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Redalyc.MORPHOLOGICAL and MOLECULAR EVIDENCE in THE Boletín de la Sociedad Botánica de México ISSN: 0366-2128 [email protected] Sociedad Botánica de México México GÁNDARA, ETELVINA; SOSA, VICTORIA; LEÓN DE LA LUZ, JOSÉ LUIS MORPHOLOGICAL AND MOLECULAR EVIDENCE IN THE DELIMITATION OF BEHRIA AND BESSERA, TWO GENERA OF THE MILLA COMPLEX (THEMIDACEAE) Boletín de la Sociedad Botánica de México, núm. 85, diciembre, 2009, pp. 113-124 Sociedad Botánica de México Distrito Federal, México Available in: http://www.redalyc.org/articulo.oa?id=57712090011 How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative Bol.Soc.Bot.Méx. 85: 113-124 (2009) TAXONOMÍA Y FLORÍSTICA MORPHOLOGICAL AND MOLECULAR EVIDENCE IN THE DELIMITATION OF BEHRIA AND BESSERA, TWO GENERA OF THE MILLA COMPLEX (THEMIDACEAE) ETELVINA GÁNDARA1, 2, VICTORIA SOSA1, JOSÉ LUIS LEÓN DE LA LUZ3 1Biología Evolutiva, Instituto de Ecología, A.C., Apartado Postal 63, 91000 Xalapa, Veracruz, México 3Centro de Investigaciones Biológicas del Noroeste S.C., Apartado Postal 128, 23000 La Paz, Baja California Sur, México 2Corresponding author. E-mail: [email protected] Abstract: Among the taxa of the Milla complex (Themidaceae), a monocot group of perennial petaloid geophytes, there are two genera with controversial taxonomic status: Behria and Bessera. Tree-based and character-based analyses were conducted on 14 populations to determine if they should be considered different taxa. In addition, ecological separation was taken into account. As outgroups representative taxa of Dandya, Jaimehintonia, Milla and Petronymphe, i.e. the rest of the genera in the Milla clade, were used. Behria is a monotypic genus restricted to Baja California and Bessera includes two species from the Pacifi c slopes of Mexico and the Trans-Mexican Volcanic Belt. The chloroplast intergenic spacer psbK-psbI was sequenced and 37 morphological characters were coded. Tree-based analyses retrieved all populations of Behria and all populations of Bessera as monophyletic groups, both forming part of a more inclusive clade. Character-based analysis detected six diagnostic characters, all of which were fl oral. The conclusion of recognizing Behria and Bessera as independent genera was based on these results and also on their different distri- butional ranges. Key words: chloroplast intergenic spacer psbK-psbI, Dandya, Jaimehintonia, petaloid geophytes, Petronymphe. Resumen: Entre los taxones del complejo Milla (Themidaceae), un grupo de monocotiledóneas petaloides geófi tas, existen dos géneros con estatus taxonómico incierto: Behria y Bessera. Con el objetivo de determinar si estos géneros pueden considerarse independientes, se realizaron análisis fi logenéticos y de atributos diagnósticos con base en atributos morfológicos y moleculares de 14 poblaciones. Adicionalmente se consideraron individuos representativos del resto de los géneros del complejo, tales como Dandya, Jaimehintonia, Milla y Petronymphe. Behria es un género monotípico restringido al sur de la Península de Baja California. Bessera comprende dos especies de la vertiente pacífi ca de México y la Faja Volcánica Transmexicana. Se secuenció el espaciador intergénico psbK-psbI del cloroplasto y se codifi caron 37 atributos morfológicos. Los análisis fi logenéticos identifi caron a las pobla- ciones de Behria y Bessera como grupos monofi léticos independientes, formando parte de un clado más inclusivo. Los resultados del análisis de atributos diagnósticos reconocieron seis atributos fl orales. Por lo tanto se concluye que estos taxones pueden ser identifi cados claramente por su morfología fl oral, además tienen diferentes patrones de distribución y se deben considerar como géneros independientes. Palabras clave: espaciador intergénico del cloroplasto psbK-psbI, Dandya, geófi tas petaloides, Jaimehintonia, Milla y Petronymphe. hemidaceae is a plant family of perennial petaloid geo- Milla complex, (2) Brodiaea-Dichelostemma-Triteleiopsis, Tphytes found mainly in western North America (Fay (3) Triteleia-Bloomeria-Muilla clevelandii, and (4) An- and Chase, 1996). The genera currently included in Themi- drostephium-other members of Muilla. daceae were formerly recognized as tribe Brodiaeae in the The Milla clade is supported by two synapomorphic Alliaceae (Dahlgren et al., 1985). In previous taxonomic character states: an ovary stipe adnate to the perianth tube, studies they were divided into two complexes: the Milla and a membranous corm (Pires and Sytsma, 2002). It is complex, centered in Mexico, and the Brodiaea complex, comprised of six genera (Behria, Bessera, Dandya, Jaime- centered in the western United States (Moore, 1953). More hintonia, Milla, and Petronymphe). Behria is a monotypic recently, Pires et al. (2001) and Pires and Sytsma (2002) genus endemic to Baja California (Greene, 1886). In con- found that the genera are grouped into four clades: (1) the trast, Bessera encompasses two species from western and 113 ETELVINA GÁNDARA, VICTORIA SOSA, JOSÉ LUIS LEÓN DE LA LUZ central Mexico (Ramírez-Delgadillo, 1992; Moore, 1953). morphological and cpDNA data (trnL-trnF and rbcL) re- Dandya has four species with a restricted geographic distri- trieved Bessera elegans and B. (Behria) tenuifl ora as sister bution in western and northern Mexico (Espejo-Serna and species, but B. tuitensis was not considered (Pires et al., López-Ferrari, 1992; Lenz, 1971b; Villarreal-Quintanilla 2001). Another study based solely on cpDNA (trnL-F, rpl16, and Encina-Domínguez, 2005). Milla includes ten species and ndhF) retrieved B. elegans and B. tuitensis as sisters, but (Lenz, 1971a; Moore, 1953; Howard, 1999) with M. bifl ora that study did not include B. (Behria) tenuifl ora (Pires and Cav., the most widely distributed member of the group, found Sytsma, 2002). To date, no study has included the two spe- from Arizona to Guatemala (Espejo-Serna and López-Ferrari, cies in Bessera and Behria tenuifl ora simultaneously. 2003; McNeal, 2003). Lastly, Jaimehintonia and Petronym- Wiens and Penkrot (2002) proposed a novel method for phe are both monotypic. The fi rst is restricted to a couple of delimiting species, based on morphological and molecular small areas in Nuevo Leon (Turner, 1993) and the latter is characters. Tree-based and character-based analyses are known from a single locality in Guerrero (Moore, 1951). performed separately and distribution patterns are also Greene (1886) described Behria as a new genus when he considered. This method utilizes populations as terminals was writing the monograph on Brodiaea. Later, Macbride rather than individuals to avoid a biased treatment of the (1918) transferred Behria tenuifl ora to Bessera arguing that polymorphisms shared between populations as homoplasies both Bessera elegans Schult. f. and Bessera (Behria) tenui- rather than synapomorphies in the tree-based morphologi- fl ora possess a red perianth and exerted stamens fused at the cal analyses. The character-based analysis fi nds diagnostic base. According to him, the only difference between the two character states representing differences among the putative taxa was the degree of fusion of fl oral segments. The dispute species. This procedure was designed to delimit closely re- on the status of Behria continued, some authors treat Behria lated taxa. tenuifl ora as part of Bessera (Moore, 1953; Ramírez-Delg- The method of Wiens and Penkrot (2002) is utilized in adillo, 1992; Espejo-Serna and López-Ferrari, 1992; Pires this study to clarify whether Behria and Bessera are inde- et al., 2001), while others place this taxon in its own genus pendent taxa. Thus, tree-based and character-based analyses (Krause, 1930; Lenz, 1971b; Shreve and Wiggins, 1964; were performed with populations of these closely associated León de la Luz and Pérez-Navarro, 2004). groups, together with representative species of genera in the Whether Behria and Bessera are separate genera has not Milla clade as the study units, employing morphological yet been demonstrated. A study of Themidaceae based on characters and chloroplast DNA sequences of psbK-psbI. Table 1. Morphological data matrix. Character and character state number as in Appendix 2. Letters and numbers after the species name are the population abbreviation, indicated in Table 2. “?” = missing data, “-” = not applicable. Polymorphisms: (0, 1) = A; (0, 1, 2) = B; (0, 2) = C; (1, 2) = D; (1, 3) = E; (2, 3) = F; (2, 3, 4) = G Terminals / Characters 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 Behria tenuifl ora BCS1 A 1 0 1 1 1 C 0 0 0 D 1 2 1 1 1 0 0 2 0 0 0 2 0 1 1 0 0 0 0 2 - - - 0 1 2 B. tenuifl ora BCS2 1 1 0 1 1 1 B 0 0 0 1 1 2 1 1 1 0 0 2 0 0 0 2 0 1 1 0 0 0 0 2 - - - 0 1 2 B. tenuifl ora BCS3 ? 1 0 1 1 1 0 0 0 0 1 ? 2 1 ? 1 0 0 2 0 0 0 2 0 1 1 0 0 0 0 2 - - - 0 1 2 B. tenuifl ora BCS4 A 1 0 1 1 1 0 0 0 0 1 ? 2 1 ? 1 0 0 2 0 0 0 2 0 1 1 0 0 0 0 2 - - - 0 1 F Bessera elegans MOR1 1 1 0 1 1 1 0 0 0 0 2 1 2 1 1 1 1 2 1 1 1 - 2 1 0 1 2 0 2 0 2 0 1 1 0 1 2 B. elegans GRO1 0 1 0 1 1 1 0 1 0 1 1 E 2 1 1 1 1 2 1 1 1 - 2 0 0 0 1 0 2 0 2 0 1 1 0 1 2 B. elegans GRO2 1 1 0 1 1 1 0 1 0 0 3 1 2 1 2 1 1 2 1 1 1 - 2 1 0 1 1 0 2 0 3 0 1 1 0 1 3 B.
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