Systématique et biogéographie du groupe Caesalpinia (famille Leguminosae)

Parmi les lignées des Caesalpinioideae (dans la famille des Leguminosae), l’un des groupes importants au sein duquel les relations phylogénétiques demeurent nébuleuses est le « groupe Caesalpinia », un clade de plus de 205 espèces, réparties présentement entre 14 à 21 genres. La complexité taxonomique du groupe Caesalpinia provient du fait qu’on n’arrive pas à résoudre les questions de délimitations génériques de Caesalpinia sensu lato (s.l.), un regroupement de 150 espèces qui sont provisoirement classées en huit genres. Afin d’arriver à une classification générique stable, des analyses phylogénétiques de cinq loci chloroplastiques et de la région nucléaire ITS ont été effectuées sur une matrice comportant un échantillonnage taxonomique du groupe sans précédent (~84% des espèces du groupe) et couvrant la quasi-totalité de la variation morphologique et géographique du groupe Caesalpinia. Ces analyses ont permis de déterminer que plusieurs genres du groupe Caesalpinia, tels que présentement définis, sont polyphylétiques ou paraphylétiques. Nous considérons que 26 clades bien résolus représentent des genres, et une nouvelle classification générique du groupe Caesalpinia est proposée : elle inclut une clé des genres, une description des 26 genres et des espèces acceptées au sein de ces groupes. Cette nouvelle classification maintient l’inclusion de douze genres (Balsamocarpon, Cordeauxia, Guilandina, Haematoxylum, Hoffmanseggia, Lophocarpinia, Mezoneuron, Pomaria, Pterolobium, Stenodrepanum, Stuhlmannia, Zuccagnia) et en abolit deux (Stahlia et Poincianella). Elle propose aussi de réinstaurer deux genres (Biancaea et Denisophytum), de reconnaître cinq nouveaux genres (Arquita, Gelrebia, Hererolandia, Hultholia et Paubrasilia), et d’amender la description de sept genres (Caesalpinia, Cenostigma, Coulteria, Erythrostemon, Libidibia, Moullava, Tara). Les résultats indiquent qu’il y aurait possiblement aussi une 27e lignée qui correspondrait au genre Ticanto, mais un échantillonage taxonomique plus important serait nécéssaire pour éclaircir ce problème. Les espèces du groupe Caesalpinia ont une répartition pantropicale qui correspond presque parfaitement aux aires du biome succulent, mais se retrouvent aussi dans les déserts, les prairies, les savanes et les forêts tropicales humides. À l’échelle planétaire, le biome succulent consiste en une série d’habitats arides ou semi-arides hautement fragmentés et caractérisés par l’absence de feu, et abrite souvent des espèces végétales grasses, comme les Cactacées dans les néo-tropiques et les Euphorbiacées en Afrique. L’histoire biogéographique du groupe Caesalpinia a été reconstruite afin de mieux comprendre l’évolution de la flore au sein de ce biome succulent. Ce portrait biogéographique a été obtenu grâce à des analyses de datations moléculaires et des changements de taux de diversification, à une reconstruction des aires ancestrales utilisant le modèle de dispersion-extinction-cladogenèse, et à la reconstruction de l’évolution des biomes et du port des plantes sur la phylogénie du groupe Caesalpinia. Ces analyses démontrent que les disjonctions trans-continentales entre espèces sœurs qui appartiennent au même biome sont plus fréquentes que le nombre total de changements de biomes à travers la phylogénie, suggérant qu’il y a une forte conservation de niches, et qu’il est plus facile de bouger que de changer et d’évoluer au sein d’un biome différent. Par ailleurs, contrairement à nos hypothèses initiales, aucun changement de taux de diversification n’est détecté dans la phylogénie, même lorsque les espèces évoluent dans des biomes différents ou qu’il y a changement de port de la plante, et qu’elle se transforme, par exemple, en liane ou herbacée. Nous suggérons que même lorsqu’ils habitent des biomes très différents, tels que les savanes ou les forêts tropicales humides, les membres du groupe Caesalpinia se retrouvent néanmoins dans des conditions écologiques locales qui rappellent celles du biome succulent. Finalement, bien que la diversité des espèces du biome succulent ne se compare pas à celle retrouvée dans les forêts tropicales humides, ce milieu se distingue par un haut taux d’espèces endémiques, réparties dans des aires disjointes. Cette diversité spécifique est probablement sous-estimée et mérite d’être évaluée attentivement, comme en témoigne la découverte de plusieurs nouvelles espèces d’arbres et arbustes de légumineuses dans la dernière décennie. Le dernier objectif de cette thèse consiste à examiner les limites au niveau spécifique du complexe C. trichocarpa, un arbuste des Andes ayant une population disjointe au Pérou qui représente potentiellement une nouvelle espèce. Des analyses morphologiques et moléculaires sur les populations présentes à travers les Andes permettent de conclure que les populations au Pérou représentent une nouvelle espèce, qui est génétiquement distincte et comporte des caractéristiques morphologiques subtiles permettant de la distinguer des populations retrouvées en Argentine et en Bolivie. Nous décrivons cette nouvelle espèce, Arquita grandiflora, dans le cadre d’une révision taxonomique du genre Arquita, un clade de cinq espèces retrouvées exclusivement dans les vallées andines.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

Morphological Trait Evolution in Solanum (Solanaceae): Evolutionary Lability of Key Taxonomic Characters

Solanum is one of the world\u27s largest and economically most important plant genera, including 1245 currently accepted species and several major and minor crops (e.g., tomato, potato, brinjal eggplant, scarlet eggplant, Gboma eggplant, lulo, and pepino). Here we provide an overview of the evolution of 25 key morphological traits for the major and minor clades of this giant genus based on stochastic mapping using a well-sampled recently published phylogeny of Solanum. The most evolutionarily labile traits (showing \u3e100 transitions across the genus) relate to plant structure (growth form and sympodial unit structure), herbivore defence (glandular trichomes), pollination (corolla shape and colour), and dispersal (fruit colour). Ten further traits show evolutionary lability with 50–100 transitions across the genus (e.g., specialised underground organs, trichome structure, leaf type, inflorescence position and branching, stamen heteromorphism). Our results reveal a number of highly convergent traits in Solanum, including tubers, rhizomes, simple leaves, yellow corollas, heteromorphic anthers, dioecy, and dry fruits, and some unexpected pathways of trait evolution that could be explored in future studies. We show that informally named clades of Solanum can be morphologically defined by trait combinations providing a tool for identification and enabling predictive phylogenetic placement of unsampled species

Phylogenomic discordance suggests polytomies along the backbone of the large genus Solanum

Premise Evolutionary studies require solid phylogenetic frameworks, but increased volumes of phylogenomic data have revealed incongruent topologies among gene trees in many organisms both between and within genomes. Some of these incongruences indicate polytomies that may remain impossible to resolve. Here we investigate the degree of gene-tree discordance in Solanum, one of the largest flowering plant genera that includes the cultivated potato, tomato, and eggplant, as well as 24 minor crop plants. Methods A densely sampled species-level phylogeny of Solanum is built using unpublished and publicly available Sanger sequences comprising 60% of all accepted species (742 spp.) and nine regions (ITS, waxy, and seven plastid markers). The robustness of this topology is tested by examining a full plastome dataset with 140 species and a nuclear target-capture dataset with 39 species of Solanum (Angiosperms353 probe set). Results While the taxonomic framework of Solanum remained stable, gene tree conflicts and discordance between phylogenetic trees generated from the target-capture and plastome datasets were observed. The latter correspond to regions with short internodal branches, and network analysis and polytomy tests suggest the backbone is composed of three polytomies found at different evolutionary depths. The strongest area of discordance, near the crown node of Solanum, could potentially represent a hard polytomy. Conclusions We argue that incomplete lineage sorting due to rapid diversification is the most likely cause for these polytomies, and that embracing the uncertainty that underlies them is crucial to understand the evolution of large and rapidly radiating lineages.Peer reviewe

Phylogenomic Discordance Suggests Polytomies Along the Backbone of the Large Genus Solanum

Premise of the study Evolutionary studies require solid phylogenetic frameworks, but increased volumes of phylogenomic data have revealed incongruent topologies among gene trees in many organisms both between and within genomes. Some of these incongruences indicate polytomies that may remain impossible to resolve. Here we investigate the degree of gene-tree discordance in Solanum, one of the largest flowering plant genera that includes the cultivated potato, tomato, and eggplant, as well as 24 minor crop plants. Methods A densely sampled species-level phylogeny of Solanum is built using unpublished and publicly available Sanger sequences comprising 60% of all accepted species (742 spp.) and nine regions (ITS, waxy, and seven plastid markers). The robustness of this topology is tested by examining a full plastome dataset with 140 species and a nuclear target-capture dataset with 39 species of Solanum (Angiosperms353 probe set). Key results While the taxonomic framework of Solanum remained stable, gene tree conflicts and discordance between phylogenetic trees generated from the target-capture and plastome datasets were observed. The latter correspond to regions with short internodal branches, and network analysis and polytomy tests suggest the backbone is composed of three polytomies found at different evolutionary depths. The strongest area of discordance, near the crown node of Solanum, could potentially represent a hard polytomy. Conclusions We argue that incomplete lineage sorting due to rapid diversification is the most likely cause for these polytomies, and that embracing the uncertainty that underlies them is crucial to understand the evolution of large and rapidly radiating lineages

Global urban environmental change drives adaptation in white clover

Urbanization transforms environments in ways that alter biological evolution. We examined whether urban environmental change drives parallel evolution by sampling 110,019 white clover plants from 6169 populations in 160 cities globally. Plants were assayed for a Mendelian antiherbivore defense that also affects tolerance to abiotic stressors. Urban-rural gradients were associated with the evolution of clines in defense in 47% of cities throughout the world. Variation in the strength of clines was explained by environmental changes in drought stress and vegetation cover that varied among cities. Sequencing 2074 genomes from 26 cities revealed that the evolution of urban-rural clines was best explained by adaptive evolution, but the degree of parallel adaptation varied among cities. Our results demonstrate that urbanization leads to adaptation at a global scale

Reinstatement of Ticanto (Leguminosae-Caesalpinioideae) – the final piece in the Caesalpinia group puzzle.

A recent molecular phylogenetic analysis of the Caesalpinia group demonstrated that it comprises 26 genera, but the recognition of a putative 27 genus, , remained in doubt. This study presents a phylogenetic analysis of ITS and five plastid loci revealing a robustly supported monophyletic group representing the Ticanto clade, sister to the morphologically distinct genus . Based upon this evidence, along with a morphological evaluation, the genus is here reinstated. Descriptions are provided for all nine species of , together with a key to the species, maps, and colour photographs. Nine new combinations are made: (Hand.-Mazz.) R. Clark & Gagnon, (L.) R. Clark & Gagnon, (S. J. Li, Z. Y. Chen & D. X. Zhang) R. Clark & Gagnon, (Metcalf) R. Clark & Gagnon, R. Clark & Gagnon, (Hemsl.) R. Clark & Gagnon, (Craib) R. Clark & Gagnon, (Champion ex Benth.) R. Clark & Gagnon and (S. J. Li, D. X. Zhang & Z.Y. Chen) R. Clark & Gagnon. The final major question in the delimitation of segregate genera from within and the Caesalpinia group is thus resolved

A new cryptic species in a new cryptic genus in the Caesalpinia group (Leguminosae) from the seasonally dry inter-Andean valleys of South America

The generic affiliation of the Andean species Caesalpinia trichocarpa, C. mimosifolia, and their close relatives has remained uncertain in all recent studies of Caesalpinia s.l. (Leguminosae, subfamily Caesalpinioideae). A new densely sampled phylogeny based on four DNA sequence regions (rps16, trn D-trnT, ycf6- psbM, ITS) strongly supports the monophyly of an Andean clade. We propose that despite the lack of obvious diagnostic morphological synapomorphies, this Andean group should be considered as a distinct genus, here described as the new genus Arquita. Phylogenetic analyses also suggest a problem with species delimitation in this group. Within C. trichocarpa, accessions from disjunct geographic areas in Argentina, Bolivia and Peru each form a robustly supported, unresolved clade that includes C. mimosifolia. The morphological and genetic cohesiveness of the C. trichocarpa complex is investigated using morphometric phenetic analyses of qualitative and quantitative flower and leaf traits, and reconstruction of a densely sampled phylogeny using three plastid and one nuclear ribosomal DNA sequence loci. Our results suggest that the most geographically isolated of these clades, narrowly endemic to two inter-Andean valleys in central-north Peru and separated by ∼1350 km, and extensive high Andean cordilleras above 4000 m, from the nearest populations in Bolivia, represents a genetically highly distinct and morphologically cryptic lineage here described as a new species (Arquita grandiflora). A full taxonomic account of the new genus Arquita and its component species is provided, with a distribution map and a key to the species

Revisiting the cytomolecular evolution of the Caesalpinia group (Leguminosae): a broad sampling reveals new correlations between cytogenetic and environmental variables

The pantropical Caesalpinia group includes 225 species in 27 monophyletic genera, and the group has undergone recent phylogenetic, taxonomic and biogeographic revisions. Previous works have reported a diverse pattern of heterochromatin distribution related to ecological niche/geographic distribution, and variation in genome size also correlated with environmental variables. In order to investigate the relationship between cytogenetic and ecological traits using the Caesalpinia group as a model, new cytomolecular data (chromosome number and morphology, CMA/DAPI staining and number and position of 5S and 35S rDNA sites) for 14 species in six genera were generated. These data were analysed by phylogenetic comparative methods. All species studied have 2n = 24 (16 M/SM + 8A), and most of them just have one pair of 5S rDNA sites and two to five pairs of 35S rDNA sites. Three heterochromatic patterns were observed on the chromosomes: (i) proximal CMA+/DAPI− bands, (ii) proximal CMA0/DAPI− bands and (iii) proximal CMA0/DAPI0 bands. The “Coulteria + Tara” and “Arquita + Balsamocarpon + Erythrostemon + Pomaria” clades (except for E. gilliesii, E. hughesii and E. mexicanus) independently showed CMA0/DAPI− bands associated with larger genomes and geographic distributions at higher latitudes. We statistically demonstrate that heterochromatin (CMA/DAPI intensity along the chromosome), genome size and latitude are autocorrelated in the Caesalpinia group. On the other hand, we found a non-significant correlation between genome size and amount of heterochromatin. We argue that environmental factors associated with different latitude may have played a role in contributing to the diversification of the heterochromatin in Caesalpinia group