others
∞ “many” organs (i.e., >12) (infinity; Unicode 221E)
– defined range of organ number (en-dash; Unicode 2013)
+ (plus) connects different whorls of the same organ category
: (colon) separates morphologically contrasting organs within one organ whorl
(x), [x], {x} fusion of different organs (different kinds of bracket allow for a nested series of up to three levels of fused organs)
r
organ reduction (superscript r)
0
organ loss (superscript zero)
♀ female (pistillate) flower (female sign; Unicode 2640)
♂ male (staminate) flower (male sign; Unicode 2642)

Prenner & al., 2008). In groups that possess closely-spaced We here argue for the addition of ovules as a fifth category
whorls of morphologically similar sepals and petals (such as of organ (see also Sattler, 1973). To avoid confusion of the letter
many monocots), both K and C are replaced with P for tepals O with a zero, we suggest that oVules should be symbolised
(the ‘P’ represents a perigon – one or more undifferentiated by V, yielding a pentapartite ‘KCAGV’ format. In addition
whorls subtending the inner floral organs). to the number of ovules per ovary, we highlight the type of

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placentation (Va = apical, Vx = axile, Vb = basal, Vc = free superscript terms. For example, in the legumes, the exception-
central, Vp = parietal, Vm = marginal placentation). ally complex five-petalled corolla of the typical papilionoid
If required, the presence or absence of flower-subtending flag blossom is designated as C1flag : 2wing : (2)keel↓ (note that the
bracts (B) and/or flower-preceding bracteoles (Bt) can be an- two keel petals are the only petals that are fused, here indicated
notated at the beginning of the floral formula, immediately by parentheses; see following paragraph).
prior to K or P (for discussions of these organs and their value Organ fusion.  — Fusion of different organs is indicated
in floral ontogeny and inflorescence terminology see Prenner, using various kinds of bracket. If several organs from differ-
2004a; Prenner & al., 2009). Similarly, it is possible to high- ent whorls are fused, this can be depicted using a specified
light the existence of an epicalyx or calyculus at the beginning sequence of three different types of brackets: “(x)” followed by
of the floral formula; for example ‘B3’ in Tofieldia. Each of “[x]” followed by “{x}”. If the fusion is restricted to the basal
these lettered abbreviations is followed by the number of or- or apical region of the specified organs, brackets are given in
gans present in that whorl, or in the case of ovules, the number subscript or superscript, respectively. For example, in the typi-
per gynoecium. An observed range of organs is indicated with cal disc flower of Asteraceae, the shared corolla–filament tube
an en-dash (e.g., A3–6). If there are “many” organs the infinity and apically fused anthers are annotated as follows: [C(5) A(5)]
symbol “∞” is traditionally used. We suggest that an appropri- (see more detailed discussion below under “Within-species
ate threshold for introducing this symbol would be more than applications: mutants and models”).
twelve organs within the whorl in question (greater numbers Ovary position.  — Ovaries can be either superior (with the
can be given if appropriate). outer floral organs inserted at the base of the ovary), inferior
Annotation within a category of organ.  — If more than (with the outer floral organs inserted on top of the ovary, which
one whorl is differentiable within a particular organ category, appears as if the ovary is congenitally “sunken” into the flower
then the different whorls of a single organ category are con- base) or semi-inferior (with the outer floral organs inserted at
nected with the plus symbol “ + ”, scoring the outer whorl or near the middle of the ovary; see Endress, 1994; Soltis &
first. For example, the early-divergent orchid Neuwiedia has Hufford, 2002; Soltis & al., 2003). We here propose to annotate
an outer androecial whorl of three stamens, two suppressed, superior ovaries by underlining the “G” for gynoecium (“G”),
and an inner whorl of three stamens, one suppressed, giving inferior ovaries by inserting the letter G with circumflex (“Ĝ”)
“A20 : 1 + 10 : 2” (cf. Kocyan & Endress, 2001). and semi-inferior ovaries with hyphens placed immediately
Where further differentiation is evident within one whorl, before and immediately after the letter G (“-G-“).
these morphs can be distinguished using the colon symbol. Dioecy and related reproductive architectures.  — As
Within each whorl, the annotated organ sequence begins with already noted, the development of floral formulae has been
the adaxial (typically upper) side of the flower, followed by strongly influenced by the classic hermaphrodite eudicot flower
the organs on the abaxial (typically lower) side of the flower. that gave us the equally classic ‘ABC’ model of floral devel-
For example, in a typical flower of the mint family (Lam- opmental control (e.g., Theissen & al., 2002). However, there
iaceae) and in the classic model organism Antirrhinum, two exist more complex spatial arrangements where male and fe-
petals form the upper lip and three petals form the lower lip. male reproductive functions are separated in different flowers
Such cases are annotated as C(2upper lip : 3lower lip)↓. Note that (monoecy) or even on different individuals (dioecy), or where
all five petals are basally fused; this feature is symbolised male or female function is suppressed in some but not all flow-
by the subscript brackets (see discussion below for details). If ers (gynodioecy and androdioecy respectively) (Barrett, 2002).
the adaxial organs are oriented lowermost as a result of 180° For separate male (staminate) and female (pistillate) flow-
rotation of the pedicel and/or ovary (i.e., resupination, as seen ers, we recommend two separate floral formulae prefixed by
in the majority of orchids), adaxial numbers should still be ♀ or ♂, and linked by the word “plus”.
given before abaxial (i.e., following the “original” orientation),
but the KCAGV floral formula should be prefixed with the
“registered” sign “®” (Fig. 1A, B). EXAMPLES OF FLORAL FORMULAE FOR
We view an obdiplostemonous androecium as one in which COMPLEX FLOWER ARCHITECTURES
the outer stamen whorl is located opposite the petals and hence
the law of alternating whorls is broken (contrary to the defini- It is tempting to use flowers with simple ratios of read-
tion of Bachelier & Endress, 2009: 502). Obdiplostemony can ily distinguished organ whorls to illustrate our format, such
be annotated with “↔” between the whorls, as in our example as Marsh Crane’s-bill, Geranium palustre (Fig. 1C, D), with
of Geranium (Fig. 1C, D) where the androecium is annotated five sepals, five petals, ten stamens in two whorls (arranged
as “A5↔5”. If there is only one whorl of stamens and the law obdiplostemonously) and a superior ovary of five fused carpels
of alternating whorls is broken (i.e., the androecium is obhap- and ten ovules arranged in axile placentation:
lostemonous), as in Primulaceae, the location of the stamens
opposite the petals can be indicated as “A5↔” (i.e., omitting K5* C5* A5↔5* G(5)* Vx10
the position of the second whorl of stamens).
Organ reduction is annotated as superscript “r”, organ loss However, we are anxious to demonstrate that the format
as superscript “0”. If further differentiation of floral organs can readily accommodate more complex floral morpholo-
within a single whorl is required, it is possible to resort to gies exhibiting less well-differentiated and/or partially fused

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TAXON 59 (1) • February 2010: 241–250 Prenner & al. • Floral formulae in taxonomic descriptions

Fig. 1. Flower (A) and floral diagram and floral formula (B) of Cypripedium calceolus, a cypripedioid orchid (A by G. Prenner, B modified from
Rudall & Bateman, 2002). Flower (C) and floral diagram and floral formula (D) of Geranium palustre (Geraniaceae) (C with permission from
[Link]; D modified from Eichler, 1878).

whorls (Fig. 1A, B). We have chosen our examples from the (3) fusion of the anthers that form a tube, superscript “ (X)”,
Asteraceae, Orchidaceae and Papilionoideae, the last of which through which the style will later emerge in order to secondar-
is not readily convertible to a floral formula, according to ily present the pollen. Sepals are transformed into specialised
Weberling (1989: 19). hairs or scales (superscript “pappus”) and the inferior ovary
The main challenge posed by typical Asteraceae is the ba- consists of a pseudomonomerous gynoecium of two fused
sal and/or proximal fusion of floral organs. Use of superscript carpels with a single ovule in basal placentation. The floral
and/or subscript brackets allows representation of (1) fusion formula for a typical radially symmetrical Asteraceae disc
of the five petals to form a corolla tube, shown as “C(5)”; (2) flower is therefore:
formation of a corolla–filament tube by fusion of stamen fila-
ments with basal parts of the corolla tube, subscript “ [X]”; and Kpappus* [C(5)* A(5)*] Ĝ(1 : 1r  )* Vb1

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Prenner & al. • Floral formulae in taxonomic descriptions TAXON 59 (1) • February 2010: 241–250

In orchid flowers, the various elements of the androecium comparative biology. Although comparison is most commonly
and gynoecium are fused into a single structure termed the confined to mature (anthetic) morphs representing different
gynostemium (column), and three to five of their presumed species, the method is equally applicable to comparisons
ancestral number of six stamens are suppressed (e.g., Rudall within species, either of contrasting mutant floral morphs (e.g.,
& Bateman, 2002, 2004). Orchids also show considerable di- Bateman & Rudall, 2006; Nutt & al., 2006) or of contrasting
versification within particular organ whorls, notably the dif- ontogenetic stages (e.g., Tsou & Mori, 2007).
ferentiation of a labellum from among the three petals. We Inferring groundplans.  — Eichler’s (1875) conservatism
here take as our example the resupinate flower (annotated with in deploying floral formulae usefully highlights two different
the prefixed registration sign) of Lady’s Slipper, Cypripedium uses of floral formulae, distinguishing empirical formulae
calceolus (Fig. 1A, B): that accurately reflect the actual appearance of one particular
depicted taxon from theoretical floral formulae that aim to
® K(2) : 1↓ C1labellum : 2↓ [A30 + 10 : 2↓ Ĝ(3)*] Vp∞ convey the fundamental essence of a range of evolutionarily
related floral morphologies. ‘Essential’ formulae are some-
Lastly, the floral formula for a typical papilionoid ‘flag times termed a floral groundplan. Motivations for inferring
blossom’ can be translated into a floral formula: groundplans differ. Some authors are seeking the ‘ancestral’
(in modern parlance, plesiomorphic) condition for the clade
K(5)↓ C1flag : 2wing : (2)keel↓ A(5 + 4) : 1↓ G1↓ Vm1–∞ under scrutiny. Others are seeking a more dynamic compari-
son, either between putative plesiomorphic and apomorphic
The different types of petals (adaxial flag, lateral wing pet- mature groundplans (perhaps exploring gains or losses of or-
als and abaxial keel petals) are annotated in superscript. The gans) or, less commonly, between juvenile and mature floral
chosen formula denotes a species in which five stamen filaments architectures in an ontogenetic series within a single species.
of the outer whorl and four stamen filaments of the inner whorl Groundplans are best inferred by vertically tabulating flo-
are fused to an adaxially open sheath, whereas the tenth stamen, ral formulae (Tables 2, 3), which is the most effective avail-
located in the adaxial position, remains free (as in the French able method of highlighting the architectural differences that
bean, Phaseolus vulgaris). In brooms (Genista and its relatives), distinguish between different floral morphs. For example,
all ten stamen filaments form a closed filament tube, annotated in Table 2 we have summarised floral formulae distributed
as A(5+5), whereas in other species of the same subfamily, such throughout the text of the recent study of floral diversity in the
as the pagoda tree (Styphnolobium japonicum), all ten stamens legume subfamily Caesalpinioideae by Prenner & Klitgaard
remain free, annotated as A5+5 (for details see Prenner, 2004b). (2008), who demonstrated that even the most complex floral
The gynoecium consists of a single superior ovary that contains morphologies found in this group, such as that of the early-
one to many ovules in marginal placentation. divergent Duparquetia orchidacea, can easily be encapsulated
In summary, we are confident that any flower can be con- in the form of a floral formula (Table 2). Displaying all rel-
verted into a floral formula with little if any loss of architec- evant formulae together in a tabular format enables the reader
tural information. to easily track trends in flower evolution within this florally
diverse subfamily. There is a widely accepted legume floral
groundplan of K5 C5 A5 + 5 G1 Vm1–∞, but the caesalpin-
THE DIVERSE USES OF FLORAL FORMULAE ioids show great diversity within this groundplan, using as
their main evolutionary theme the fusion and/or suppression
Comparative biology.  — Thus far, we have discussed of organs (Table 2).
floral formulae as a means of summarising isolated, static In Table 3 we have summarised floral formulae distributed
floral morphologies. In fact, they have an additional role in throughout the text of the recent study of floral organogeny and

Table 2. Floral formulae of selected Caesalpinioideae compared with the floral groundplan of Leguminosae (derived from
Tucker, 1998, 2000, 2002; Prenner & Klitgaard, 2008).
Caesalpinioideae
Duparquetia orchidacea B Bt K4C:10↓ C3:2r↓ A(4):10+50↓ G1↓ Vm2–5
Cercis canadensis B Bt K(5)↓ C5↓ A5+5↓ G1↓ Vm∞
Petalostyles labicheoides B Bt K3:(2)C↓ C5↓ A3:2r+50↓ G1C↓ Vm∞
0 0 0
Labichea lanceolata B Bt K3:(2) ↓ C4:1 ↓ A2:3 +5 ↓ G1↓ Vm2–3
Dialium guineense B Bt K5↓ C1:40↓ A2:30+50↓ G1↓ Vm2
C C r r r 0
Tamarindus indica B Bt K3:(2) ↓ C3:2 ↓ A(3):2 ↓+4 :1 G1↓ Vm8–10
Brownea latifolia B BtC K3:(2)C↓ C5* A(5+4):10↓ G1↓ Vm∞
C C r 0
Amherstia nobilis B Bt K3:(2) ↓ C3:2 ↓ A(5+4):1 ↓ G1↓ Vm1–6
Leguminosae groundplan B Bt K5↓ C5↓ A5+5↓ G1↓ Vm1– ∞

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TAXON 59 (1) • February 2010: 241–250 Prenner & al. • Floral formulae in taxonomic descriptions

diversity in Lecythidaceae by Tsou & Mori (2007). Although of angiosperm flowers. In addition, mapping floral formulae
each whorl, most notably the gynoecium, exhibits variation across the terminal branches of a phylogeny, whether mor-
between the scored taxa, the most common formula is K6 C6 phological or molecular, readily identifies positions on the
A∞ Ĝ(2–8) Vx1–∞. It is tempting to view this as a fundamental tree at which specific transitions in floral morphology take
groundplan that provided the basis for evolutionary excur- place, noting any potentially correlated shifts occurring on the
sions into monosymmetry associated with smaller numbers same branch of the tree. It also determines the polarity of the
and fusion of the sepals, and with deviations from the more transitions. For example, Rudall & Bateman (2002: figs. 13,
common number of six petals (Table 3). This is of course only 14) mapped shifts in partial and complete suppression of be-
a first approximation which can act as a testable hypothesis in tween three and five of the six plesiomorphic stamens among
a phylogenetic background. subfamilies of Orchidaceae, thereby clarifying how this suite
The opportunity exists in vertical tabulation to place first of characters had evolved within the family.
a full reference formula and then in subsequent formulae use a Within-species applications: ontogenetic series.  — The
full stop to indicate identical codings to the reference formula, most extensive use of floral formulae in recent literature was
annotating only deviations from the reference formula (thus by Tsou & Mori (2007) in their study of floral organogeny
mimicking the tabulation of nucleotide and amino acid data and diversity in Lecythidaceae. Inspired by Sattler (1973),
in comparative molecular studies). the authors included symbols for postgenital fusion “<x>”
Phylogenetic applications: inferring plesiomorphic ar- and congenital fusion “(x)”. Furthermore, they created an
chitectures and character evolution.  — A modern researcher “ontogenetic floral diagram” to convey information on organ
studying groundplans is most likely to be seeking a plesiomor- sequence and changes of floral symmetry during the course of
phic ‘ur-architecture’ for a particular clade of species. This is floral ontogeny (see also Sattler, 1973). Such changes of sym-
best inferred by considering floral morphological characters metry patterns during development were already highlighted
within a phylogenetic framework based on morphological by Endress (1999), who listed and discussed a broad range of
and/or molecular data. In theory, a floral formula could itself examples of shifts in symmetry during ontogeny. We believe
be viewed as constituting as a single phylogenetic character, that such changes during ontogeny remain under-studied, and
paralleling similar discussions regarding whether sequences that the translation into ontogenetic formulae could yield a
obtained from a single genic region constitute a single ‘hyper- better understanding of floral symmetry in general. Another
multistate’ character (e.g., Doyle, 1992). However, we would potentially fruitful excercise would be to annotate different
argue that, like a DNA sequence, a floral formula encompasses states of paedomorphosis during ontogeny into the format of
too much information to act as a single, highly multistate, an ontogenetic floral formula.
character in a morphological cladistic matrix. Within-species applications: mutants and models.  —
The greatest strength of floral formulae lies in provid- Although floral formulae have traditionally been used at the
ing an ideal checklist for characters that can be inserted into species level and above, they are equally applicable to any
normal text, and should be coded in any phylogenetic study situation where comparison reveals a substantial qualitative

Table 3. Floral formulae in subfamilies Planchonioideae and Lecythidoideae (Lecythidaceae) (modified from Tsou &
Mori, 2007; ovule numbers after Prance & Mori, 1977, 1979).
Planchonioideae
Barringtonia racemosa B Bt K(4)* C4* A(∞)* Ĝ(2–4)* Vx2–3
Lecythioideae
Grias peruviana B Bt K(4)* C4* A(∞)* Ĝ(4–5)* Vx2–4
Gustavia macarenensis B Bt K(4)* C8* A∞* Ĝ(4–8)* Vx7–93
Couroupita guianensis B Bt K6↓ C6* A∞↓ Ĝ(6–7) Vx30–115
Cariniana domestica B Bt K6↓ C6* A∞↓ Ĝ(3) Vx13–25
Cariniana micrantha B Bt K6↓ C6* A∞↓ Ĝ(3) Vx13–25
Cariniana decandra B Bt K(5)* C5* A(8–10)* Ĝ(3) Vx∞
Allantoma lineata B Bt K(5)* C5* A(30)* Ĝ(4)* Vx∞
Couratari sandwithii B Bt K6↓ C6↓ A∞↓ Ĝ(3)* Vx?
Bertholletia excelsa B Bt K6↓ C6↓ A∞↓ Ĝ(4)↓ Vx5
Corythophora amapaensis B Bt K6↓ C6↓ A∞↓ Ĝ(3–4)↓ Vx5–8
Eschweilera rankiniae B Bt K6↓ C6↓ A∞↓ Ĝ(2)↓ Vx∞
Lecythis pisonis B Bt K6↓ C6↓ A∞↓ Ĝ(4)↓ Vx10–25
Generalised groundplan B Bt K6 C6 A∞ Ĝ(2–8) Vx1– ∞

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Prenner & al. • Floral formulae in taxonomic descriptions TAXON 59 (1) • February 2010: 241–250

shift in floral morphology. For example, Bateman & Rudall CONCLUSIONS: THE FUTURE OF FLORAL
(2006) explored naturally occurring floral mutants of orchids, FORMULAE
comparing mutant with wildtype morphs in the hope of bet-
ter understanding the basis of floral evolution in the family. The floral formula is a powerful didactic tool.  — A flo-
Comparing two varieties of the Green-flowered Helleborine, ral formula constitutes a compact, consistent, unified, semi-
Epipactis phyllanthes, yields the following contrast: quantitative description that requires only universally available
typescript and Unicode character code. As such, it is a powerful
Epipactis phyllanthes var. vectensis: tool to encourage consistency of floral description, effectively
® ↓ K3* C1labellum : 2↓ [A20 : 1 + 30↓ Ĝ(3)*] Vp∞ offering a checklist of available floral components and generat-
ing what might be termed an ‘identikit’ flower (especially in
Epipactis phyllanthes var. phyllanthes: combination with floral diagrams). Preparing floral formulae
® * K3* C3* [A20 : 1 + 30↓ Ĝ(3)*] Vp∞ encourages both more rigorous analysis of floral form and use
of an explicit comparative approach (e.g., Stützel, 2002; Leins
This comparison highlights a transition within the corolla & Erbar, 2008). Once the few basic principles governing the
between monosymmetric and polysymmetric floral morphs. format of a floral formula have been learned, formulae also be-
Similarly, we can compare the orchid Cephalanthera damaso- come a valuable teaching tool, encouraging students to examine
nium with a contrasting morph that was originally described more closely the flowers put before them (e.g., Burrows, 2009).
as a novel genus by Chen (1965) but was reinterpreted as a Floral formulae are more concise than floral dia-
homeotic mutant of Cephalanthera by Bateman & Rudall grams.  — Some observers may argue that floral diagrams
(2006): convey considerably more information regarding floral ar-
chitecture than do floral formulae (Fig. 1). We consider both
Cephalanthera damasonium: floral formulae and floral diagrams to be under-used tools
® ↓ K3* C1labellum : 2↓ [A20 : 1 + 30↓ Ĝ(3)*] Vp∞ in systematic studies of angiosperms, viewing them not as
alternatives but as mutually supportive and complementary
‘Tangtsinia’ nanchuanica: methods of summarising floral architectures. Our revised
        * K3* C3* [A3 + 30* Ĝ(3)*] Vp∞ format for floral formulae upgrades the information content
and flexibility of floral formulae. Only the relative sizes and
Here, another transition to monosymmetry in the corolla orientations of organs within and between whorls cannot read-
is mirrored by a similar transition in the outer (expressed) sta- ily be depicted in a floral formula and, to compensate for
men whorl, suggesting a degree of developmental correlation this one deficit, features reflecting the axial extension of the
and encouraging the particular model of floral developmental flower – ovary position and the partial or complete fusion of
control in orchids advanced by Bateman & Rudall (2006), and organs – are more readily summarised as formulae than as
tested by Mondragón-Palomino & Theissen (2009). diagrams. The greatest strength of floral formulae is their
Astonishingly, even the best-known angiosperm model or- compactness and their exclusive use of typeface; unlike floral
ganisms, Arabidopsis and Antirrhinum, possess controversial diagrams, they are readily employed within bodies of text.
floral architecture (e.g., Endress, 1992). Even in Arabidopsis Floral formulae should become a recommended element
– the cornerstone of the ‘ABC’ model of floral developmental of formal taxonomic descriptions.  — As defined in the glos-
control and the most intensively researched angiosperm spe- sary to the ICBN (Vienna Code ; McNeill & al., 2006, footnote
cies on the planet – there remain three significantly different to recommendation 8A.4), the protologue is “everything associ-
interpretations of floral organisation, all of which show pe- ated with a name at its valid publication, i.e., description or di-
culiarities that are important for discussion of floral structure agnosis, illustrations, references, synonymy, geographical data,
(cf. Meyerowitz & al., 1991; Endress, 1992; Meyerowitz, 1994; citation of specimens, discussion and comments.” Bateman
Luo & al., 1996; see also Ronse De Craene, 2002). The flowers (2009, in press) recently highlighted the incongruity evident in
can be viewed as either (1) four-whorled with different organ the ICBN between the plethora of articles and recommendations
numbers, (2) six-whorled with two organs each and the organs that regulate nomenclature and the designation of types on the
in the median plane are in pairs (except for the sepals), or (3) one hand and the character content of the associated diagnosis/
five-whorled with four organs each; two outer stamens lost description on the other. There are no minimum requirements
and two carpels reduced: for the content of a botanical diagnosis (other than that it should
be written in Latin: McNeill & al., 2006, article 36.1, 36A), an
4-whorled:2 K4 ┼ C4* A6┼ G(2)┼ Vp∞ astonishing freedom that is in practice a recipe for anarchy in
6-whorled:2 K2 + 2┼ C4* A2 + 4 ┼ G(2)┼ Vp∞ both diagnosis and subsequent identifications using that diag-
5-whorled:2 K4 ┼ C4* A2 : 20 + 4 ┼ G(2 : 2sterile)┼ Vp∞ nosis. Bateman recommended that the content of diagnoses
should be considered more seriously by the ICBN, taking into
Selecting among these three alternative interpretations consideration recent initiatives in DNA barcoding (e.g., Tautz &
has downstream implications for the popular causal models al., 2003; Savolainen & al., 2005) that could lead to diagnoses
conceived as general explanations of flower development in wholly lacking in morphological information, written in the
angiosperms (Meyerowitz & al., 1991; Meyerowitz, 1994). language of nucleotides rather than Latin.

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TAXON 59 (1) • February 2010: 241–250 Prenner & al. • Floral formulae in taxonomic descriptions

We believe that a requirement (or, failing that, a recommen- Ellstrand, N.C. 1983. Floral formula inconstancy within and among
dation) for routine inclusion of floral formulae in angiosperm plants and populations of Ipomopsis aggregata (Polemoniaceae).
Bot. Gaz. 144: 119–123.
diagnoses would bring some much-needed consistency to di- Endress, P.K. 1992. Evolution and floral diversity: The phylogenetic
agnoses, as well as acting as a de facto checklist for features surroundings of Arabidopsis and Antirrhinum. Int. J. Pl. Sci. 153:
of the flower that merit more detailed description. S106–S122.
Future developments.  — We will continue to explore and, Endress, P.K. 1994. Diversity and evolutionary biology of tropical
where possible, further expand the utility of floral formulae. flowers. Cambridge: Cambridge Univ. Press.
Two areas where further progress could be made are to explore Endress, P.K. 1999. Symmetry in flowers: Diversity and evolution.
Int. J. Pl. Sci. 160 (6 Suppl.): S3–S23.
in greater detail the application of floral formulae to ontoge- Endress, P.K. 2001. Evolution of floral symmetry. Curr. Opin. Pl.
netic series, seeking to encapsulate heterochronic as well as Biol. 4: 86–91.
heterotopic shifts in morphology (e.g., Gould, 1977; Alberch Goebel, K. 1887. Outlines of classification and special morphology of
& al., 1979; Bateman, 1994), and to adapt the formulae for ap- plants. Oxford: Clarendon Press.
plication to the (mostly dioecious) gymnosperms, both living Gould, S.J. 1977. Ontogeny and phylogeny. Cambridge, Massachu-
setts: Belknap.
and fossil. It is also our intention to encourage other authors
Grisebach, A. 1854. Grundriss der systematischen Botanik für aka-
to critique our present suggestions in print, with the ultimate demische Vorlesungen. Göttingen: Verlag der Dietrichschen Bu-
goal of achieving a consensus notation for floral formulae. chhandlung.
Judd, W.S., Campbell, C.S., Kellogg, E.A., Stevens, P.F. & Dono-
ghue, M.J. 2007. Plant systematics: A phylogenetic approach, 3rd
ACKNOWLEDGEMENTS ed. Sunderland, Massachusetts: Sinauer.
Kocyan, A. & Endress, P.K. 2001. Floral structure and development of
Apostasia and Neuwiedia (Apostasioideae) and their relationships
We thank Louis Ronse De Craene, Erik Smets and Dmitry to other Orchidaceae. Int. J. Pl. Sci. 162: 847–867.
Sokoloff for helpful suggestions, though the opinions expressed here Leins, P. & Erbar, C. 2008. Blüte und Frucht: Morphologie, Ent-
remain our own. This contribution was written while RMB was sala- wicklungsgeschichte, Phylogenie, Funktion und Ökologie, 2. Aufl.
ried via NERC grant NE/E004369/1 (PI Jason Hilton, University of Stuttgart: E. Schweizerbart’sche Verlagsbuchhandlung.
Birmingham). Luo, D., Carpenter, R., Vincent, C., Copsey, L. & Coen, E. 1996.
Origin of floral asymmetry in Antirrhinum. Nature 383: 794–799.
Martius, C.F. 1828. Über die Architectonik der Blüthen. Isis (Oken)
21: 522–529.
LITERATURE CITED McNeill, J., Barrie, F. R., Burdet, H.M., Demoulin, V., Hawk-
sworth, D.L., Marhold, J., Nicolson, D.H., Prado, J., Silva, P.C.,
Alberch, P., Gould, S.J., Oster, G.F. & Wake, D.B. 1979. Size and Skog, J.E., Wiersema, J.H. & Turland, N.J. 2006. International
shape in ontogeny and phylogeny. Paleobiology 5: 296–317. code of botanical nomenclature (Vienna Code): Adopted by the
Bachelier, J.B. & Endress, P.K. 2009. Comparative floral morphol- Seventeenth International Botanical Congress; Vienna, Austria,
ogy and anatomy of Anacardiaceae and Burseraceae (Sapindales), July 2005. Regnum Vegetabile 146. Ruggell: Gantner.
with a special focus on gynoecium structure and evolution. Bot. J. Méndez, M. & Gómez, J.M. 2006. Phenotypic gender in Horma-
Linn. Soc. 159: 499–571. thophylla spinosa (Brassicaceae): A perfect hermaphrodite with
Barrett, S.C.H. 2002. The evolution of plant sexual diversity. Nature tetradynamous flowers, is variable. Pl. Syst. Evol. 262: 225–237.
Rev. Genet. 3: 274–284. Meyerowitz, E.M. 1994. The genetics of flower development. Sci.
Bateman, R.M. 1994. Evolutionary-developmental change in the Amer. 271: 56–65.
growth architecture of fossil rhizomorphic lycopsids: Scenarios Meyerowitz, E.M., Bowman, J.L., Brockman, L.L., Drews, G.N.,
constructed on cladistic foundations. Biol. Rev. 69: 527–597. Jack, T., Sieburth, L.E. & Weigel, D. 1991. A genetic and mo-
Bateman, R.M. 2009. What’s in a name? [Parts 1 and 2]. J. Hardy lecular model for flower development in Arabidopsis thaliana.
Orchid Soc. 6: 53–63, 88–99. Development Suppl. 1: 157–167.
Bateman, R.M. In press. The perils of addressing long-term challenges Mondragón-Palomino, M. & Theissen, G. 2009. Why are orchid
in a short-term world: Making descriptive taxonomy predictive. flowers so diverse? Reduction of evolutionary constraints by para-
In: Hodkinson, T., Jones, M.B., Waldren, S. & Parnell, J. (eds.), logues of class B floral homeotic genes. Ann. Bot. 104: 583–594.
Climate change and systematics. Systematics Association special Nutt, P., Zierrnann, J.G., Hintz, M., Neuffer, B. & Theissen, G.
volume 79. Cambridge: Cambridge Univ. Press. 2006. Capsella as a model system to study the evolutionary rel-
Bateman, R.M. & Rudall, P.J. 2006. The Good, the Bad, and the evance of floral homeotic mutants. Pl. Syst. Evol. 259: 217–235.
Ugly: Using naturally occurring terata to distinguish the possible Prance, G.T. & Mori, S.A. 1977. What is Lecythis? Taxon 26: 209–222.
from the impossible in orchid floral evolution. Aliso 22: 481–496. Prance, G.T. & Mori, S.A. 1979. Lecythidaceae, part 1, The actino-
Burrows, G. 2009. The virtual floral formula. Pl. Sci. Bull. 55: 149–150. morphic-flowered New World Lecythidaceae. Flora Neotropica
Cassel, F.P. 1820. Morphonomia botanica: sive Observationes circa Monograph 21. New York: New York Botanical Garden.
proportionem et evolutionem partium plantarum. Colonia Agrip- Prenner, G. 2004a. New aspects in floral development of Papilionoi-
pina [Cologne]: M. DuMont-Schauberg. deae: Initiated but suppressed bracteoles and variable initiation of
Chen, S.-C. 1965. A primitive new orchid genus Tangtsinia and its sepals. Ann. Bot. 93: 537–545.
meaning in phylogeny. Acta Phytotax. Sin. 16: 82–85. Prenner, G. 2004b. The asymmetric androecium in Papilionoideae
Doyle, J.J. 1992. Gene trees and species trees: Molecular systematics (Leguminosae): Definition, occurrence, and possible systematic
as one-character taxonomy. Syst. Bot. 17: 144–163. value. Int. J. Pl. Sci. 165: 499–510.
Eichler, A.W. 1875. Blüthendiagramme construiert und erläutert, 1. Prenner, G., Box, M.S., Cunniff, J. & Rudall, P.J. 2008. The branch-
Theil. Leipzig: Engelmann. ing stamens of Ricinus and the homologies of the angiosperm sta-
Eichler, A.W. 1878. Blüthendiagramme construiert und erläutert, 2. men fascicle. Int. J. Pl. Sci. 169: 735–744.
Theil. Leipzig: Engelmann. Prenner, G. & Klitgaard, B.B. 2008. Towards unlocking the deep

249
Prenner & al. • Floral formulae in taxonomic descriptions TAXON 59 (1) • February 2010: 241–250

nodes of Leguminosae: Floral development and morphology of the Stützel, T. 2002. Botanische Bestimmungsübungen. Stuttgart: Ulmer.
enigmatic Duparquetia orchidacea (Leguminosae, Caesalpinioi- Tautz, D., Arctander, P., Minelli, A., Thomas, R.H. & Vogler, A.P.
deae). Amer. J. Bot. 95: 1349–1365. 2003. A plea for DNA taxonomy. Trends Ecol. Evol. 18: 70–74
Prenner, G., Vergara-Silva, F. & Rudall, P.J. 2009. The key role of Theissen, G., Becker, A., Winter, K.-U., Münster, T., Kirchner, C.
morphology in modelling inflorescence architecture. Trends Pl. & Saedler, H. 2002. How the land plants learned their floral ABCs:
Sci. 14: 302–309. The role of MADS-box genes in the evolutionary origin of flow-
Ronse De Craene, L.P. 2002. Floral development and anatomy of ers. Pp. 173–205 in: Cronk, Q.C.B., Bateman, R.M. & Hawkins,
Pentadiplandra (Pentadiplandraceae): A key genus in the identi- J.A. (eds.), Developmental genetics and plant evolution. London:
fication of floral morphological trends in core Brassicales. Canad. Taylor & Francis.
J. Bot. 80: 443–459. Tian, X.-H., Zhao, L., Ren, Y. & Zhang, X.-H. 2007. Number of
Rudall, P.J. & Bateman, R.M. 2002. Roles of synorganisation, zygo- floral organs in Circaeaster agrestis (Circaeasteraceae) and pos-
morphy and heterotopy in floral evolution: The gynostemium and la- sible homeosis among floral organs. Pl. Syst. Evol. 265: 259–265.
bellum of orchids and other lilioid monocots. Biol. Rev. 77: 403–441. Tsou, C.-H. & Mori, S.A. 2007. Floral organogenesis and floral evo-
Rudall, P.J. & Bateman, R.M. 2004. Evolution of zygomorphy in lution of the Lecythidoideae (Lecythidaceae). Amer. J. Bot. 94:
monocot flowers: Iterative patterns and developmental constraints. 716–736.
New Phytol. 162: 25–44. Tucker, S.C. 1998. Floral ontogeny in legume genera Petalostylis,
Sachs, J. 1873. Lehrbuch der Botanik nach dem gegenwaertigen Stand Labichea, and Dialium (Caesalpinioideae: Cassieae): A series in
der Wissenschaft, 3. Aufl. Leipzig: Engelmann. floral reduction. Amer. J. Bot. 85: 184–208.
Sattler, R. 1973. Organogenesis of flowers: A photographic text-atlas. Tucker, S.C. 2000. Floral development in tribe Detarieae (Legumi-
Toronto: Univ. of Toronto Press. nosae: Caesalpinioideae): Amherstia, Brownea, and Tamarindus.
Savolainen, V., Cowan, R.S., Vogler, A.P., Roderick, G.K. & Lane, Amer. J. Bot. 87: 1385–1407.
R. 2005. Towards writing the encyclopaedia of life: An introduction Tucker, S.C. 2002. Floral ontogeny of Cercis (Leguminosae: Cae-
to DNA barcoding. Philos. Trans., Ser. B 360: 1805–1811. salpinioideae: Cercideae): Does it show convergence with papil-
Soltis, D.E., Fishbein, M. & Kuzoff, R.K. 2003. Reevaluating the ionoids? Int. J. Pl. Sci. 163: 75–87.
evolution of epigyny: Data from phylogenetics and floral ontogeny. Weberling, F. 1989. Morphology of flowers and inflorescences. Trans.
Int. J. Pl. Sci. 164 (5 Suppl): S251–S264. R.J. Pankhurst from the original of 1981. Cambridge: Cambridge
Soltis, D.E. & Hufford, L. 2002. Ovary position diversity in Saxifra- Univ. Press.
gaceae: Clarifying the homology of epigyny. Int. J. Pl. Sci. 163:
277–293.

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