Summary
The genus Phlomis is one of the largest genera of subfamily Lamioideae (Lamiaceae) with more than 100 recognised species distributed in Asia, southern Europe and northern Africa. Morphological studies have suggested that the species can be split into two morphologically distinct groups that have previously been recognised as either sections or separate genera. The current study represents the first molecular phylogenetic study of the genus in which taxa representative of the entire genus are included. In contrast to considerable morphological and ecological variation among taxa, sequence variation in the trnL intron, the trnL – F intergenic spacer, and the rps16 intron was low, resulting in sparse phylogenetic resolution, particularly among the South West Asian and Mediterranean taxa. However, the results support a split of the genus into two separate groups that are recognised here as the genera Phlomoides and Phlomis in order to decrease taxonomic complexity. The close relationship between Phlomis s.l. and Eremostachys, pointed out by several authors, is confirmed, and in a noteworthy new finding, the small Asian genus Notochaete, is more closely related to the Phlomis s.l. lineage than previously assumed. The two monotypic genera Pseuderemostachys and Lamiophlomis as well as Notochaete hamosa are transferred to Phlomoides and three new combinations are made. Character optimisation analyses based on geographic distributions of all taxa show that the groups obtained by molecular phylogenetic analysis correlate well with biogeography. The results suggest that the Phlomis s.l. lineage has a Central Asian origin in an area around Western China.
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References
Adylov, T. A., Kamelin, R. V. & Makhmedov, A. M. (1986). Notae de Lamiaceis (Lamiaceae). Novosti Sist. Vyssh. Rast. 23: 110 – 114 (in Russian).
_____, _____ & _____ (1987). Phlomoides. In: A. I. Vvedensky (ed.), Conspectus Florae Asiae Mediae, pp. 91 – 107. Academiae Scientarum USSR, Tashkent.
Albaladejo, R. G., Aparicio, A. & Silvestre, S. (2004). Variation patterns in the Phlomis × composita (Lamiaceae) hybrid complex in the Iberian Peninsula. Bot. J. Linn. Soc. 145: 97 – 108.
_____, Aguilar, J. F., Aparicio, A. & Feliner, G. N. (2005). Contrasting nuclear-plastidial phylogenetic patterns in the recently diverged Iberian Phlomis crinita and P. lychnitis lineages (Lamiaceae). Taxon 54: 987 – 998.
Aparicio, A. (1997). Fitness components of the hybrid Phlomis × margaritae Aparicio & Silvestre (Lamiaceae). Bot. J. Linn. Soc. 124: 331 – 343.
_____ & Albaladejo, R. G. (2003). Microsporogenesis and meiotic abnormalities in the hybrid complex of Phlomis composita (Lamiaceae). Bot. J. Linn. Soc. 143: 79 – 85.
_____, _____, Porras, M. & Ceballos, G. (2000). Isozyme evidence for natural hybridzation in Phlomis (Lamiaceae): Hybrid origin of the rare P. × margaritae. Ann. Bot. 85: 7 – 12.
Azizian, D. & Cutler, D. F. (1982). Anatomical, cytological and phytochemical studies on Phlomis L. and Eremostachys Bunge (Labiatae). Bot. J. Linn. Soc. 85: 249 – 281.
_____ & Moore, D. M. (1982). Morphological and palynological studies in Phlomis L, Eremostachys Bunge and Paraphlomis Prain (Labiatae). Bot. J. Linn. Soc. 85: 225 – 248.
Bentham, G. (1830). Synopsis of the genera and species of Indoan Labiatae. In: N. Wallich, Plantae asiaticae rariores 1. Treuttel & Würtz, London, Paris, Strasbourg.
_____ (1832 – 1836). Labiatarum Genera et Species. James Ridgeway and Sons, London.
Boissier, E. (1879). Flora Orientalis vol. 4 part 1. H. Georg, Genevae, Basileae, Lugduni.
Briquet, J. (1897). Phlomis L. In: A. P. Engler & K. Prantl (ed.), Die natürlichen Pflanzenfamilien, pp. 248 – 249. W. Engelmann, Leipzig.
Cantino, P. D. (1992). Evidence for a polyphyletic origin of the Labiatae. Ann. Missouri Bot. Gard. 79: 361 – 379.
Corriveau, J. L. & Coleman, A. W. (1988). Rapid screening method to detect potential biparental inheritance of plastid DNA and results for over 200 angiosperm species. Amer. J. Bot. 75: 1443 – 1458.
Fang, L.-Q., Pan, Y.-Z. & Gong, X. (2007). A karyomorphological study in the monotypic genus Lamiophlomis and five species in Phlomis (Lamiaceae). Acta Phytotax. Sin. 45: 627 – 632.
Farris, J. S., Albert, V. A., Kallersjo, M., Lipscomb, D. & Kluge, A. G. (1996). Parsimony jackknifing outperforms neighbor-joining. Cladistics 12: 99 – 124.
Goloboff, P. A., Farris, J. S. & Nixon, K. (2003a). TNT: Tree analysis Using New Technology. Version 1.0. Program and documentation available at http://www.zmuc.dk/public/phylogeny/tnt.
_____, _____, Källersjö, M., Oxelman, B., Ramírez, M. J. & Szumik, C. A. (2003b). Improvements to resampling measures of group support. Cladistics 19: 324 – 332.
_____, Carpenter, J. M., Arias, J. S. & Esquivel, D. R. M. (2008). Weighting against homoplasy improves phylogenetic analysis of morphological data sets. Cladistics 24: 1 – 16.
Hall, T. A. (1999). BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucl. Acids Symp. Ser. 41: 95 – 98.
Harley, R. M., Atkins, S., Budantsev, A. L., Cantino, P. D., Conn, B. J., Grayer, R., Harley, M. M., De Kok, R., Krestovskaya, T., Morales, R., Paton, A. J., Ryding, O. & Upson, T. (2004). Labiatae. In: J. W. Kadereit (ed.), The Families and Genera of Vascular Plants vol. VII Flowering Plants — Dicotyledons. Lamiales (except Acanthaceae including Avicenniaceae), pp. 167 – 275. Springer, Berlin, Heidelberg.
Hedge, I. C. (1990). Labiateae. In: S. I. Ali & Y. J. Nassir (eds), Flora of Pakistan, pp. 193 – 217. BCC and T Press, University of Karachi.
Herder, F. (1868). Enumeratio plantarum in regionibus Cis- et Transiliensibus a CL. semenovio anno 1857 collectarum. Labiatae. Bull. Soc. Imp. Naturalistes Moscou 41: 379 – 394.
Holmgren, P. K. & Holmgren, N. H. (1998). Index Herbariorum: A Global Directory of Public Herbaria and Associated Staff. New York Botanical Garden's Virtual Herbarium. http://sweetgum.nybg.org/ih/.
Hooker, J. D. (1885). Phlomis L., Notochaete Benth. & Eremostachys Bunge. Flora of British India, 691 – 695. L. Reeve & Co., London.
Huber-Morath, A. (1982). Phlomis L. In: P. H. Davis (ed.), Flora of Turkey, 102 – 126. Edinburgh University Press, Edinburgh.
Kamelin, R. V. & Makhmedov, A. M. (1990a). [The system of the genus Phlomoides (Lamiaceae)]. Bot. Zhurn. (Moscow & Leningrad) 75: 241 – 250 (in Russian).
_____ & _____ (1990b). [A new system of the genus Phlomis (Lamiaceae)]. Bot. Zhurn. (Moscow & Leningrad) 75: 1163 – 1167 (in Russian).
Knorring, O. E. (1954a). Phlomis L. In: B. K. Shiskin (ed.), Flora of the USSR (English version), pp. 41 – 77. Keter Publishing House, Jerusalem.
_____ (1954b). Eremostachys Bge. In: B. K. Shiskin (ed.), Flora of the USSR (English version), pp. 3 – 41. Keter Publishing House, Jerusalem.
Kudo. Y. (1929). Labiatarum Sino-Japonicarum Prodromus. Mem. Fac. Sci. & Agric., Taihoku Imp. Univ. 2: 1 – 332.
Li, H. W. & Hedge, I. C. (1994a). Lamiophlomis Kudô. In: P. H. Raven & Z. Y. Wu (eds.), Flora of China, 156. Science Press, Beijing & Missouri Botanical Garden Press, St. Louis.
_____ & _____ (1994b). Phlomis Linnaeus. In: P. H. Raven & Z. Y. Wu (eds), Flora of China, 143 – 155. Science Press, Beijing & Missouri Botanical Garden Press, St. Louis.
_____ & _____ (1994c). Notochaete Bentham. In: P. H. Raven & Z. Y. Wu (eds.), Flora of China, 139. Science Press, Beijing & Missouri Botanical Garden Press, St. Louis.
_____ & _____ (1994d). Eremostachys Bunge. In: P. H. Raven & Z. Y. Wu (eds.), Flora of China, 140 – 141. Science Press, Beijing & Missouri Botanical Garden Press, St. Louis.
Lindqvist, C. & Albert, V. A. (2002). Origin of the Hawaiian endemic mints within North American Stachys (Lamiaceae). Amer. J. Bot. 89: 1709 – 1724.
Link, J. H. F. (1829). Handbuch zür Erkennung der nutzbarsten und am häufigsten vorkommenden Gerwächse. Berlin.
Linné, C. Von. (1735). Regnum Vegetabile; Didynamia, Phlomis. Systema Naturae Ed. I. Lugduni batavorum.
_____ (1753). Phlomis. Species Plantarum II: 584 – 587. Laurentii Salvii, Holmiae.
Makhmedov, A. M. (1986). Generis Phlomoides Moench (Lamiaceae) species nova ex Asia media. Novosti Sist. Vyss. Rast. 23: 115 – 116.
_____ (1990). Systema Generis Eremostachys Bunge (Lamiaceae). Novosti Sist. Vyss. Rast. 27: 135 – 137.
Mateu, I. (1986). Revisión del género Phlomis L. (Labiatae) en la Península Ibérica e Islas Baleares. Acta Bot. Malacit. 11: 177 – 204.
Moench, C. (1794). Methodus plantas horti botanici et agri Marburgensis a straminum situ describendi. Marburgi Cattorum.
Müller, K. (2005). SeqState. Nees Institute for Biodiversity of Plant, University of Bonn, Bonn.
Nixon, K. C. (2002). Winclada, Version 1.00.08. Published by the author.
Oxelman, B., Liden, M. & Berglund, D. (1997). Chloroplast rps16 intron phylogeny of the tribe Sileneae (Caryophyllaceae). Pl. Syst. Evol. 206: 393 – 410.
Pan, Y.-Z., Fang, L.-Q., Hao, G., Cai, J. & Gong, X. (2009). Systematic position of Lamiophlomis and Paraphlomis (Lamiaceae) based on nuclear and chloroplast sequences. J. Syst. Evol. 47: 535 – 542.
Pau, C. (1918a). Plantas de Melilla. Bol. Soc. Aragonesa Ci. Nat. 17: 123 – 133.
_____ (1918b). Una ligera visita botaníca a Tous. Butl. Inst. Catalana Hist. Nat. 18: 158 – 161.
_____ (1922). Nueva contribución al estudio de la flora de Almería. Mem. Mus. Ci. Nat. Barcelona, Ser. Bot. 1: 6 – 34.
Popov, M. G. P. (1941). Op-yt monografii roda Eremostachys Bge. Tentamen monographiae generis Eremostachys Bge. Nouv. Mem. Soc. Nat. Mosc. 19: 1 – 166.
Rechinger, K. H. (1982a). Eremostachys. In: Flora Iranica pp. 256 – 292. Akademische Druck-u. Verlagsanstalt, Graz.
_____ (1982b). Phlomis. In: Flora Iranica pp. 292 – 316. Akademische Druck-u. Verlagsanstalt, Graz.
Rivas-Martínez, S., Asensi, A., Molero, J. & Valle, F. (1991). Endemismos vasculares de Andalucía. Rivasgodaya 6: 5 – 76.
Ryding, O. (2008). Pericarp structure and phylogeny of the Phlomis group (Lamiaceae subfam. Lamioideae). Bot. Jahrb. Syst. 127: 299 – 316.
Scheen, A.-C. & Albert, V. A. (2007). Nomenclatural and taxonomic changes within the Leucas clade (Lamioideae; Lamiaceae). Syst. Geogr. Pl. 77: 229 – 238.
_____ & _____ (2009). Molecular phylogenetics of the Leucas group (Lamioideae; Lamiaceae). Syst. Bot. 34: 173 – 181.
_____, Lindqvist, C., Fossdal, C. G. & Albert, V. A. (2007). Molecular phylogenetics of tribe Synandreae, a North American lineage of lamioid mints (Lamiaceae). Cladistics 23: 1 – 16.
_____, Bendiksby, A. M., Ryding, O., Mathiesen, C., Albert, V. A. & Lindqvist, C. (2010). Molecular phylogenetics, character evolution and suprageneric classification of Lamioideae (Lamiaceae). Ann. Missouri Bot. Gard. 97: 191 – 217.
Simmons, M. P. & Ochoterena, H. (2000). Gaps as characters in sequence-based phylogenetic analyses. Syst. Biol. 49: 369 – 381.
Taberlet, P., Gielly, L., Pautou, G. & Bouvet, J. (1991). Universal primers for amplification of three non-coding regions of chloroplast DNA. Pl. Molec. Biol. 17: 1105 – 1109.
Taylor, J. M. (1998). Phlomis. The Neglected Genus. National Council for the Conservation of Plants and Gardens, UK.
Thompson, J. D. (2005). Plant evolution in the Mediterranean. Oxford University Press, New York.
Wagstaff, S. J. & Olmstead, R. G. (1997). Phylogeny of Labiatae and Verbenaceae inferred from rbcL sequences. Syst. Bot. 22: 165 – 179.
_____, Hickerson, L., Spangler, R., Reeves, P. A. & Olmstead, R. G. (1998). Phylogeny in Labiatae s. l., inferred from cpDNA sequences. Pl. Syst. Evol. 209: 265 – 274.
Acknowledgements
The authors thank the A, O, S, UPS, and US herbaria and Jim Mann Taylor (NCCPG National Collection of Phlomis) for providing plant material, and Jan Wesenberg for translation of Russian literature. This research was supported by the Research Council of Norway (grant no. 154145), the University of Oslo (Norway) and University at Buffalo (SUNY, USA).
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Mathiesen, C., Scheen, AC. & Lindqvist, C. Phylogeny and biogeography of the lamioid genus Phlomis (Lamiaceae). Kew Bull 66, 83–99 (2011). https://doi.org/10.1007/s12225-011-9257-0
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DOI: https://doi.org/10.1007/s12225-011-9257-0