Phylogenetic position of Echeveria heterosepala (Crassulaceae): a rare species with diagnostic characters of Pachyphytum
<p><strong>Background: </strong><em>Echeveria</em> and <em>Pachyphytum</em> are two closely related Neotropical genera in the Crassulaceae. Several species in <em>Echeveria</em> possess characters cited as diagnostic for <em>Pachyphytum</em> such as a clearly defined stem, a nectary scale on the inner face of petals and as inflorescence a scorpioid cyme or cincinnus. <em>Pachyphytum</em> has been identified as monophyletic while <em>Echeveria</em> as polyphyletic in previous molecular phylogenetic analysess.</p><p><strong>Hypothesis: </strong>The objective of this paper is to identify the phylogenetic position of a rare species with restricted distribution in <em>Echeveria</em>, <em>E. heterosepala</em> that possesses the diagnostic characters of <em>Pachyphytum</em> to better understand the generic limits between these two genera. We expect this species to be closely related to <em>Pachyphytum</em>.</p><p><strong>Methods</strong>: Bayesian inference and Maximum Likelihood analyses were carried out using 47 taxa, including as ingroup, species of <em>Echeveria, Graptopetalum, Lenophyllum, Pachyphytum, Sedum</em>, <em>Thompsonella</em> and <em>Villadia</em> and as outgroup, species in<em> Dudleya</em>. Analyses were conducted based on plastid (<em>rpl16, trnL-F</em>) and nuclear (ETS, ITS) markers. Ancestral character reconstruction was carried out under a parsimony criterion based on the molecular trees retrieved by the phylogenetic analyses. Four morphological characters were considered: defined stem, type of inflorescence, nectary scale in petals and position of sepals.</p><strong>Results</strong>: Accessions of <em>E</em>. <em>heterosepala</em> were retrieved in a well-supported clade group comprising exclusively <em>Echeveria</em> species. Therefore this species belongs to <em>Echeveria</em> although possessing characters similar to <em>Pachyphytum</em> and moreover it was not identified closely related to this genus. None of the traits considered have taxonomic importance. The node at the <em>Pachyphytum</em> clade identified unambiguous character states such as stem present, straight sepals, nectary scale on petals, however these character states were identified in the rest of the clades as well. Remarkably, the monophyly of <em>Pachyphytum</em> was corroborated, while <em>Echeveria</em> remains poorly understood.