Evolució, filogènia i sistemàtica del complex

Abstract

Aquesta tesi se centra en el complex <i>Arctium-Cousinia</i> (Compositae, Cardueae-Carduinae) que comprèn un dels gèneres més amplis de les Compostes, <i>Cousinia</i> Cass. (aproximadament 600 espècies), i altres gèneres menors com <i>Arctium</i> L., <i>Hypacanthium</i> Juz. i <i>Schmalhausenia</i> C. Winkl. Les espècies d'aquest complex es distribueixen principalment per les muntanyes d'Àsia Central (Tien Shan i Pamir-Alai) i la regió Irano-Turània. Al seu torn, <i>Arctium</i> conté algunes espècies cosmopolites. Existeixen importants controvèrsies quant a la delimitació i adscripció d'alguns taxons d'aquest complex d'espècies i s'obtenen dades contradictòries en funció del tipus de dades considerades.<br/><br/>La filogènia generada al present treball, confirma la monofilia del complex <i>Arctium-Cousinia</i> així com la seva subdivisió en dos llinatges també monofilètics anomenats Arctioide i Cousinioide. Al seu torn posa de manifest la manca de monofilia dels gèneres <i>Arctium</i> i <i>Cousinia</i> i de <i>Cousinia</i> subgèneres <i>Cynaroides</i> i <i>Hypacanthodes</i>. La solució que es proposa és la redefinició del gènere Arctium al qual s'han transferit totes les espècies del llinatge Arctioide i s'han proposat els canvis nomenclaturals pertinents. Paral·lelament, mitjançant la tècnica del rellotge molecular, s'ha datat el complex<i> Arctium-Cousinia</i> en 19.1 ma, la separació dels llinatges Arctioide i Cousinioide en 14.8 ma i la diversificació dels esmentats llinatges en 9.7 ma i 8,7 ma, respectivament.<br/><br/>L'estudi sobre la genètica poblacional d'<i>Arctium minus</i>, realitzat mitjançant l'ús de microsatèl·lits dissenyats a aquest treball, ha mostrat que els patrons de variabilitat genètica de l'esmentada espècie són consistents amb les seves característiques biològiques: la seva capacitat d'autofecundació i el seu curt cicle de vida expliquen la seva baixa diversitat genètica, alts valors d'endogàmia i excés d'homozigots. La major part de la variabilitat genètica es troba entre poblacions a causa d'un baix flux gènic entre elles juntament amb la seva capacitat facultativa d'autofecundació. S'han detectat indicis de què <i>A. Minus</i> s'hauria refugiat al sud de la seva actual àrea de distribució durant les glaciacions del Pleistocè i posteriorment hauria recolonitzat Europa. Aquests patrons s'haurien vist emmascarats per influència recent de l'activitat humana i per esdeveniments de dispersió de llavors a llarga distància.<br/><br/><b>English summary:</b><br/>This thesis is focused on the Arctium-Cousinia complex (Compositae, Cardueae-Carduinae), including Arctium L., Cousinia Cass. as one of the largest genera of the Compositae, Hypacanthium Juz. and Schmalhausenia C. Winkl. This group of genera has its highest diversity in the Irano-Turanian region and the mountains of Central Asia (Tien Shan and Pamir-Alaj). Arctium s.str. is Eurosiberian in distribution. Significant difficulties in generic delimitation in this complex arise from the incongruent distribution of several morphological, pollen, karyological and molecular characters.<br/><br/>The phylogeny obtained bears out the monophyly of the Arctium-Cousinia complex and its division in two monophyletic lineages named Arctioid and Cousinioid. Our analyses also show that Arctium and Cousinia as well as Cousinia subgenera Cynaroides and Hypacanthodes are not monophyletic. As a solution, a broadly monophyletic genus Arctium is proposed, including all the species belonging to the Arctioid lineage and we provide nomenclatural changes when necessary. In addition, using the molecular clock approach the Arctium-Cousinia complex is dated in 19.1 mya, the split of the Arctioid and Cousinioid lineages in 14.8 mya and the diversification of the lineages mentioned in 9.7 mya and 8,7 mya, respectively.<br/><br/>Regarding the population genetics of Arctium minus (Hill) Bernh., investigated using microsatellite markers developed for this study, the observed patterns of genetic variation are consistent with its life style. Low gene diversity, high inbreeding values and significant homozygote excess were recovered, factors linked with the self-pollination ability of A. minus and its short-life cycle. Most of the genetic variability is found among instead of within populations due to low gene flow altogether with its self-pollination ability. There are few signs which may suggest that A. minus adapted to the Pleistocene climatic changes in the European populations with putative refugia in the south of its present distribution from where the species might radiate northwards. It is also suggested that original biogeographic patterns in A. minus may have been blurred by human activity and by long distance seed dispersal.

This thesis is focused on the Arctium-Cousinia complex (Compositae, Cardueae-Carduinae), including Arctium L., Cousinia Cass. as one of the largest genera of the Compositae, Hypacanthium Juz. and Schmalhausenia C. Winkl. This group of genera has its highest diversity in the Irano-Turanian region and the mountains of Central Asia (Tien Shan and Pamir-Alaj). Arctium s.str. is Eurosiberian in distribution. Significant difficulties in generic delimitation in this complex arise from the incongruent distribution of several morphological, pollen, karyological and molecular characters.<br/><br/>The phylogeny obtained bears out the monophyly of the Arctium-Cousinia complex and its division in two monophyletic lineages named Arctioid and Cousinioid. Our analyses also show that Arctium and Cousinia as well as Cousinia subgenera Cynaroides and Hypacanthodes are not monophyletic. As a solution, a broadly monophyletic genus Arctium is proposed, including all the species belonging to the Arctioid lineage and we provide nomenclatural changes when necessary. In addition, using the molecular clock approach the Arctium-Cousinia complex is dated in 19.1 mya, the split of the Arctioid and Cousinioid lineages in 14.8 mya and the diversification of the lineages mentioned in 9.7 mya and 8,7 mya, respectively.<br/><br/>Regarding the population genetics of Arctium minus (Hill) Bernh., investigated using microsatellite markers developed for this study, the observed patterns of genetic variation are consistent with its life style. Low gene diversity, high inbreeding values and significant homozygote excess were recovered, factors linked with the self-pollination ability of A. minus and its short-life cycle. Most of the genetic variability is found among instead of within populations due to low gene flow altogether with its self-pollination ability. There are few signs which may suggest that A. minus adapted to the Pleistocene climatic changes in the European populations with putative refugia in the south of its present distribution from where the species might radiate northwards. It is also suggested that original biogeographic patterns in A. minus may have been blurred by human activity and by long distance seed dispersal.