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, 107 (1), 1-37

Molecular Phylogenetics of Subfamily Ornithogaloideae (Hyacinthaceae) Based on Nuclear and Plastid DNA Regions, Including a New Taxonomic Arrangement

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Molecular Phylogenetics of Subfamily Ornithogaloideae (Hyacinthaceae) Based on Nuclear and Plastid DNA Regions, Including a New Taxonomic Arrangement

Mario Martínez-Azorín et al. Ann Bot.

Abstract

Background and aims: The taxonomic arrangement within subfamily Ornithogaloideae (Hyacinthaceae) has been a matter of controversy in recent decades: several new taxonomic treatments have been proposed, based exclusively on plastid DNA sequences, and these have resulted in classifications which are to a great extent contradictory. Some authors have recognized only a single genus Ornithogalum for the whole subfamily, including 250-300 species of variable morphology, whereas others have recognized many genera. In the latter case, the genera are inevitably much smaller and they are better defined morphologically. However, some are not monophyletic as circumscribed.

Methods: Phylogenetic analyses of Ornithogaloideae were based on nucleotide sequences of four plastid regions (trnL intron, trnL-F spacer, rbcL and matK) and a nuclear region (ITS). Eighty species covering all relevant taxonomic groups previously recognized in the subfamily were sampled. Parsimony and Bayesian analyses were performed. The molecular data were compared with a matrix of 34 morphological characters.

Key results: Combinations of plastid and nuclear data yielded phylogenetic trees which are better resolved than those obtained with any plastid region alone or plastid regions in combination. Three main clades are found, corresponding to the previously recognized tribes Albuceae, Dipcadieae and Ornithogaleae. In these, up to 19 clades are described which are definable by morphology and biogeography. These mostly correspond to previously described taxa, though some need recircumscription. Morphological characters are assessed for their diagnostic value for taxonomy in the subfamily.

Conclusions: On the basis of the phylogenetic analyses, 19 monophyletic genera are accepted within Ornithogaloideae: Albuca, Avonsera, Battandiera, Cathissa, Coilonox, Dipcadi, Eliokarmos, Elsiea, Ethesia, Galtonia, Honorius, Loncomelos, Melomphis, Neopatersonia, Nicipe, Ornithogalum, Pseudogaltonia, Stellarioides and Trimelopter. Each of these has a particular syndrome of morphological characters. As a result, 105 new combinations are made and two new names are proposed to accommodate the taxa studied in the new arrangement. A short morphological diagnosis, synonymy, details of distribution and an identification key are presented.

Figures

Fig. 1.
Fig. 1.
Bayesian inference analysis obtained from ITS sequences in the phylogenetic analysis of Ornithogaloideae. Phylogenetic relationships shown by one of the post-burn-in topologies. Taxa are identified following the traditional treatment of Ornithogalum and other related genera (cf. Obermeyer, 1978; Zahariadi, 1980; Müller-Doblies and Müller-Doblies, 1996). The new proposed generic circumscription of the present study is shown on the right side. Branch lengths are shown above branches, and posterior probabilities and bootstrap percentages are shown below. Arrows indicate branches that collapse in the 50 % majority rule consensus tree obtained by Bayesian analysis.
Fig. 2.
Fig. 2.
Bayesian inference analysis of the combination of rbcL, trnL-F, matK and ITS sequences in the phylogenetic analysis of Ornithogaloideae. Phylogenetic relationships shown by one of the post-burn-in topologies. Taxa are identified following the traditional treatment of Ornithogalum and other related genera (cf. Obermeyer, 1978; Zahariadi, 1980; Müller-Doblies and Müller-Doblies, 1996). The new proposed generic circumscription of the present study is shown on the right side. Branch lengths are shown above branches, and posterior probabilities and bootstrap percentages are shown below. Arrows indicate branches that collapse in the 50 % majority rule consensus tree obtained by Bayesian analysis.
Fig. 3.
Fig. 3.
Recontruction of character distribution in a post-burn-in tree obtained by a Bayesian inference, after combination of all the DNA regions studied: (A) main colour of tepals; (B) darker central band on tepals; (C) fusion of tepals and adnation of filaments to tepals; (D) seed morphology. All characters are unordered.
Fig. 3.
Fig. 3.
Recontruction of character distribution in a post-burn-in tree obtained by a Bayesian inference, after combination of all the DNA regions studied: (A) main colour of tepals; (B) darker central band on tepals; (C) fusion of tepals and adnation of filaments to tepals; (D) seed morphology. All characters are unordered.
Fig. 4.
Fig. 4.
Morphological variation in several genera of Ornithogaloideae: (A) Honorius boucheanum (from Sturm, Deutschlands Flora in Abbildungen: tab. 33. 1796, as Ornithogalum boucheanum); (B) Nicipe juncifolia (from Jacquin, Plantarum Rariorum Horti Caesarei Schoenbrunnensis Descriptiones et Icones, 1: tab. 90. 1797, as O. juncifolium); (C) Elsiea corymbosa (from Leighton in J. S. African Bot. 10: 56. 1944); (D) Ethesia prasina (from Lindl., Bot. Reg. 2: tab. 158. 1816, as O. prasinum); (E) E. xanthochlora (from Pretoria National Herbarium PREART 0002737, as O. xanthochlorum); (F) Trimelopter unifolium (from Jacquin, Icones Plantarum Rariorum, 2(16): tab. 429. 1795, as O. fuscatum); (G and H) T. dyeri (pictures by A. P. Dold); (I) Battandiera amoena (picture by Stephen L. Jury); (J) capsule of B. amoena; (K) capsules and seeds of B. pulchra; (L) Stellarioides longebracteata; (M) S. sessiliflora; (N) Capsules and seeds of S. longebracteata; (O) capsules and seeds of S. sessiliflora.
Fig. 5.
Fig. 5.
Inflorescence and flower features of most of the Eurasian genera of Ornithogaloideae: (A) Cathissa reverchonii; (B) C. concinna; (C) C. unifolia; (D) Melomphis arabica; (E) Loncomelos narbonense; (F) L. pyrenaicus; (G) Ornithogalum baeticum; (H) O. bourgaeanum; (I) O. divergens.
Fig. 6.
Fig. 6.
Capsule (lateral and section views) and seed features of most of the Eurasian genera of Ornithogaloideae: (A) Cathissa reverchonii; (B) C. concinna; (C) C. unifolia; (D) Melomphis arabica; (E) Loncomelos narbonense; (F) L. pyrenaicus; (G) Ornithogalum baeticum; (H) O. bourgaeanum; (I) O. divergens.

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