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Euphorbia is among the largest genera of angiosperms, with about 2000 species that are renowned for their remarkably diverse growth forms. To clarify phylogenetic relationships in the genus, we used maximum likelihood, bayesian, and parsimony analyses of DNA sequence data from 10 markers representing all three plant genomes, averaging more than 16kbp for each accession. Taxon sampling included 176 representatives from Euphorbioideae (including 161 of Euphorbia). Analyses of these data robustly resolve a backbone topology of four major, subgeneric clades--Esula, Rhizanthium, Euphorbia, and Chamaesyce--that are successively sister lineages. Ancestral state reconstructions of six reproductive and growth form characters indicate that the earliest Euphorbia species were likely woody, non-succulent plants with helically arranged leaves and 5-glanded cyathia in terminal inflorescences. The highly modified growth forms and reproductive features in Euphorbia have independent origins within the subgeneric clades. Examples of extreme parallelism in trait evolution include at least 14 origins of xeromorphic growth forms and at least 13 origins of seed caruncles. The evolution of growth form and inflorescence position are significantly correlated, and a pathway of evolutionary transitions is supported that has implications for the evolution of Euphorbia xerophytes of large stature. Such xerophytes total more than 400 species and are dominants of vegetation types throughout much of arid Africa and Madagascar.
Euphorbia subg. Euphorbia is the largest and most diverse of four recently recognized subgenera within Euphorbia and is distributed across the tropics and subtropics. Relationships within this group have been difficult to discern due mainly to homoplasious morphological characters and inadequate taxon sampling in previous phylogenetic studies. Here we present a phylogenetic analysis of E. subg. Euphorbia, using one nuclear and two plastid regions, for the most complete sampling of molecular sequence data to date. We assign 661 species to the subgenus and show that it is comprised of four highly supported clades, including a single New World clade and multiple independent lineages on Madagascar. Using this phylogenetic framework we discuss patterns of homoplasy in morphological evolution and general patterns of biogeography. Finally, we present a new sectional classification of E. subg. Euphorbia comprising 21 sections, nine of them newly described here.
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RESUMENSe calcula que en México las Euphorbiaceae están representadas por 43 géneros y 782 especies, más 32 taxa intraespecíficos: 8 subespecies y 24 variedades. Se propone una lista preliminar de la familia para el país. Los géneros más grandes son Euphorbia (241 spp., 31%), Croton (124 spp., 16%), Acalypha (108 spp., 14%), Jatropha (48 spp., 6%) y Phyllanthus (41 spp., 5%). Juntos representan 72% de las especies de las Euphorbiaceae conocidas de México. Hay un género endémico del país, Enriquebeltrania y se estima que 57% de las especies mexicanas son endémicas a su territorio. Empleando el concepto de Megaméxico, este valor aumenta a 78%. Los taxa introducidos están representados por un género (Ricinus) y 10 especies. Se destacan dos centros de diversidad que corresponden con las vertientes pacífica y atlántica. A nivel de género, las euforbiáceas mexicanas muestran una gran afinidad con las de Centro y Sudamérica; a nivel de especie, en su mayoría son endémicas, pero de las que no lo son, 200 existen también en Centroamérica, 138 en los Estados Unidos, 85 en Sudamérica y 55 en las Antillas. Casi ninguna de las especies mexicanas se encuentra de manera natural en el Viejo Mundo. A pesar de su gran importancia, tanto sistemática como florística, el conocimiento de las Euphorbiaceae en México es aún sumamente pobre.Palabras clave: diversidad, Euphorbiaceae, México. ABSTRACTIt is estimated that in Mexico the Euphorbiaceae are represented by 43 genera and 782 species. In addition, there are 32 infraspecific taxa: 8 subspecies and 24 varieties. A preliminary list of the taxa is provided. The most diverse genera are Euphorbia (241 spp., 31%), Croton (124 spp., 16%), Acalypha (108 spp., 14%), Jatropha (48 spp., 6%), and Phyllanthus (41 spp., 5%). Together, these represent 72% of the species present in Mexico. There is one endemic genus, Enriquebeltrania. It is estimated that 57% of the Mexican species are endemic to the country. Employing the concept of Megamexico, 78% of the species are endemic. There are ten introduced species and one introduced genus (Ricinus). Within the country, the family is most diverse along the Pacific and Atlantic slopes. At the level of genus, the Mexican Euphorbiaceae show a strong affinity with Central and South America. At the level of species, most Mexican Euphorbiaceae are endemic, but of the species that are not, 200 also occur in Central America, 138 in the United States, 85 in South America, and 55 in the Antilles. Almost no Mexican species are found naturally in the Old World. Despite their great systematic and floristic importance, an adequate understanding of the Mexican Euphorbiaceae is still lacking.
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