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Understanding the Structure of Flowers—The Wonderful Tool of Floral Formulae: a Response to Prenner & Al

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Understanding the Structure of Flowers—The Wonderful Tool of Floral Formulae: a Response to Prenner & Al TAXON 63 (5) • October 2014: 1103–1111 Ronse De Craene & al. • Floral formulae METHODS 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, [email protected] 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 to describe flowers. Prenner & al. (2010: 241) also suggested that floral formulae should be an inherent component of plant Flowers are remarkably conservative in their numbers diagnoses and taxonomic descriptions, “functioning as a logi- of parts. Whichever flower of Erodium (Geraniaceae) a keen cal phenotypic counterpart to the DNA barcode”. It is clear that observer will look at (see Fig. 1A), he will always find the same floral formulae have a clear advantage over lengthy descrip- number of whorls and the same number of organs within each tions, as the structure of a flower can be presented in a suc- whorl of the flower (five sepals and petals, two whorls of five cinct way, accessible to any reader. As such floral formulae stamens, one of which is sterile, and five carpels: Fig. 1A). are a powerful incentive for consistent observations and also Flower morphologists have sought for ways to succinctly a valuable teaching tool (Prenner & al., 2010). Prenner & al. describe flowers, and a perfect tool was conceived in the form of advocate the use of symbols that can be reproduced easily with floral formulae. The first floral formulae were created in the first a non-sophisticated computer, enabling its universal use. decades of the nineteenth century and have been progressively Floral diagrams, two-dimensional representations of the developed especially in Germany. For an overview of the history ground-plan of the flower, are the major counterpart to flo- of the use of floral formulae we refer to Prenner & al. (2010). ral formulae in floral descriptions. As discussed in Ronse De Although floral formulae (together with floral diagrams) Craene (2010), floral formulae have a disadvantage against have been integral part of the basic teaching in botany, the floral diagrams in that it is difficult to describe the position recent depreciation of botany at universities has not helped in of organs in the flower, and impossible to show any spatial the recognition of the value of this methodology. Few recent arrangements or any conspicuous special structures, such as papers or textbooks use floral formulae, often in a very simpli- nectaries, coronas or appendages. However, floral formulae fied and frequently inconsistent form, such as Judd & al. (2002), have the advantage that they do not necessitate any pictorial Stützel (2006), Tsou & Mori (2007), Leins & Erbar (2010), or means. They also open possibilities for comparative research Simpson (2010). and the understanding of trends in flower evolution. The value of floral formulae has been highlighted by We agree with Prenner & al. (2010) that floral formulae Prenner & al. (2010) who proposed an updated list of symbols (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 Version of Record (identical to print version). 1103 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. 1104 Version of Record (identical to print version). TAXON 63 (5) • October 2014: 1103–1111 Ronse De Craene & al. • Floral formulae research. Therefore, we believe that a broader use of these tools structures of unknown/unclear origin), K for calyx, Corolla, should be encouraged. Ronse De Craene (2010: 39) came up Perigon, Androecium, Gynoecium), and a use of lower case with a list of abbreviations comparable to Prenner & al. (2010), letters for specific structures that are part of organs, such as although it was less elaborate. However, the advantages of flo- locules, ovules (including their position on the placenta), pollen ral formulae for educational purposes are clear and lecturers sacs, styles and stigmas. may adapt a personal approach (E. Smets, pers. comm.). As The inclusion of details of the flower, such as the number suggested by Prenner & al. (2010), their proposed system is of pollen sacs or locules forces the user to accurately observe open for discussion, and Taxon is probably a good medium to the flower to record as many details as possible. Not including start this discussion. The ultimate aim is to reach consensus these details in a consistent manner may lead to inconsisten- among botanists for the use of a balanced and universal system. 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 A REEVALUATION OF FLORAL FORMULAE characters. Several qualitative features can be shown through superscript symbols and Prenner & al. use this to great effect. The following points are a criticism of the paper by Prenner For example, C stands for the number of petals, while c stands & al. (2010) and suggestions for improvement. We also refer to for petaloidy, which can be used as a special feature of the Table 1 for clarity. perigon. A calyx can often be pigmented, as, e.g., in Fuchsia The floral formulae proposed by Prenner & al. (2010) are (Onagraceae) and this can be shown as Kc. Pc stands for a very detailed and contain more information than classical for- petaloid perigon, when it is not possible to differentiate sepals mulae, including extrafloral organs associated with flowers, from petals, as in tulips (Weberling, 1989). such as bracts and bracteoles, and gynoecial characters, such Inclusion of bracts and bracteoles in floral formulas is use- as the placentation and ovules. However, it is best to maintain ful, although strictly speaking they belong to the inflorescence. a hierarchical arrangement with subcategories. Treating parts Bracteoles are generally paired in eudicots and single in mono- of organs (i.e., ovules) at the same level as the organs leads to cots (Ronse De Craene, 2010). However, in some cases with a confusion and makes the floral formulae appear too complex. multitude of bracts surrounding the flower a strict distinction A better approach includes a restriction of capital letters to between bracts and bracteoles is not possible (e.g., Reaumuria the main organs associated with the flower (see Table 1) Bract, in Tamaricaceae: Ronse De Craene, 1990). Boundaries of flow- Bracteole, Epicalyx
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