A taxonomic revision of Bromeliaceae subfam. Tillandsioideae is presented based on a multi-locus DNA sequence phylogeny (viz., plastid DNA loci rpoB-trnC-petN, trnK-matK-trnK, and ycf1, and the nuclear DNA gene PHYC) and new or re-evaluated morphology (e.g., leaf, inflorescence, sepal, petal, ovary, stigma, stamen, pollen, ovule, and seed morphology). This enables the circumscription of monophyletic units using synapomorphic combinations of diagnostic morphological characters. Stigma morphology has proven to be indicative for super-specific taxa in Tillandsioideae. One new stigma type and several subtypes of previously described stigmas were found. The four tribes proposed earlier are mostly confirmed, but Catopsideae replaces the formerly used name Pogospermeae for the monotypic tribe of Catopsis. In addition, the two new subtribes Cipuropsidinae and Vrieseinae are proposed within tribe Vrieseeae. Several new genera are established to render taxonomic units monophyletic and morphologically well circumscribed. They represent segregates of either Mezobromelia (Gregbrownia: 4 spp.), Tillandsia (viz., Barfussia: 3 spp., Josemania: 5 spp., Lemeltonia: 7 spp., Pseudalcantarea: 3 spp., and Wallisia: 4 spp. and 1 hybrid), or Vriesea (viz., Goudaea: 2 spp., Jagrantia: 1 sp., Lutheria: 4 spp., Stigmatodon: 18 spp., and Zizkaea: 1 spp.). The new subgenera Tillandsia subg. Pseudovriesea and T. subg. Viridantha are established, and T. subg. Aerobia is resurrected. An identification key to all accepted genera of Bromeliaceae subfam. Tillandsioideae is provided. Furthermore, to clarify nomenclatural uncertainties, typifications are proposed for Catopsis subg. Tridynandra, Thecophyllum [unranked] Biflorae, Tillandsia subg. Aerobia, T. sect. Caricifoliae, T. sect. Conostachys, T. sect. Cyathophora, T. sect. Eriophyllum, T. sect. Macrocyathus, T. sect. Platystachys Baker auct. non al., Tillandsia sect. Strepsia, Vriesea subg. Conostachys Mez auct. non al., T. lindenii K. Koch auct. non al., and T. macropetala.
This is the first phylogenetic analysis of the Megamexican Bromeliaceae genus Hechtia and includes 82.6 % of the known taxa. We used plastid (ycf1, rpl32-trnL intergenic spacer), and nuclear (PRK) DNA regions, as well as morphological characters. We generated 244 new sequences for a total of 62 taxa (including 12 species of the outgroup). Results of combined data using parsimony and Bayesian inference reveal the monophyly of Hechtia, as well as identify five well supported clades: (1) a clade (H. tillandsioides complex) as the sister group to the rest of Hechtia; (2) a clade including the species of the H. guatemalensis complex, distributed in Southern Megamexico; the remaining taxa of the genus are retained in a clade which consists of three well-supported clades; (3) the H. glomerata complex distributed in the Gulf of Mexico drainage; (4) a clade of two species (H. deceptrix and H. epigyna) that share an inferior ovary and are distributed north of the Tehuantepec Isthmus in the Sierra Madre Oriental; and (5) an internally poorly resolved clade with the remaining species containing several well-supported, geographically restricted clades. At this time it is uncertain what part of Megamexico was first invaded by the ancestor of Hechtia. Regardless, it becomes clear that from the original point of invasion in what is now Megamexico, it radiated into restricted geographical realms with secondary radiations occurring within them, which resulted in some recurrent particular evolutionary trends most likely associated with the invasion of dry, highly seasonal climates, or cooler areas subject to occasional frosts. Lateral inflorescences and flower morphology suggesting pollination syndromes other than melittophily (psychophily/trochilophily) have evolved more than once in Hechtia.
We performed a phylogenetic analysis using maximum parsimony and Bayesian inference of three plastid DNA markers and the external transcribed spacer (ETS) of nuclear ribosomal DNA to assess the species composition of the Tillandsia utriculata complex and their phylogenetic relationships, and to reconstruct patterns of character evolution and biogeography. The results showed that species of the T. utriculata complex are nested in a clade composed mainly of Mexican and Central American species of T. subgenus Tillandsia (Mexican Clade), and are organized in two lineages: the T. utriculata clade and the T. limbata clade. The ancestor of the core Mexican Clade was probably a T. utriculata-like epiphyte (Group II-type remote flowers and flexuous rachises). The T. utriculata clade is defined morphologically by the presence of acute petals. In this clade, there are two lineages: one of high-elevation, saxicolous, grey-leaved plants from the Mexican Plateau; and one which is more widespread and found from the Gulf of Mexico to Venezuela. The T. limbata clade probably arose in western Mesoamerica and is defined by rounded petals. These species are found mainly in tropical dry forests, but one species colonized wet environments of eastern Mesoamerica. Finally, analyses based on the ETS region allowed us to distinguish between T. utriculata and T. pringlei.
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