Abstract
The circumscription and phylogenetic relationships of the Anemone section Begoniifolia have long been in dispute. Here, we use molecular data and palynological and cytological observations to test the monophyly of the section and evaluate phylogenetic relationships within the section. Three pollen types were observed in the section: Anemone sumatrana is 3-colpate; A. hokouensis, A. howellii, and A. orthocarpa are 6-colpate; and A. begoniifolia and A. xingyiensis are pantoporate. All six species have the chromosome number 2n = 16 and essentially uniform karyotypes (2n = 10 m + 4st + 2t). Nuclear ITS and plastid atpB-rbcL DNA sequences were analyzed using maximum parsimony, maximum likelihood, and Bayesian inference methods. Within Anemone s. l., two strongly supported clades corresponding to subgenera Anemonidium (x = 7) and Anemone (x = 8) were recovered. Our results indicate that section Begoniifolia is better treated at subsectional rank and is one of four subsections (Begoniifoliae, Rupicolae, Rivulares, and Vitifoliae) within section Rivularidium of subgenus Anemone. Subsection Begoniifoliae is relatively well supported and includes A. begoniifolia, A. hokouensis, A. howellii, A. orthocarpa, A. sumatrana, and A. xingyiensis. We present a formal reclassification of section Rivularidum, including two new combinations: subsections Begoniifoliae and Rivulares.
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Acknowledgments
We thank Prof. Y. F. Deng, Dr. J. Cai, Dr. Tran Van Tien, and Mr. J. P. Luo for their help in collecting plant material. This work was supported by the Main Direction Program of Knowledge Innovation of the Chinese Academy of Sciences (KSCX2-EW-Z-1).
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Appendices
Appendix 1
Voucher information of cytological and palynological material of Anemone section Begoniifolia. The sample marked with one asterisk was only palynologically examined, and those marked with two asterisks were both palynologically and cytologically examined.
Anemone begoniifolia H. Lév. and Vaniot, Duyun, Guizhou, China, L. Y. Wang 5*(IBSC); Xingyi, Guizhou, China, Y. Zhang and M. Tang 75 (IBSC); Nanchuan, Chongqing, China, Y. Zhang 91 (IBSC). A. hokouensis C. Y. Wu and W. T. Wang, Hekou, Yunnan, China, Y. Zhang and M. Tang 89**(IBSC). A. howellii J. F. Jeffrey and W. W. Smith, Anlong, Guizhou, China, Y. Zhang and M. Tang 87**(IBSC); Libo, Guizhou, China, Y. F. Deng s.n. (IBSC); Libo, Guizhou, China, M. Tang and J. P. Luo 24 (IBSC). A. orthocarpa Hand.-Mazz., Xingyi, Guizhou, China, Y. Zhang and M. Tang 76**(IBSC). A. sumatrana de Vriese, Lac Duong, Lam Dong, Vietnam, Tran Van Tien s.n. **(IBSC).
Appendix 2
List of taxa and sources of plant material used for the current molecular study [taxa, GenBank accession numbers (ITS, and atpB-rbcL), source, and voucher data]. * indicates taxa sequenced for this study; all other sequences are referenced in Hoot et al. (2012).
*Anemoclema glaucifolium (Franch.) W. T. Wang, KJ819801, KJ819821, Zhongdian, Yunnan, China, Q. E. Yang 3104 (IBSC); KJ819802, KJ819822, Lijiang, Yunnan, China, Q. E. Yang 2649 (IBSC). Anemone americana (DC.) H. Hara, AY055386, AY055407. A. anemonoides (H. Rasm.) J. C. Manning and Goldblatt, JF810691, JF810701. A. antucensis Poepp., AY056049, AF311735. (2012). A. balliana (Britton) Hoot, FJ639900, FJ639864. *A. begoniifolia H. Lév. and Vaniot, KJ819810, KJ819830, Nanchuan, Chongqing, China, Y. Zhang 91 (IBSC). A. berlandieri Pritz., FJ639876, FJ639841. A. blanda Schott and Kotschy, AY055402, AY055422. A. bracteata (Harv. ex J. Zahlbr.) J. C. Manning and Goldblatt, JF810692, JF810702. A. caffra Harv., AY055399, AY055420. A. canadensis L., AY055387, AY055408. A. caroliniana Walt., AY055403, AY055423. A. chilensis (Gay) Reiche, FJ639901, FJ639865. A. coronaria L., FJ639877, FJ639842. A. crassifolia Hook. f., AY055398, AY055419. *A. davidii Franch., KJ819811, KJ819831, Emei Shan, Sichuan, China, Y. Zhang 93 (IBSC). A. decapetala Ard., FJ639878, FJ639843. A. demissa Hook. f. and Thoms., FJ639879, FJ639844. *A. demissa Hook. f. and Thoms., KJ819799, KJ819819, Luquan, Yunnan, China, J. Cai et al. 13PXB025 (IBSC). A. drummondii S. Watson, AY055404, AY055424. A. edwardsiana Tharp, FJ639880, FJ639845. A. fanninii Harv., JF810685, JF810695. A. flaccida F. Schmidt, AY055391, AY055412. *A. flaccida F. Schmidt, KJ819809, KJ819829, Emei Shan, Sichuan, China, Y. Zhang 90, (IBSC). *A. griffithii Hook. f. and Thoms., KJ819798, KJ819818, Medog, Xizang, China, J. Cai et al. 13CS7679 (IBSC). A. halleri All., FJ639908, FJ639872. A. helleborifolia DC., FJ639881, FJ639846. A. hepatica L., FJ639904, FJ639868. A. hortensis L., FJ639882, FJ639847. *A. hokouensis C. Y. Wu and W. T. Wang, KJ819808, KJ819828, Hekou, Yunnan, China, Y. Zhang and M. Tang 89 (IBSC). *A. howellii J. F. Jeffrey and W. W. Smith, KJ819805, KJ819825, Libo, Guizhou, China, M. Tang and J. P. Luo24 (IBSC). A. hupehensis Lemoine, AY055397, AY055418. *A. imbricata Maxim., KJ819793, KJ819813, Zayu, Xizang, China, J. Cai et al. 13CS6946 (IBSC). A. integrifolia Humb. and Bonpl. ex DC., FJ639906, FJ639870. A. integrifolia Humb. and Bonpl. ex DC., FJ639907, FJ639871. A. keiskeana T. Ito ex Maxim., AY055390, AY055411. A. knowltonia Burtt Davy, JF810693, JF810703. A. lithophila Rydb., FJ639883, FJ639848. A. major (Phil.) Meigen, FJ639902, FJ639866. A. mexicana Kunth FJ639884, FJ639849. A. narcissiflora L., AY055393, AY055414. A. nemorosa L., FJ639885, AF386091, Meyer et al. (2010); Ehrendorfer and Samuel (2001). *A. obtusiloba D. Don, KJ819797, KJ819817, Markam, Xizang, China, J. Cai et al. 13CS7032 (IBSC). *A. obtusiloba D. Don, KJ819795, KJ819815, Zayu, Xizang, China, J. Cai et al. 13CS6953 (IBSC). A. occidentalis S. Watson, AY055400, AY055426. *A. orthocarpa Hand.-Mazz., KJ819812, KJ819832, Xingyi, Guizhou, China, Y. Zhang and M. Tang 76 (IBSC.) A. parviflora Michx., FJ639887, FJ639851. A. pavonina Lam., FJ639888, FJ639852. A. quinquefolia L., JF810686, JF810696. A. ranunculoides L., FJ639889, FJ639853. A. richardsonii Hook. f., AY055388, AY055409. A. rigida Gay, FJ639890, FJ639854, Meyer et al. (2010). A. rivularis Buch.-Ham. ex DC., AY055396, AY055417. *A. rivularis Buch.-Ham. ex DC., KJ819807, KJ819827, Anlong, Guizhou, China, Y. Zhang and M. Tang 88 (IBSC). *A. rupicola Camb., KJ819796, KJ819816, Medog, Xizang, China, J. Cai et al. 13CS7010 (IBSC). A. somaliensis Hepper, JF810687, JF810697. * A. sumatrana de Vriese, KJ819804, KJ819824, Lac Duong, Lam Dong, Vietnam, Tran Van Tien s.n. (IBSC). A. sylvestris L., FJ639894, FJ639858 Hoot et al. (2012). *A. sylvestris L., KJ819803, KJ819823, Chifeng, Nei Mongol, China, Y. Liu 178 (IBSC). A. tenuicaulis (Cheeseman) Parkin and Sledge, AY055389, AY055410. A. tenuifolia var. capensis, JF810688, JF810698. A. tenuifolia var. tenuifolia, JF810689, JF810699. A. tetrasepala Royle, FJ639895, FJ639859. A. thomsonii Oliv., JF810690, JF810700. *A. tomentosa (Maxim.) Péi, KJ819800, KJ819820, Lushi, Henan, China, J. P. Luo and L. Wang 152 (IBSC). A. transsilvanica (Fuss) Heuff, FJ639905, FJ639869. A. trullifolia Hook. f. and Thoms., AY055395, AY055416. *A. trullifolia var. linearis (Brühl) Hand.-Mazz., KJ819794, KJ819814, Medog, Xizang, China, J. Cai et al. 13CS6952 (IBSC). A. tuberosa Rydb., FJ639897, FJ639861. A. vesicatoria (L. f.) Prantl, AY055401, AY055421. A. virginiana L., FJ639898, FJ639862. A. vitifolia Buch.-Ham. ex DC., FJ639899, FJ639863. *A. xingyiensis Q. Yuan and Q. E. Yang, KJ819806, KJ819826, Xingyi, Guizhou, China, Y. Zhang and M. Tang 77 (IBSC). Clematis hexapetala Pall., AY055385, AY055406. C. ochroleuca Aiton, FJ639903, FJ639867.
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Zhang, Y., Hong, Y., Ren, C. et al. Palynology, cytology, and molecular systematics of Anemone section Begoniifolia (Ranunculaceae). Plant Syst Evol 301, 411–424 (2015). https://doi.org/10.1007/s00606-014-1082-0
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DOI: https://doi.org/10.1007/s00606-014-1082-0