Systematics of Utricularia sect. Foliosa Kamieński. (Lentibulariaceae). Species circumscription and phylogeny. 1
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Paulo C. BALEEIRO , Richard W. JOBSON & Paulo T. SANO 1
Department of Botany – Biosciencie Institute of University of Sao Paulo. St. Rua do Matão, 277. 05508-900 São Paulo, SP Brasil.
[email protected] Phone: **55 – 011-3091-7545 2 National Herbarium of New South Wales, Mrs Macquaries Road, Sydney, NSW 2000, Australia
INTRODUCTION Uncovering biodiversity through classification is important, but not simple. Complexes of taxa and/or cryptic species are defined by different taxa/lineages harbouring limited morphological and/or genetic gaps. Recent molecular phylogenetic studies found the genus Utricularia L. (Lentibulariaceae), along with most of the sectional rankings, to be monophyletic (Jobson & Albert 2002; Müller et al., 2005), although relationships within several sections remained unresolved. One such example exists within the neotropical Utricularia sect. Foliosa, which is composed of three accepted species (sensu Taylor 1989) including U. amethystina Salzm ex A.St.-Hil & Girard, U. tridentata Sylvén and U. tricolor A.St.-Hil. of which the first contains 31 previously recognised species, synonymized after the nomenclatural revision of Taylor (1989). In addition, the last molecular work proposed for the whole Utricularia, suggests the inclusion inside U. sect. Foliosa of the 10 U. sect. Psyllosperma P.Taylor species, based on a questionable paraphyletic species, using just one specimen of U. sect Foliosa. Recently, Baleeiro et al. (in press, 2015) examined the U. sect. Foliosa complex using multivariate analyses which indicated the necessity of the resurrection of synonymised binomials and the addition of potential new taxa. Utricularia is one of the richest genus among carnivorous plants and South America (Cerrado Domain and Guiana Shield) is its centre of diversity (Fig. 1).
MATERIAL AND METHODS Prior to the molecular work, a initial revision of all binomials, including those under synonyms, was undertaken, and specimens were examined like SPF, RB, R, NY, US, F and MO. We show part of Multivariate analysis results for most of morphotypes of U. sect. Foliosa. Our morphometrics study used floral and vegetative characters to asses relationships between morphotypes. We used Principal Component Analysis (PCA) , Canonical Variant Analysis (CVA) , and cluster analyses (Fig. 3). This work was followed up by a molecular phylogeny of U. sect. Foliosa and most members of sister U. sect. Psyllosperma using three chloroplast (rps16, trnL -F, trnD-T), and one nuclear (ITS) region. Analysis of the combined data shows strong support for a sister relationship between monophyletic clades representing both sections. Parsimony, Likelihood and Bayesian analyses were performed with all three methods producing compatible topologies. Here we present the strict consensus tree derived from maximum parsimony analysis using a matrix of the concatenated four regions.
RESULTS AND DISCUSSION Our results show that U. sect. Psyllosperma is a sister to U. sect. Foliosa (JK=100%) (Fig. 2). Within U. sect. Foliosa we find strong support for two major clades, that are also supported by morphological synapomorphies such as presence or absence of trichomes or papillae on the calyx and corolla. One of the clades contains U. amethystina s.s., U. tricolor and U. tridentata, while four new species, and seven resurrected species are found across both clades. The analysis also shows a sister relationship between U. amethystina s.s. and U. tricolor. Our morphometric and molecular results have contributed to a new revision of U. sect. Foliosa that now recognizes 13 species, uncovering previously underestimated diversity. Within these clades, patterns of geographic distribution across Central and South America indicate speciation was possibly driven by isolation factors such as specific phyto-physiognomy (Fig. 1).
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Figure 2. Strict consensus tree from the combined rps16, trnLF, trnDT and ITS analysis. Jackknife support values are shown.
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Figure 01. Centre of diversity for Utricularia A Cerrado Domain and B Guiana Shield, and distribution maps of U. sect. Foliosa species: C. U. amethystina, D. U. tricolor, E. U. tridentata, F. U. lindmanii and U. sp nov., G. U. damazioi, H. U. velascoensis, U. hirtella complex and U. trinervia.
CONCLUSION Both molecular and morphological data were effective in uncovering the diversity hidden in just one taxa U. amethystina, and better explain the taxa relationships of both U. sect. Foliosa and sect. Psyllosperma. C
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REFERENCES Baleeiro PC, Jobson RW & Sano P. 2015. Morphometric approach to address taxonomic problems: The case of Utricularia sect. Foliosa Kamiénski (Lentibulariaceae Rich.). Journal of Systematic and Evolution. (In Press). Jobson RW & Albert VA. 2002. Molecular Rates Parallel Diversification Contrasts between Carnivorous Plant Sister Lineages. Cladistics 18: 127-136. Müller K & Borsch T. 2005. Phylogenetics of Utricularia (Lentibulariaceae) and molecular evolution of the trnK intron in a lineage with high substitutional rates. Plant Systematics and Evolution 250: 3967. Taylor P. 1989. The genus Utricularia – A taxonomic monograph. Kew Bull. Add. Ser., 14: 1-724.
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Figure 3. Multivariate analysis results of Baleeiro et al (in press) for members of U. sect. Foliosa. A. Cluster analysis, B. PCA, C. CVA and D. CVA of more closely related morphotypes.
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