Phylogeny of Central American Bromelioideae (Bromeliaceae)
Claudia J. Ramírez-Díaz1, Ivón Ramírez-Morillo1, Germán Carnevali1, Juan P. Pinzón-Esquivel2, Georg Zizka3 & Nestor Eduardo Raigoza Flores1
1Unidad de Recursos Naturales-Herbario (CICY), Centro de Investigación Científica de Yucatán, A.C., Calle 43 No. 130 Colonia Chuburná de Hidalgo, CP 97205, Mérida, Yucatán, México;
2Departamento de Botánica, Campus de Ciencias Biológicas y Agropecuarias, Universidad Autónoma de Yucatán, Carretera Mérida-Xmatkuil km. 15.5, Apdo, Postal 4-115 Itzimná, CP 97100 Mérida, Yucatán, México; 3Senckenberg Research Institute and Natural History Museum
Frankfurt, Senckenberganlage 25, 60325 Frankfurt/Main, Germany. claujrd11@gmail.com
Most Central American members of Bromelioideae belong to clades that originated and diversified in South America, whereas others diversified in Central America. Molecular evidence suggests that the Central American Clade I (sensu Sass & Specht, 2010, hereafter the Androlepis Alliance, AA) a morphologically variable group that has been classified in four genera, comprises nine species.
Cladistic analyses under MP and BI of nuclear (g3pdh, rpb2, ETS) and plastid (trnL-F) DNA sequences of 220 Bromelioideae taxa, were conducted to test the monophyly and composition of the clade. We also performed a more restricted analysis to test the monophyly of constituent species as currently circumscribed; in this analysis of nuclear and cpDNA (g3pdh, agt1, rpl32 and ycf1) regions of 60 genotypes mostly belonging to the AA were included. We also evaluated the significance of some morphological characters, such as sexual expression and anther ornamentation, in the diversification of this group. We found the AA to comprise at least twelve species.
Topologies from nuclear DNA show strong topological incongruence relative to topologies from plastid DNA. Furthermore, our finding of non-monophyly and morphological intermediacy of AA species suggests hybridization and subsequent plastid capture in the diversification of these species.
Key words: Androlepis Alliance, anther ornamentation, hybridization, plastid capture, sex expression.
Floral morphology still underexploited: the example of the "Cryptanthoid complex"
(Bromeliaceae: Bromelioideae)
Elton Martinez Carvalho Leme1, Sascha Heller2, Georg Zizka3 & Heidemarie Halbritter4
1Research Associate, Marie Selby Botanical Gardens; 2Goethe University; 3Senckenberg Research Institute and Natural History Museum Frankfurt; 4Department of Botany and Biodiversity Research, University of Vienna. leme@tjrj.jus.br
The huge morphological and ecological diversity in Bromeliaceae, together with the high degree of convergent and parallel evolution, and the current limits demonstrated for molecular approaches, despite all advances achieved in the last years, make the taxonomy of the family an exceptional challenge for the taxonomist today. The documentation of under-utilized leaf and floral morphological characters and the combination of geographical range, habitat, as well as leaf trichomes, calyx and corolla conformation, petal appendages, stigma, pollen, fruits and seeds, to name a few, pointed toward the need of a new circumscription of the genera of the Cryptanthus complex. It reveals that geographical range, habitat, and morphological characters, when accessed in detail and combined, played an important role to support genera and species alliances and renew the relevance of morphology based taxonomy in Bromeliaceae.
Key words: Bromelioideae, Cryptanthoid complex, genera alliances, morphology, under-utilized characters.
Limitation of molecular studies in Bromelioideae taxonomy Gregory K. Brown1
1University of Wyoming. gkbrown@uwyo.edu
Molecular studies have had a profound impact on Bromelioideae taxonomy. For example, DNA data have confirmed that Bromelioideae, sensu Smith and Downs (1979), is monophyletic;
consistently place Bromelia, Ochagavia, Deinacanthon, Fascicularia, and Fernseea near the base of the subfamily; and demonstrate the non-monophyletic status of several long-recognized genera such as Aechmea. Unfortunately, the slow evolving bromeliad genome provides a limited number of informative sites and is proving to be largely deficient in resolving species-level relationships, and thus, a robust and well-supported phylogeny for most of Bromelioideae. The problems of low nucleotide substitution rates are compounded by limited sample sizes for most genera, especially larger ones (e.g., Cryptanthus, Hohenbergia, Neoregelia, Orthophytum). Further magnifying these impediments for developing a well-supported phylogeny, and thus a Bromelioideae taxonomy that reflects phylogenetic relationships, is the widespread lack of extensive, detailed morpho-anatomical data for most species. Recommendations are presented for how Bromelioideae phylogenetic studies should be conducted to overcome the now recognized limitations.
Key words: Bromelioideae, phylogeny, sample-size, sequence-data, taxonomy.
Systematics and evolution of Aechmea subg. Chevaliera
Jefferson Rodrigues Maciel1, Rafael Batista Louzada2, Ana Maria Benko-Iseppon3, Georg Zizka4 &
Marccus Alves2
1Jardim Botânico do Recife, Km 7,5 da BR 232, s/n, Curado, 50000-230, Recife, PE, Brazil;
2Universidade Federal de Pernambuco, Laboratório de Morfo-Taxonomia Vegetal, Av. Moraes Rego, s.n., CDU, 50670-930, Recife, PE, Brazil. www.morfotaxonomia.com; 3Universidade Federal de Pernambuco, Centro de Biociências, Departamento de Genética, Laboratório de Genética e Biotecnologia Vegetal, Av. Prof. Moraes Rego, 1235, CDU, 50.670-420, Recife, PE, Brazil;
4Senckenberg Research Institute and Natural History Museum Frankfurt, Department of Botany and Molecular Evolution, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany
Goethe-University, Department of Ecology, Evolution and Diversity, Max-von-Laue-Str. 13, D-60439 Frankfurt am Main, Germany. jeffersonmaciel@recife.pe.gov.br
A better understanding of the phylogeny of Aechmea subgenus Chevaliera is hampered by the limited taxonomic sampling in molecular studies of Bromelioideae. In order to understand the evolution of Aechmea subgenus Chevaliera, we conducted a molecular phylogenetic study using three DNA regions (ETS, matK and phyC). We ran analyses of reconstruction of ancestral states of morphological characters, divergence time of lineages, reconstruction of ancestral area, reconstruction of ancestral habitat, and reconstruction of evolution in environmental space. Our sampling included 22 of the up to 30 species recorded in the subgenus. Aechmea subg. Chevaliera turned out to be clearly polyphyletic. Amazonian species previously included in A. subg. Chevaliera were found in separate clades of Bromelioideae phylogeny. Two distinct groups comprised the majority of species up to now grouped in Chevaliera. The first of which (Aechmea multiflora group) includes seven species; the second (A. sphaerocephala group) comprises 11 species. Both groups are endemics of Atlantic Forest. Morphological and evolutionary dynamics analyses shows that they share the same morphological and geographical spaces, but they had different dynamics of occupation of Atlantic Forest in the last three millions of years.
Key words: Atlantic Forest, biogeography, Bromelioideae, diversification, taxonomy.
New results from the Ochagavia-Fascicularia group using genome skimming
Juraj Paule1, Roswitha Schmickl2, Tomáš Fér3, Sabine Renger1, Georg Zizka1
1Department Botany and Molecular Evolution, Senckenberg Research Institute Frankfurt and Goethe University, Frankfurt/Main, Germany; 2Institute of Botany, The Czech Academy of Sciences, Pruhonice and Department of Botany, Faculty of Science, Charles University, Prague, Czech Republic; 3Department of Botany, Faculty of Science, Charles University, Prague, Czech Republic
Ochagavia (4 species) and Fascicularia (1 species) form a well-supported clade of the early diverging Bromelioideae. The two genera are morphologically similar, but can be easily discerned on the basis of generative characters. Besides the species distributed on the Chilean mainland, the group includes Ochagavia elegans, endemic to the oceanic Isla Robinson Crusoe of the Chilean Juan Fernández Islands. The somewhat enigmatic species morphologically clearly is assigned to Ochagavia, but in previous molecular phylogenetic studies formed mostly a sister clade to the remainder of Fascicularia and Ochagavia. A genomic approach, including whole plastomes (ca 160k bp) and nuclear rDNA cistron (ca 6k bp) was used to analyse the evolution of this clade.
Key words: early-diverging Bromelioideae, Chile, Isla Robinson Crusoe, Ochagavia elegans, plastome.
Genome size evolution in Bromelioideae Juraj Paule1, Sascha Heller1 & Georg Zizka1
1Department Botany and Molecular Evolution, Senckenberg Research Institute and Natural History Museum, Senckenberganlage 25, 60325 Frankfurt/Main, Germany; Institute Ecology, Evolution and Diversity, Goethe University, Max-von-Laue-Straße 13, 60438 Frankfurt/Main, Germany.
jpaule@senckenberg.de
The subfamily Bromelioideae is one of the most diverse groups among the Neotropical plant family Bromeliaceae. Previously, key innovations have been identified which account for the extraordinary radiation and species richness of this subfamily, especially in the so called “core” Bromelioideae.
However, in order to extend our understanding of the evolutionary mechanisms involved, genomic processes underlying “fast” speciation (e.g., polyploidy, dysploidy) also need to be tested. Here, using PI and DAPI staining and flow cytometry we estimated genome size and GC-content of 268 plants covering 30 genera and 192 species sampled in order to cover the most recent phylogeny of the subfamily Bromelioideae. Published and newly collected data from the subfamily were analyzed and interpreted in a phylogenetic framework. The data suggest significant differences between the subfamilies. With respect to polyploidy, contrasting patterns were detected for “core” (“tank”) and
“early diverging” (“tank-less”) Bromelioideae. The results exemplify that even in fast radiating plant groups diverse evolutionary mechanisms could act in closely related clades.
Key words: Bromeliaceae, C-value, flow cytometry, polyploidy, tank habit.