Open Access
Translator Disclaimer
20 December 2011 A molecular phylogenetic study of the Ephedra distachya / E. sinica complex in Eurasia
Nobuko Kakiuchi, Masayuki Mikage, Stefanie Ickert-Bond, Maria Maier-Stolte, Helmut Freitag
Author Affiliations +

This study provides new information on phylogenetic relationships in the Ephedra distachya / E. sinica complex (Ephedraceae, gymnosperms) based on sequence variation and morphology-based delimitation of individual species. Altogether we have included 50 samples from this complex and closely related species with 39 samples sequenced for the first time. Our sampling scheme provides a much broader sampling both in respect of the number of species as well as the number of samples per species than has been accomplished in previous studies. Sequences for E. dahurica and E. pseudodistachya are given here for the first time. We sequenced the nuclear ribosomal internal transcribed spacer regions (nr ITS1 ITS2) as well as the chloroplast intergenic spacer between rnL and trnF (cp trnL-F), but only ITS1 was sufficiently informative for phylogeny reconstruction. Our data show, for the first time, (1) a well supported E. strobilacea / E. sarcocarpa / E. transitoria clade as sister to the “L” or “Asia 2” clade, furthermore (2) a distinct E. distachya clade, (3) a composite E. dahurica clade that also includes E. sinica and a fraction of E. intermedia, (4) a well supported E. regeliana clade and (5) an unresolved agglomeration of E. intermedia, E. przewalskii, E. lomatolepis and E. pseudodistachya. Most noteworthy is the support derived from the molecular data for (1) the separation of the hitherto doubtful E. pseudodistachya from E. distachya and E. dahurica and (2) the high degree of sequence similarity of E. sinica and E. dahurica. Based on morphological, molecular, ecological and biogeographical evidence, E. sinica is reduced to a subspecies of E. dahurica.

See the PDF.



J. Amann 1739: Stirpium rariorum in imperio Rutheno sponte provenientium icones et descriptiones collectae 1. — Petropoli: Typographia Academiae Scientiarum. Google Scholar


M. N. Chaudhri , I. H. Vegter & C. M. de Wal 1972: Index herbariorum. Part 2, Collectors 3. — Regnum Veg. 86. Google Scholar


C. Y. Cheng 1978: Ephedraceae. — Pp. 468–489 in: W. C. Cheng & L. K. Fu (ed.), Flora Reipublicae Popularis Sinicae 7. — Beijing: Science Press. Google Scholar


Chinese Pharmacopoeia Commission (ed.) 2005: Pharmacopoeia of the People’s Republic of China 1. — Beijing: People's Medical Publ. House. Google Scholar


S. K. Czerepanov 1995: Vascular plants of Russia and adjacent states (the former USSR). — Cambridge: Cambridge University. Google Scholar


C. R. Florin 1933: Über einige neue oder wenig bekannte asiatische Ephedra-Arten der Sektion Pseudobaccatae Stapf. — Kongl. Svenska Vetenskapsakad. Handl., ser. 3, 12(1): 1–49. Google Scholar


H. Freitag & M. Maier-Stolte 1993: Ephedra. — P. 49 in: T. G. Tutin , N. A. Burges , A. O. Chater , J. R. Edmondson , V. H. Heywood , D. M. Moore , D. H. Valentine , S. M. Walters & D. A. Webb (ed.), Flora europaea, ed. 2, 1. — Cambridge: Cambridge University. Google Scholar


H. Freitag & M. Maier-Stolte 1994: Ephedraceae. — Pp. 5–16, 39–52 in: K. Browicz (ed), Chorology of trees and shrubs in Southwest Asia and adjacent regions 10. — Poznan: Polish Scientific Publishers. Google Scholar


H. Freitag & M. Maier-Stolte 2009: Ephedra. — In: FloraGreif. Virtual flora of Mongolia. — Greifswald University: Published at floragreif  Google Scholar


B. Frenzel 1992: Atlas of the palaeoclimates and palaeoenvironments of the northern hemisphere. Late Pleistocene—Holocene. — Budapest: Hungarian Academy of Sciences; Jena: G. Fischer. Google Scholar


L. G. Fu , Y. F. Yu & H. Riedl 1999: Ephedraceae. — Pp. 97–103 in: Z. Y. Wu & P. H. Raven (ed.). Flora of China 4. — Beijing: Science Press & St Louis: Missouri Botanical Garden. Google Scholar


A. V. Galanin 2008: Ephedra. — Pp. 74 in: A. V. Galanin , A. B. Belikovich & O. B. Khrbko (ed.), Flora of Dahuria 1. — Vladivostok: Dal'nauka. Google Scholar


J. G. Gmelin 1747: Flora sibirica sive historia plantarum Siberiae 1. — St. Petersburg: Academia Scientiarum. Google Scholar


R. Govaerts 2010a: World Checklist of Ephedraceae. — Royal Botanic Gardens, Kew: published at  http:// Scholar


R. Govaerts 2010b: Ephedraceae. — In: Euro+Med PlantBase. — Published at Scholar


V. I. Grubov 1982: Opredelitel' sosudistykh rastenii Mongolii — Leningrad: Nauka [Engl, ed.: 2000, Key to the vascular plants of Mongolia. Enfield & Plymouth: Science Publishers]. Google Scholar


I. A. Gubanov 1996: Konspekt flory vneshnej Mongolii (sosudistye rasteniya) — Moskva: Valang. Google Scholar


J. Huang , D. E. Giannasi & R. A. Price 2005: Phylogenetic relationships in Ephedra (Ephedraceae) inferred from chloroplast and nuclear DNA sequences. — Molec. Phylogen. Evol. 35: 48–59. Google Scholar


S. M. Ickert-Bond & C. Rydin 2011: Micromorphology of the seed envelope of Ephedra (Gnetales) and its relevance for the timing of evolutionary events. — Int. J. Pl. Sci. 172: 36–48. Google Scholar


S. M. Ickert-Bond & M. F. Wojciechowski 2004: Phylogenetic relationship in Ephedra ( Gnetales): evidence from nuclear and chloroplast DNA sequence data. — Syst. Bot. 29: 834–849. Google Scholar


S. M. Ickert-Bond , C. Rydin & S. S. Renner 2009: A fossil-calibrated relaxed clock for Ephedra indicates an Oligocene age for the divergence of Asian and New World clades and Miocene dispersal into South America. — J. Syst. Evol. 47: 444–456. Google Scholar


Japanese Ministry of Health, Labour and Welfare 2006: Japanese Pharmacopoeia, ed. 15. — Published at  http://  Google Scholar


C. Jarvis 2007: Order out of chaos: Linnean plant names and their types. — London: The Linnean Society & The Natural History Museum. Google Scholar


N. Kakiuchi & M. Mikage 2007: Collaborative survey of Ephedra plants in Buryatia, Russia. — Newslett. Himalayan Bot. 39: 1–8. Google Scholar


G. Lang 1994: Quartäre Vegetationsgeschichte Europas: Methoden und Ergebnisse. — Stuttgart: G. Fischer. Google Scholar


Y. Liu 1985: Flora in desertis Reipublicae Populorum Sinarum 1. — Beijing: Science Press [in Chinese]. Google Scholar


C. F. Long , N. Kakiuchi , A. Takahashi , K. Komatsu , S. Q. Cai & M. Mikage 2004: Phylogenetic analysis of the DNA sequence of the non-coding region of nuclear ribosomal DNA and chloroplast of Ephedra plants in China. — Pl. Med. 70: 1080–1085. Google Scholar


D. R. Maddison & W. P. Maddison 2005: MacClade v. 4.08. — Sunderland, Mass.: Sinauer Associates. Google Scholar


C. A. Meyer 1846: Versuch einer Monographie der Gattung Ephedra — St. Petersburg: Druckerei der Kaiserlichen Akademie der Wissenschaften. Google Scholar


M. Mikage 2009: Report of achievement for Grant-in-Aids for Project No. 17405014 from the Ministry of Education, Culture, Sport Science and Technology of Japan. — Published at ja/p/17405014  Google Scholar


K. Müller 2006: Incorporating information from lengthmutational events into phylogenetic analysis. — Molec. Phylogen. Evol. 38: 667–676. Google Scholar


V. A. Nikitin 1957: Ephedraceae — Pp. 503–504 in: P. N. Ovchinnikow (ed.). Flora Tadzhikskoi SSR 1. — Moskva & Leningrad: Izd. Akedamii Nauk SSSR. Google Scholar


J. Nouviant 1993: Ephedra negrii. — Pp. 65 in: M. Kerguelen , Index synonymique de la Flore de France. — Paris: Muséum National d'Histoire Naturelle. Google Scholar


J. Nouviant 1997: Recherches sur Ephedra en Europe III: Entre Ephedra distachya et helvetica. — Bull. Murith. Soc. Valais, Sci. Nat. 115: 68–75. Google Scholar


Nouviant J:1998: Recherches sur Ephedra in Europe V: Typification de E. monostachya et clé de détermination des Ephedra européens. — Bull. Murith. Soc. Valais, Sci. Nat. 116: 81–87. Google Scholar


M. G. Pachomova 1968: Ephedra vvedenskyi Pachom. sp. nov. —Pp. 198–199 in: S. S. Kovalevskaya (ed.), Conspectus florae Asiae Mediae 1. — Tashkent: Academia Scientiarum USSR. Google Scholar


M. G. Pachomova 1971: Ephedra. — Pp. 25–33 in: V. I. Grubov , A. E. Macenko & M. G. Pakhomova (ed.), Plantae Asiae Centralis 6. — Leningrad: Nauka. Google Scholar


G. A. Peschkova 2005: Synopsis of the Siberian Ephedra species (Ephedraceae). — Bot. Zhurn. 90: 423–426. [in Russian]. Google Scholar


W. Pfeiffer & A. Stamatakis 2010: Hybrid MPI/Pthreads parallelization of the RAxML Phylogenetics Code. — In: Ninth IEEE International Workshop on High Performance Computational Biology (HiCOMB 2010) Atlanta. — Published on the CIPRES Science Gateway at  http:  Google Scholar


E. Regel 1880: Ephedra glauca Rgl. — Trudy Imp. St.Peterburgsk. Bot. Sada 6: 484. Google Scholar


R. Riedl 1967: Die Gattung Ephedra in Europa. — Sci. Pharm. 36: 225–228. Google Scholar


F. Ronquist & J. P. Huelsenbeck 2003: MrBayes 3.0: Bayesian phylogenetic inference using mixed models. — Bioinformatics 19: 1572–1574. Google Scholar


C. Rydin & P. Korall 2009: Evolutionary relationships in Ephedra (Gnetales), with implications for seed plant phylogeny. — Int. J. Pl. Sci. 170: 1031–1043. Google Scholar


C. Rydin , A. Khodabandeh & P. K. Endress 2010: The female reproductive unit of Ephedra (Gnetales): comparative morphology and evolutionary perspectives. — Bot. J. Linn. Soc. 163: 387–430. Google Scholar


P. Sharma & P. L. Uniyal 2009: Ephedra sumlingensis (Ephedraceae) Sharma & Uniyal a new species from Himachal Pradesh, India. — Bull. Bot. Surv. India 50: 179–182. Google Scholar


P. Sharma , P. L. Uniyal & O. Hammer 2010: Two new species of Ephedra (Ephedraceae) from the western Himalayan Region. —  Syst. Bot. 35: 730–735. Google Scholar


M. P. Simmons & H. Ochoterena 2000: Gaps as characters in sequence-based phylogenetic analyses. — Syst. Biol. 49: 369–381. Google Scholar


M. P. Simmons , K. Müller & A. P. Norton 2007: The relative performance of indel coding methods in simulations. — Molec. Phylogen. Evol. 44: 724–740 Google Scholar


V. N. Siplivinskiy 1973: Chto takoe Ephedra dahurica Turcz.? [What is Ephedra dahurica Turcz.?]. — Novosti Sist. Vyssh. Rast. 1973: 355–357. Google Scholar


A. Stamatakis , P. Hoover & J. Rougemont 2008: A fast bootstrapping algorithm for the RAxML Web-Servers. — Syst. Biol. 57: 758–771. Google Scholar


O. Stapf 1889: Die Arten der Gattung Ephedra. —  Denkschr. Kaiserl. Akad. Wiss., Wien, Math.-Naturwiss. Kl. 56: 1–112. Google Scholar


O. Stapf 1927: Ma huang of China (Epheda sinica Stapf). — Bull. Misc. Inf. Kew. 3: 133–134. Google Scholar


P. Taberlet , F. Gielly , G. Pautou & J. Boubet 1991: Universal primers for amplification of three non-coding region of chloroplast DNA. — Pl. Molec. Biol. 17: 1105–1109. Google Scholar


A. L. Takhtajan & M. G. Pachomova 1967: Ephedra aurantiaca Takht. & Pachom. sp. n. — Bot. Mater. Gerb. Inst. Bot. Akad. Nauk Uzbeksk. S.S.R. 18: 53. Google Scholar


S. W. Tang [ed. by Shang Z. J., based on the 12 century edition by Cheng A.] 2002: Daguan Bencao. — Hefei: Anhui Academic Publisher [in Chinese]. Google Scholar


B. Thiers 2008+ [continuously updated]: Index herbariorum: A global directory of public herbaria and associated staff. — New York Botanical Garden: published at Scholar


N. S. Turczaninow 1854: Flora baicalensi-dahurica [17]. — Bull. Soc. Imp. Naturalistes Moscou 27: 353–422. Google Scholar


L. Wang , N. Kakiuchi & M. Mikage 2010: Studies on Ephedra plants in Asia 6. Geographical changes of anatomical features and alkaloid content of Ephedra sinica. — J. Nat. Med. 64: 63–69. Google Scholar


C. Y. Yang 1993: Ephedraceae. — Pp. 87–109 in: C. Y. Yang (ed.). Flora xinjiangensis 1. — Wulumuqi: Xinjiang Science, Technology and Hygiene Publishing. Google Scholar


Y. Yang 2002: Classification of Chinese Ephedra plants and discussion on their difference. — Doctoral thesis. Graduate School of the Chinese Academy of Science. [in Chinese]. Google Scholar


Y. Yang , D. Fu & G. Zhu 2003: A new species of Ephedra (Ephedraceae) from China. —  Novon 13: 153–155. Google Scholar
© 2011 BGBM Berlin-Dahlem.
Nobuko Kakiuchi, Masayuki Mikage, Stefanie Ickert-Bond, Maria Maier-Stolte, and Helmut Freitag "A molecular phylogenetic study of the Ephedra distachya / E. sinica complex in Eurasia," Willdenowia 41(2), 203-215, (20 December 2011).
Published: 20 December 2011

cp trnL-F
Ephedra dahurica
Ephedra intermedia
Ephedra pseudodistachya
Ephedra strobilacea
Get copyright permission
Back to Top