Abstract
Delimitation of sections is controversial within the genus Cytisus L. (Fabaceae, Genisteae). A morphological study has been conducted on 19 taxa from sections Alburnoides, Spartopsis and Verzinum to clarify their discrimination. Thirty-five quantitative and qualitative characters were recorded on a maximum of 15 dry or living flowers per taxon. Three multiple correspondence factor analyses (MCFA) were performed on a matrix based on 22 of the 35 recorded morphological characters to (1) compare the variability within and between individuals and (2) distinguish groups among the studied taxa. MCFA showed that both flowers sampled from the same plant or different individuals could represent the morphological variability of a taxon. MCFA also clustered the 19 taxa into three groups corresponding to sections Alburnoides, Spartopsis and Verzinum as defined by Cristofolini and Troia (Taxon 44:733–746, 2006). However, floral morphology has not been sufficient to discriminate taxa within sections. A key of the three studied sections based on floral characters is given.
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Acknowledgments
The authors would like to thank Eurogeni and INRA Angers-Nantes, centre d’Angers, who funded this work, as well as Véronique Kapusta, who managed the ‘Genisteae’ team. The authors are also thankful to the curators of herbaria B, BR, E, G, K, L, MA, MPU, P, RNG, SALA and VAL for providing herbarium specimens that were used in the present study. Many thanks to Joël Mathez and Frédéric Andrieu from the Conservatoire Botanique National du Bassin Méditerranéen for their help in understanding the reproductive biology of Cytisus baeticus.
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Appendices
Appendix 1
List of voucher specimens observed for each taxon (x2, x3, x4 after the name of a specimen indicate that two, three or four flowers were observed on that specimen, abbreviation of herbarium following).
Cytisus arboreus. Algeria: Battandier and Trabut 195 (L), Birch Wolfe s.n. (K), Munby s.n. (K), Trethewy 50 (K); France: Penchinat 242 (L), Penchinat 1431 (L).
Cytisus ardoinoi. France: Hepper 9377 (K), Moggridge s.n. (K) x3.
Cytisus baeticus. Algeria: Alleizette s.n. (BR), Birch Wolfe s.n. (K); Morocco: Bramwell et al. 516 (RNG), Davis 51235 (RNG), Lewalle 8620 (BR), Sutton et al. 56 (RNG), Trethewy 5 (K); Spain: Bisby et al. 45 (K), Bisby, Davis 61512 (E), Fernández Casas 2942 (BR), Fernández González et al. 1141 (VAL), González and López s.n. (VAL), Gros s.n. (VAL), Mateo s.n. (VAL), Mauricio 8377 (VAL).
Cytisus balansae. Algeria: Davis 52364 (RNG) x2, Davis 52501 (RNG) x2; Morocco: Aedo et al. 4190 (RNG), Aedo et al. 4190 (VAL) x2; North Africa: Powell s.n. (MPU) x3.
Cytisus cantabricus. Spain: Aedo s.n. (MA), Laínz s.n. (RNG), Aizpuru and Catalán 14162 (VAL), Aizpuru and Catalán 14162 (SALA), Bellés s.n. (VAL), Boccilla Martínez s.n. (VAL), Brummitt and Chater 97 (K), Fabregat s.n. (VAL), Fernández Casas s.n. (MA), Harrold and McBeath 156 (E), Herrera Gallastegui 15199 (VAL), Mateu et al. s.n. (VAL), Vermeulen-Fernández 125 (L), Valle s.n. (SALA), Valle et al. s.n. (SALA).
Cytisus catalaunicus. France: Andr. et al. 1645 (L) x2, Heukels 72 (L) x2; Morocco: Jury 13694 (RNG) x3; Spain: Font s.n. (VAL) x2, Kruseman s.n. (L) x2, Llistosella s.n. (VAL), Sennen 539 (RNG), Sennen s.n. (L) x2.
Cytisus galianoi. Spain: Aedo et al. 388 (MA) x2, Archibald 3489 (E) x2, Jury 17128 (RNG) x2, Stocken 219.63 (E) x2, Valdès et al. 621 (RNG), Valdès et al. 896 (RNG) x3, Valdès et al. 951 (RNG) x2, Valdès et al. 951.88 (G).
Cytisus grandiflorus. Morocco: Jury et al. 19589 (RNG), López andand Muñoz Garmendia 9127 (RNG); Portugal: Castro 4310 (MPU), Castro 157/36 (VAL), Moller 2151 (BR), Rothmaler 17070 (VAL), Rothmaler s.n. (VAL); Spain: Aguilella and Garcia-Fayon s.n. (VAL), Arán and Toha 5446 (VAL), Bisby et al. 58 (RNG), Bisby et al. 63 (RNG), Castroviejo et al. 9345 (BR), Güemes s.n. (VAL), Lange 469 (BR), Willkomm 10 (BR).
Cytisus insularis. Spain: Ortiz 49419 (MA).
Cytisus malacitanus. Spain: Arán andand Toha 5075 (VAL) x2, Bisby 53 (RNG), D. B. L. 33 (K) x2, Davis 61704 (E) x2, Fernández González 1142 (VAL) x2, Lange 466 (K) x2, Mateo and Lazaro s.n. (VAL), Sennen 1380 (RNG), Sennen 2585 (RNG) x2.
Cytisus maurus. Morocco: Fernández Casas et al. 5020 (RNG) x3, Jury et al. 16836 (RNG), Mathez 7124 (MPU), Optima Iter V 1289 (RNG) x2.
Cytisus megalanthus. Morocco: Maire s.n. (MPU) x2.
Cytisus multiflorus. Spain: Barra et al. 2312 (RNG) x2, Brummitt and Chater 172 (K) x2, Unknown collector s.n. (K) x2, Gavilán 16148 (RNG) x2; Unknown country: Guinea s.n. (RNG) x2.
Cytisus oromediterraneus. Portugal: Optima Iter VI 1075 (RNG); Spain: Aguilella s.n. (VAL) x3, Alonso andand Paz s.n. (VAL), Capell s.n. (VAL), Garín 17069 (RNG), Jury and Nieto Feliner 11177 (RNG), Mateo 566 (VAL), Mateo s.n. (VAL) x2, Miles 136 (RNG), Puente et al. s.n. (VAL), Rivas-Martínez 16149 (VAL), Segura Zubizarreta 40.104 (VAL).
Living specimen from INRA (FRANCE): SE002-013, sampled on 23 June 2010.
Cytisus reverchonii. Spain: Reverchon 1095 (P) x5, Reverchon 1095 (L).
Cytisus scoparius. France: Hekker s.n. (L); Morocco: Wilde et al. 2915 (L) x2; The Netherlands: Ooststroom 8344 (L) x4; Portugal: Bourgeau 1812 (P) x2, Bourgeau 1813 (P); Spain: Bisby et al. 87 (RNG) x2, Guinea 1179 (RNG), López 1647 (RNG) x2.
Cytisus striatus. Morocco: Lambinon and Van den Sande 94/Ma/389 (RNG), Mejías and Silvestre s.n. (RNG); Spain: Adey 178 (RNG), Adey 45b (RNG), Bisby et al. 1314 (RNG), Bourgeau 2414 (K), Font i Quer and Gros 256/21 (B), Font i Quer andand Gros 256/21 (RNG), Gros 200/30 (RNG), Optima Iter VI 201 (RNG), Porta et Rigo 112 (B), Porta et Rigo 112 (E), Rouy 71 (E) x2, Scholz and Hiepko 821 (B).
Cytisus transiens. Morocco: Mathez 1229 (MPU) x2, Mathez 1315 (MPU) x2.
Cytisus valdesii. Morocco: Ait Lafkih et al. 513 (RNG) x2, Fernández Casas et al. 4694 (RNG) x2, Herrero et al. 2968 (RNG) x2, Jury et al. 18121 (RNG) x2, López and Muñoz Garmendia 8957 (RNG) x2.
Appendix 2
Characters and character states for the 35 recorded characters. Number between square brakets: position of the character in the matrix of characters; numbers between parenthesis: code for this character state in the matrix of characters; * indicates that the character has been used for statistical analyses.
Calyx
-
Length of the upper lip* [1] (mm): ≥1 and <2 (1), ≥2 and <3 (2), ≥3 and <4 (3), ≥4 and <5 (4), ≥5 and <6 (5), ≥6 and <7 (6), ≥7 and <8 (7), ≥8 and <9 (8)
-
Length of the lower lip* [2] (mm): ≥ and <2 (1), ≥2 and <3 (2), ≥3 and <4 (3), ≥4 and <5 (4), ≥5 and <6 (5), ≥6 and <7 (6), ≥7 and <8 (7), ≥8 and <9 (8)
-
Depth of the teeth in the upper lip [3]: <1 mm (1), = 1 mm (2), >1 mm (3)
-
Depth of the teeth in the lower lip [4]: <1 mm (1), = 1 mm (2), >1 mm (3)
-
Angle between the two lips [5]: <45° (1), = 45° (2), >45° and <90° (3), = 90° (4), >90° and <135° (5), = 135° (6), >135° (7)
-
Hairiness* [6]: glabrous (1), ciliate (2), pubescent (3).
Standard
-
Length* [7] (mm): ≥6 and <8 (1), ≥8 and <10 (2), ≥10 and <12 (3), ≥12 and <14 (4), ≥14 and <16 (5), ≥16 and <18 (6), ≥18 and <20 (7), ≥20 and <22 (8), ≥22 and <24 (9), ≥24 and <26 (10)
-
Width* [8] (mm): ≥6 and <8 (1), ≥8 and <10 (2), ≥10 and <12 (3), ≥12 and <14 (4), ≥14 and <16 (5), ≥16 and <18 (6), ≥18 and <20 (7), ≥20 and <22 (8), ≥22 and <24 (9), ≥24 and <26 (10)
-
Position of the maximal width [9]: upper part (1), middle part (2), lower part (3) of the petal
-
Shape* [10]: oblong (1), eliptical (2), orbicular (3), suborbicular (4), obovate (5), rhombic (6), ovate (7)
-
Shape of the upper part* [11]: emarginated (1), slightly emarginated (2), neither emarginated nor incurvated (3), slightly incurvated (4), incurvated (5)
-
Hairiness [12]: glabrous, constant character.
Wing petals
-
Length* [13] (mm): ≥6 and <8 (1), ≥8 and <10 (2), ≥10 and <12 (3), ≥12 and <14 (4), ≥14 and <16 (5), ≥16 and <18 (6), ≥18 and <20 (7), ≥20 and <22 (8), ≥22 and <24 (9), ≥24 and <26 (10)
-
Width* [14] (mm): ≥1 and <3 (1), ≥3 and <5 (2), ≥5 and <7 (3), ≥7 and <9 (4), ≥9 and <11 (5), ≥11 and <13 (6)
-
Position of the maximal width [15]: upper part (1), middle part (2), lower part (3) of the petal, petal oblong (4)
-
Width of the apex* [16]: narrower (1), slightly narrower (2), as wide as (3), wider (4) than the rest of the petal
-
Shape of the apex* [17]: emarginated (1), rounded and curved (2), rounded and flat (3), very rounded (4), rounded (5), pointed (6)
-
Shape* [18]: obovate (1), rhombic (2), eliptical (3), sub-elliptical (4), upper part oblong (5), oblong (6), lower part oblong and lower part falcate (7), falcate (8), bean-shaped (9)
-
Curve* [19]: curvate (1), slightly curvate (2), erect (3)
-
Hairiness [20]: glabrous, constant character.
Keel petals
-
Length* [21] (mm): ≥6 and <8 (1), ≥8 and <10 (2), ≥10 and <12 (3), ≥12 and <14 (4), ≥14 and <16 (5), ≥16 and <18 (6), ≥18 and <20 (7), ≥20 and <22 (8), ≥22 and <24 (9), ≥24 and <26 (10)
-
Width* [22] (mm): ≥1 and <3 (1), ≥3 and <5 (2), ≥5 and <7 (3), ≥7 and <9 (4), ≥9 and <11 (5), ≥11 and <13 (6)
-
Position of the maximal width [23]: upper part (1), middle part (2), lower part (3) of the petal, petal oblong (4)
-
Shape* [24]: obovate (1), sub-eliptical (2), oblong (3), lower part oblong and upper part falcate (4), falcate (5), bean-shaped (6)
-
Width of the apex [25]: narrower (1), slightly narrower (2), as wide as (3), wider (4) than the rest of the petal
-
Shape of the apex [26]: truncate (1), rounded and large (2), rounded and narrow (3), pointed (4)
-
Curve [27]: curvate (1), slightly curvate (2), erect (3)
-
Axis of the apex if falcate [28]: forward (1), diagonal (2), upward (3)
-
Hairiness* [29]: absence (1), hairs on the basis of the lower margin (2), hairs on the lower margin (3).
Relative sizes of the petals
-
Relative size of wings and keel* [30]: wings > keel (1), wings = keel (2), wings < keel (3)
-
Relative size of wings and standard* [31]: standard > wings (1), standard = wings (2), standard < wings (3)
-
Relative size of standard and keel* [32]: standard > keel (1), standard = keel (2), standard < keel (3).
Style
-
Shape of the lower part [33]: erect, constant character
-
Shape of the apex* [34]: slightly curved (1), curved (2), convolute within the keel (3), forming a loop (4)
-
Shape of the curve* [35]: incurvated (1), angular (2).
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Auvray, G., Gouron, C. & Malécot, V. Floral morphology to discriminate taxa between and within Cytisus sect. Alburnoides, sect. Spartopsis and sect. Verzinum (Genisteae, Fabaceae). Plant Syst Evol 298, 1827–1835 (2012). https://doi.org/10.1007/s00606-012-0683-8
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DOI: https://doi.org/10.1007/s00606-012-0683-8