Systématique et biogéographie du groupe Caesalpinia (famille Leguminosae)

Amongst the lineages of the Caesalpinioideae (in the family Leguminosae), one of the largest groups where phylogenetic relationships remains unclear is the Caesalpinia Group, a clade of ca. 200 species, currently considered to comprise between 14 and 21 genera. The taxonomic complexity of the Caesalpinia Group stems from persisting doubts on the generic delimitations within Caesalpinia sensu lato, a group of 150 species that are provisionally classified into eight genera. In order to establish a stable generic classification, phylogenetic analyses of five chloroplastic loci and the nuclear ribosomal ITS locus were carried out on a matrix containing an unprecedented taxonomic sampling of the Caesalpinia Group (~84% of species of this group included), with virtually all of the morphological variation and geographic distribution represented. These analyses allowed us to determine that several genera of the Caesalpinia Group, as currently defined, are polyphyletic or paraphyletic. We consider that there are 26 well-resolved clades that represent distinct genera, and a new generic classification system is proposed, which includes a key to genera, the description of the 26 genera and all species accepted within these groups. A total of twelve previously accepted genera are maintained in this classification (Balsamocarpon, Cordeauxia, Guilandina, Haematoxylum, Hoffmanseggia, Lophocarpinia, Mezoneuron, Pomaria, Pterolobium, Stenodrepanum, Stuhlmannia, and Zuccagnia), whereas two genea are abolished (Stahlia and Poincianella). In addition, two genera are re-instated (Biancaea and Denisophytum), five new genera are described, (Arquita, Gelrebia, Hererolandia, Hultholia and Paubrasilia), and the description of seven genera are emended (Caesalpinia, Cenostigma, Coulteria, Erythrostemon, Libidibia, Moullava, Tara). Our results also indicate that there could possible be a 27th lineage corresponding to the genus Ticanto, but an increased taxonomic sampling is needed to adequately address this issue. The Caesalpinia Group has a pantropical distribution that corresponds almost perfectly to the geographical distribution of the Succulent Biome, but are also found in deserts, grassland prairies, savannahs, and tropical rainforests. On a planetary scale, the Succulent Biome consists of a series of semi-arid to arid habitats that are highly fragmented, and which are characterised by the absence of fire, such as deserts and dry forests. This biome often harbours succulent plant taxa, such as the Cactaceae in the Neotropics and the Euphorbiaceae in Africa. The biogeographical history of the Caesalpinia Group was reconstructed in order to gain insight into the evolution of the flora within this Succulent biome. This biogeographical portrait of this group was reconstructed using molecular dating analysis, diversification rate shifts tests, the reconstruction of ancestral areas using the dispersal-extinction-cladogenesis model (DEC), as well as through ancestral character reconstruction of the biomes and habits. These analyses demonstrate that intercontinental disjunctions between sister species belonging to the same biome are more frequent than the total number of biome shifts across the phylogeny, suggesting that there is a strong conservation of niches, and that it is easier to move than to switch to and evolve in a different biome. Furthermore, contrary to our initial hypothesis, no changes in diversification rates were detected in our phylogenies, even when species switched biomes or evolved a different plant habit, e.g. becoming lianas or herbaceous perennials. We suggest that even when members of the Caesalpinia Group inhabit different biomes, such as savannahs or tropical rainforests, they are still tracking local ecological conditions that are typical of the Succulent biome.   Finally, while total plant species diversity in the Succulent Biome does not compare to the diversity found in tropical rainforests, this biome distinguishes itself by a high number of endemic species, distributed in disjunct patches across the world. This species diversity is probably under-estimated and needs to be carefully re-evaluated, as shown in several recent descriptions of new tree and shrub species from the Succulent biome, all published in the last decade.  The last objective of this thesis is to examine the species limits in Caesalpinia trichocarpa, a shrub from the Andes that has a disjunct population in Peru, which potentially represents a new species. Morphological and molecular analyses of populations occurring across the Andes, including Bolivia and Argentina, allow us to conclude that the populations in Peru represent a new species, which is genetically distinct and has subtle morphological characteristics that allow it to be distinguished from populations found in Argentina and Bolivia. We describe this news species, Arquita grandiflora, in a taxonomic revision of the genus Arquita, a clade of five species found exclusively in Andean valleys.

Keyphrases: Caesalpinia, Andes, Leguminosae, taxonomy, systematics, biogeography, Succulent biome, species delimitation, generic delimitation, seasonally dry tropical forests, phylogeny