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(Ericales) with Special Emphasis on the Genus Purdiaea Planch

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(Ericales) with Special Emphasis on the Genus Purdiaea Planch Org. Divers. Evol. 2, 127–137 (2002) © Urban & Fischer Verlag http://www.urbanfischer.de/journals/ode Phylogenetic relationships of Cyrillaceae and Clethraceae (Ericales) with special emphasis on the genus Purdiaea Planch. Arne A. Anderberg1,*, Xiaoping Zhang1, 2 1 Department of Phanerogamic Botany, Swedish Museum of Natural History, Stockholm 2 Department of Biology,Anhui Normal University, China Received 14 February 2002 · Accepted 15 April 2002 Abstract The phylogenetic relationships and systematic position of the three genera of Cyrillaceae (Ericales), Cyrilla, Cliftonia, and Purdiaea, were inves- tigated by jackknife analysis of a combination of DNA sequences from the plastid genes atpB, ndhF,and rbcL.The results show that Cyrilla and Cliftonia together are the sister group of Ericaceae, whereas Purdiaea groups with Clethra of Clethraceae.Together, Clethra and Purdiaea form the sister group of (Cyrilla+Cliftonia)+Ericaceae. It is concluded that Purdiaea should be moved to Clethraceae. A cladistic analysis based on morphological data was performed to investigate relationships among the species of Purdiaea. The results indicate that Purdiaea belizensis from Central America is sister to all other species of the genus, and that Purdiaea nutans from northern South America is sister to the remaining group of species which are all Cuban endemics, among which P. cubensis from Pinar del Rio in western Cuba is sister to the eight species oc- curring in the Oriente province in eastern Cuba. Key words: Clethraceae, Cyrillaceae, DNA, Ericales, morphology, phylogeny Introduction ers, more detailed sampling was needed for a better un- derstanding of phylogenetic relationships within Cyril- In a recent paper on phylogenetic relationships among laceae and with other families in this clade. Many authors families of the Ericales (Anderberg et al. 2002), a mono- (e.g. Cronquist 1981, Dahlgren 1989, Takhtajan 1997) phyletic group formed by the six ericoid families Actini- have pointed to the similarity between Clethraceae and diaceae, Roridulaceae, Sarraceniaceae, Clethraceae, Cyrillaceae, and particularly the genera Clethra L. and Cyrillaceae, and Ericaceae was identified. The first three Purdiaea share a number of features, which Anderberg of these families constituted the sister group of the re- (1993) discussed as possible symplesiomorphic similari- maining three among which Clethraceae was found to be ties. Such character states include the presence of invert- sister to Cyrillaceae+Ericaceae. Cyrillaceae, in its tradi- ed, versatile, poricidal anthers and sympetalous corollas tional sense, has three genera. Cyrilla L. is found from (Fig. 1 A–C), that stand in contrast to the non-inverted southern North America to northern South America. In anthers opening with slits, and to the choripetalous flow- this genus a number of species has been described, but ers of Cyrilla and Cliftonia (Fig. 1 D–E), although most Thomas (1960) recognized only one variable species. Clethra also have choripetalous flowers. Mabberley Cliftonia Gaertn. f. has a single species in southeastern (1997) listed Clethraceae as a synonym of Cyrillaceae, North America, and Purdiaea Planch. has 11 species in including Clethra as a fourth genus of Cyrillaceae, and it South and Central America, most of which grow in east- is noteworthy that one species of Purdiaea, viz. P. be- ern Cuba. The study of Anderberg et al. (2002) included lizensis (A. C. Smith & Standley) J. L. Thomas was orig- only Cyrilla as representative of Cyrillaceae. Since this inally described as a second genus of Clethraceae, albeit genus is different in some respects from the two other with some hesitation (Schizocardia Smith & Standley genera in the family, e.g. in having haplostemonous flow- 1932, Standley & Record 1936, Thomas 1961). *Corresponding author: Arne A. Anderberg, Department of Phanerogamic Botany, Swedish Museum of Natural History, P.O. Box 50007, SE-104 05 Stockholm, Sweden; e-mail: [email protected] 1439-6092/02/02/02-127 $ 15.00/0 Org. Divers. Evol. (2002) 2, 127–137 128 Anderberg & Xiaoping Fig. 1. Flower morphology.A: Clethra alnifolia.A1: Stamen.A2. Part of opened flower with two stamens removed. B: Purdiaea belizensis. B1: Stamen. B2: Part of opened flower with four stamens removed. C: Purdiaea nutans. C1: Stamen. C2: Part of opened flower with four stamens removed. - A: Brumbach 8605 (S). B: Monro & Cafferty 2615 (MO). C: Harling & Andersson 22028 (S). Org. Divers. Evol. (2002) 2, 127–137 Phylogenetic relationships of Cyrillaceae and Clethraceae 129 Fig. 1 (continued). Flower morphology. D: Cliftonia monophylla. D1: Stamen. D2: Petal. E. Cyrilla racemiflora. E1: Stamen. E2: Petals. - D: Bilt- more herbarium 5979b (S). E: Segel & Thompson 193 (S). If the observed similarity between Purdiaea and phylla (Lam.) Britt.) to the plastid gene data set (atpB, ndhF, Clethra is symplesiomorphic, the former would proba- and rbcL) of a previous study (Anderberg et al. 2002), com- bly be the sister group of the two other genera in the prising a larger sample from all families recognized in the Cyrillaceae. In the present investigation, our aims were order Ericales by APG (1998) plus Sladeniaceae and Pentaphy- to test the position of the family in relation to the sur- lacaceae. Two mitochondrial genes (atp1 and matR) that were also included in the study by Anderberg et al. (2002) did not rounding families identified by Anderberg et al. (2002), give PCR products with Purdiaea, and could therefore not be the monophyly of Cyrillaceae, and the relationships used in the present study. The morphological investigations within genus Purdiaea. were based on herbarium material from the following herbaria, abbreviated as in Holmgren et al. (1990): BM, BP, GH, MO, NY, S. Requests for loans of herbarium material were also sent Materials and methods to three Cuban herbaria, but no replies were obtained. Taxa Molecular methods We added eight new sequences from three species (Purdiaea DNA was extracted from leaves taken from herbarium speci- nutans Planch., P. stenophylla Grieseb., and Cliftonia mono- mens [Purdiaea nutans, Harling and Andersson 22028 (S)], ma- Org. Divers. Evol. (2002) 2, 127–137 130 Anderberg & Xiaoping terial collected in silica gel [Purdiaea stenopetala Grieseb., found in the ndhF sequences were treated as missing information. Cuba, Gutiérrez & Nilsson 9 (S)], or fresh leaves [Cliftonia Analyses were performed with all codon positions included, as monophylla, Anderson 19705 (S)]. Leaves were ground with a this has been demonstrated to give the best resolution and a high- Mini-Bead Beater (BioSpec Products, Bartlesville, Oklahoma, er number of supported groups than analyses of first and second U.S.A.) and subsequently treated with the DNEasy plant DNA codon positions, or of transversions only (Källersjö et al. 2000). extraction kit from Qiagen (Qiagen Inc., Valencia, California, The morphological data set (Table 1) was also analysed U.S.A.), following the manufacturer’s protocol. PCR was per- with the “Xac” software, using the same settings as described formed with 10 µmol/l primers in 25-µl reactions using “Ready- above. Species delimitation in Cuban Purdiaea followed Be- to-go” PCR beads from Pharmacia Biotech (Amersham Phar- razain & Rodrigues (1992). The morphological analysis used macia Biotech, Uppsala, Sweden) following the manufacturer’s Cyrilla and Cliftonia as outgroup, based on the results of the standard protocol and suggested thermal cycling profile, gener- analysis of molecular data. One species of Clethra (C. alnifo- ally 95 °C for 5 min, followed by 35 cycles of 95 °C 30 s, 50 °C lia L.) was also included. A parsimony analysis was also per- 30 s, 72 °C 1 min 30 s, and finally 72 °C for 8 min. formed on the morphological data set, using Hennig86 (Farris For sequencing reactions the “Big Dye Terminator Sequenc- 1988) and the following options: mh* (constructing several ing” kit (Applied Biosystems, Warrington, Cheshire, UK) was initial cladograms by adding taxa in several different se- used, and fragments were separated on an ABI377 from Ap- quences, and retaining the shortest cladogram of each) with a plied Biosystems. Primers used for PCR and for sequencing subsequent bb* (generating multiple equally parsimonious were the same as those described by Källersjö et al. (2000). Se- cladograms). All characters were coded as non-additive (cc-.). quence fragments were assembled with the Staden software (Staden et. al. 1998), and aligned manually with the Assem- blyLign software (Oxford Molecular Group Inc., Campbell, Results California, U.S.A.). Alignment was unproblematic with only a few small gaps in ndhF. The new sequences have been submit- The jackknife analysis of the molecular data, with the ted to GenBank (Accession numbers AY082692-AY082699). three genes atpB, ndhF, and rbcL in combination, result- ed in a tree (Fig. 2) with the same basic tree topology as Phylogenetic analyses the one obtained for the three plastid genes by Ander- Two different data sets were analyzed. First an analysis of the berg et al. (2002), and congruent with their more re- molecular data was analyzed to find the systematic positions solved five-gene tree. The close relationship between the of Purdiaea and Cliftonia in relation to Cyrilla, Clethra and families Actinidiaceae, Sarraceniaceae, Roridulaceae as other groups of the Ericales. Based on the results of this initial sister to a clade with Clethraceae, Cyrillaceae, and Eri- analysis, a second analysis based on morphological data was caceae received 67% jackknife support, and the clade performed to investigate species relationships in Purdiaea. with Clethraceae, Cyrillaceae, and Ericaceae got 89% The molecular data from a combination of atpB, ndhF, and support. In the present analysis, the two species of Pur- rbcL sequences was analyzed with parsimony jackknifing (Farris diaea and the two Clethra species formed a monophylet- et al. 1996) using the computer software “Xac” (Farris 1997, ic group with 88% support, and on the next higher node Källersjö et al. 1998) with the following settings: 1000 replica- tions, each with branch-swapping and ten random addition se- Cyrilla and Cliftonia grouped together (100% support) quences.
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