Anales del Jardín Botánico de Madrid, Vol 65, No 2 (2008)

Warionia (Asteraceae): a relict genus of Cichorieae?

Liliana Katinas
División Plantas Vasculares, Museo de La Plata , Argentina

María Cristina Tellería
Laboratorio de Sistemática y Biología Evolutiva, Museo de La Plata , Argentina

Alfonso Susanna
Instituto Botánico de Barcelona , Spain

Santiago Ortiz
Laboratorio de Botánica, Facultade de Farmacia, Universidade de Santiago , Spain


The genus Warionia, with its only species W. saharae, is endemic to the northwestern edge of the African Sahara desert. This is a somewhat thistle-like aromatic plant, with white latex, and fleshy, pinnately- partite leaves. Warionia is in many respects so different from any other genus of Asteraceae, that it has been tentatively placed in the tribes Cardueae, Cichorieae, Gundelieae, and Mutisieae. Until now, a comprehensive study of Warionia to have a complete context for discussing its taxonomic position is lacking. The general morphology, anatomy, palynology and chromosome number of W. saharae are investigated here, and the species is described and illustrated. Laticifers in leaves and stems indicate a relationship with Cichorieae, and are associated with the phloem, in contact with it or with the surrounding sclerenchyma sheath. The pollen features indicate a strong relation with Cardueae, namely the structure with Anthemoid pattern where the columellae are joined to the foot layer, the ectosexine with thin columellae, the endosexine with stout and ramified columellae, the conspicuous spines with globose bases and conspicuous apical channels, and the tectum surface very perforate. Chromosomal counts resulted in 2n = 34. The morphological and palynological evidence positions Warionia between the tribes Cardueae and Cichorieae suggesting that it could be a remnant of the ancestral stock that gave rise to both tribes.


Africa; Cardueae; chromosome number; Cichorieae; laticifers; pollen; taxonomy

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Audissou, J.-A. 1999. Warionia saharae. British Cactus and Succulent Journal 17: 124-126.

Augier, J. & du Mérac, M.-L.R. 1951. La phylogénie des Composées. Revue Scientifique 89: 167-182.

Bellakhdar, J., Baayaoui, A., Kazdari, A. & Marechal, J. 1986. Herboristes et medecine traditionnelle à Tissint, oasis presaharien du sud Marocain (province de Tata). Al Biruniya 3: 7-50.

Bentham, G. & Cosson, E. 1872. Compositarum genus novum algeriense. Bulletin de la Société Botanique de France 19: 165-166.

Blackmore, S. 1984. Compositae-Lactuceae. Review of Palaeobotany and Palynology 42: 45-85. doi:10.1016/0034-6667(84)90062-9

Bremer, K. 1994. Asteraceae: Cladistics and classification. Timber Press, Portland, Oregon.

Cabrera, A.L. 1977. Mutisieae-systematic review. In: Heywood, V.H. & al. (eds.), The Biology and Chemistry of the Compositae 2: 1039-1066. Academic Press, London.

Carlquist, S. 1958. Anatomy of the Guayana Mutisieae. Part II. Memoirs of the New York Botanical Garden 10: 157-184.

Dimon, M.T. 1971. Problèmes généraux soulevés par l’étude pollinique de Composées Méditerranéennes. Naturalia monspeliensis, sér. Bot. 22: 129-144.

Dittrich, M. 1977. Cynareae-systematic review. In: Heywood, V.H. & al. (eds.), The Biology and Chemistry of the Compositae 2: 999- 1015. Academic Press, London.

Funk, V.A., Chan, R., & Keeley, S.C. 2004. Insights into the evolution of the tribe Arctoteae (Compositae: subfamily Cichorioideae s.s.) using trnL-F, ndhF and ITS. Taxon 53: 637-655. doi:10.2307/4135440

Funk, V.A., Bayer, R.J., Keeley, S., Chan, R., Watson, L., Gemeinholzer, B., Schilling, E., Panero, J.L., Baldwin, B.G., García- Jacas, N., Susanna, A. & Jansen, R.K. 2005. Everywhere but Antarctica: Using a supertree to understand the diversity and distribution of the Compositae. Biologiske Skrifter 55: 343-374.

Goertzen, L.R., Cannone, J.J., Gutell, R.R. & Jansen, R.K. 2003. ITS secondary structure derived from comparative analysis: implications of sequence alignment and phylogeny of Asteraceae. Molecular Phylogenetics and Evolution 29: 216-234. doi:10.1016/S1055-7903(03)00094-0 PMid:13678678

Gómez, F. 2001. Flora selecta marroquí. Francisco José Nevero Díez ed., España.

Hansen, H.V. 1991a. Phylogenetic studies in Compositae tribe Mutisieae. Opera Botanica 109: 1-50.

Hansen, H.V. 1991b. SEM-studies and general comments on pollen in the tribe Mutisieae (Compositae) sensu Cabrera. Nordic Journal of Botany 10: 607-623. doi:10.1111/j.1756-1051.1991.tb02102.x

Hilmi, F. 2002. Cytotoxic and anti-inflamatory sesquiterpene lactones from Warionia saharae, a traditional remedy from Morocco. PhD. thesis, Swiss Federal Institute of Technology Zurich, Zurich, Switzerland.

Holmgren P.K., Holmgren, N.H. & Barnett, L.C. 1990. Index Herbariorum, Part I: The herbaria of the world, 8th ed. New York Botanical Garden, Bronx, New York.

Humphries, C.J., Murray, B.G., Bocquet, G. & Vasudevan, K. 1978. Chromosome numbers of phanerogams from Morocco and Algeria. Botaniska Notiser 131: 391-406.

Jeffrey, C. 2007a. Compositae: Introduction with key to tribes. In: Kadereit, J.W. & Jeffrey, C. (eds.), Flowering Plants. Eudicots. Asterales, vol. VIII: 61-87. In: Kubitzki, K. (ed.), The Families and Genera of Vascular Plants. Springer Verlag, Berlin and Heidelberg.

Jeffrey, C. 2007b. Carduoid genera of uncertain placement. In: Kadereit, J.W. & Jeffrey, C. (eds.), Flowering Plants. Eudicots. Asterales, vol. VIII: 146-147. In: Kubitzki, K. (ed.), The Families and Genera of Vascular Plants. Springer Verlag, Berlin and Heidelberg.

Karis, P.O., Källersjö, M. & Bremer, K. 1992. Phylogenetic analysis of the Cichorioideae (Asteraceae), with emphasis on the Mutisieae. Annals of the Missouri Botanical Garden 79: 416-427. doi:10.2307/2399778

Karis, P.O., Eldenäs, P. & Källersjo, M. 2001. New evidence for the systematic position of Gundelia L. with notes on delimitation of Arctoteae (Asteraceae). Taxon 50: 105-114. doi:10.2307/1224514

Katinas, L., Pruski, J., Sancho, G. & Tellería, M.C. (in press). The subfamily Mutisioideae. The Botanical Review.

Lack, H.W. 2007. VIII. Tribe Cichorieae Lam. & DC. In: Kadereit, J.W. & Jeffrey, C. (eds.), Flowering Plants. Eudicots. Asterales, vol. VIII: 180-199. In: Kubitzki, K. (ed.), The Families and Genera of Vascular Plants. Springer Verlag, Berlin and Heidelberg.

Levin, D.A. 2002. The Role of Chromosomal Change in Plant Evolution. Oxford University Press, Oxford.

Lewalle, L. 1986. Warionia saharae Cosson. Al Biruniya 3: 77-78.

Lewinsohn, T.M. 1991. The geographical distribution of plant latex. Chemoecology 2: 64-68. doi:10.1007/BF01240668

Mabry, T.J. & Bohlmann, F. 1977. Summary of the chemistry of the Compositae. In: Heywood, V.H . & al. (eds.), The Biology and Chemistry of the Compositae 2: 1097-1104. Academic Press, London.

Metcalfe, C.R. & Chalk, L. 1965. Anatomy of the Dicotyledons 2. Clarendon Press, Oxford.

Oberprieler, C. & Vogt, R. 1993. Chromosome numbers of north African phanerogams. II. Willdenowia 23: 211-238.

Ortiz, S. 2000. A phylogenetic analysis of Dicoma Cass. and related genera (Asteraceae: Cichorioideae: Mutisieae) based on morphological and anatomic characters. Annals of the Missouri Botanical Garden 87: 459-481. doi:10.2307/2666141

Ortiz, S. 2001. The reinstatement of the genus Macledium Cass. (Asteraceae, Mutisieae): morphological and phylogenetic arguments. Taxon 50: 733-744. doi:10.2307/1223704

Ozenda, P. 1991. Flore et végétation du Sahara, 3º ed. Editions du Centre National de la Recherche Scientifique, Paris.

Panero, J.L. & Funk, V. 2008. The value of sampling anomalous taxa in phylogenetic studies: Major clades of the Asteraceae revealed. Molecular Phylogenetics and Evolution 47: 757-782. doi:10.1016/j.ympev.2008.02.011 PMid:18375151

Punt, W., Blackmore, S., Nilsson, S. & Le Thomas, A. 1994. Glossary of pollen and spores terminology. Laboratory of Palaeobotany and Palynology, Contribution Series 1: 1-71. University of Utrecht.

Reese, G. 1957. Uber die Polyploidiespektren in der nordsaharischen Wustenpflanzen. Flora 144: 598-634.

Riesberg, L.H., Whitton, J. & Gardner, K. 1999. Hybrid zones and the genetic architecture of a barrier to gene flow between two sunflower species. Genetics 152: 713-727.

Robinson, H. 1984. Style rotation in the Asteraceae. Taxon 33: 400- 404. doi:10.2307/1220977

Robinson, H. 1994. Notes on the tribes Eremothamneae, Gundelieae, and Moquinieae, with comparisons of their pollen. Taxon 43: 33-44. doi:10.2307/1223458

Skvarla, J.J. Turner, B.L., Patel, U.C. & Tomb, S. 1977. Pollen morphology in the Compositae and in morphologically related families. In: Heywood, V. & al. (eds.), The Biology and Chemistry of the Compositae 1: 141-248. Academic Press, London.

Skvarla, J.J., Rowley, J.R. Chissoe, W.F. & Strout, G. 2001. Incomplete exine development in aborted pollen of Eupatorium serotinum Michx. (Compositae: Eupatorieae). Taiwania 46: 103-113.

Stebbins G.L. 1971. Chromosomal evolution in higher plants. Edward Arnold, London.

Tormo, R. & Ubera, J.L. 1995. Tipos polínicos de la Tribu Cardueae en la península Ibérica. Monografías del Jardín Botánico de Córdoba 2: 1-52.

Strittmater, C.G. Dizeo de. 1973. Nueva técnica de diafanización. Boletín de la Sociedad Argentina de Botánica 15: 126-129.

Wagenitz, G. 1955. Pollenmorphologie und Systematic in der Gattung Centaurea L. s.l. Flora 142: 213-279.

Watanabe, K. (accessed 2008). Index to chromosome numbers in Asteraceae.

Watillon, C., Gaspar, T., Hofinger, M. & Ramaut, J.-L. 1987. La micropropagation de Warionia saharae Benth. & Coss. Al Biruniya 4: 35-38.

Wortley, A.H., Funk, V.A., Robinson, H., Skvarla, J. & Blackmore, S. 2007. A search for pollen morphological synapomorphies to classify rogue genera in Compositae (Asteraceae). Review of Palaeobotany and Palynology 146: 169-181. doi:10.1016/j.revpalbo.2007.03.003

Zao, Z., Skvarla, J.J. & Jansen, R.K. 2006. Mutisieae (Asteraceae) pollen ultrastructure atlas. Lundellia 9: 51-76.

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