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A – C: Ephedra distachya subsp. helvetica – A: with mature female cones; Suisse, Valais, Tourbillon hill near Sion, on S exposed rocky outcrops (with Sedum album ); phot. H. Freitag, August 1990; B: shoot with male cones; Italy, S Tyrol, Vinschgau, S-exposed slopes above Schlanders; phot. H. Freitag, July 1990; C: shoot with immature female cones, the comparatively short micropylar tubes correspond to E. “negrii” ; France, Alpes de Hautes Provence, Larange-Montéglin; phot. E. Zippel, May 2005. – D: E. dahurica with mature female cones; the distorted twigs are artefacts due to unknown injury that also occur in other species; Russia, Buryatia, c. 100 km S Ulan-Ude, in Stipa krylovii steppe near Sulfat’noe Ozero; phot. H. Freitag, September 2003.
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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 ti...
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... Ephedra distachya L. / E. sinica Stapf complex is defined here somewhat arbitrarily as a group of morphologically similar and closely related species that additionally also include E. dahurica Turcz., E. lomatolepis Schrenk and E. pseudodistachya Pachom . Taxa in this complex share the same habit, range from 10 – 40 cm in size, bear 2-seeded female cones which become fleshy at maturity, generally have a rather short micropylar tube as well as paired leaves (Fig. 1). Species delimitation in this complex is difficult, particularly in the vegetative stage which dominates in the field and in the majority of herbarium specimens. The centre of distribution is the steppe and semidesert area of Eurasia, further extending from the Atlantic and Mediterranean to the eastern margin of the continent. Recent molecular work (Long & al. ...
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Understanding the factors that contribute to population genetic structures and ecological niche divergence is crucial for elucidating the biogeographical history and speciation of plants. To disentangle the present phylogeographical patterns and evolutionary history of two closely related parapatric species from Ephedra (Ephedra sinica and E. inter...
Citations
... Rbcl Huang et al., 2005Wang et al., 2005 12. rps4 Ickert-Bond and Wojciechowski 2004;Rydin et al., 2004 13. trn K Zhu et al., 2012 14. Trn L Kakiuchi et al., 2011;Long et al., 2004 15. ...
... Thus such characters had proven to be useful for monitoring genetic diversity in gene pools and to elucidate the factors controlling changes in life history of natural populations such as breeding system, inbreeding depression, effective population size, population structure and gene flows (Lynch and Milligan, 1994). In genus Ephedra, molecular taxonomy had helped in solving the taxonomic complexity and misidentification problems and reduction to synonyms (Kakiuchi et al., 2011;Long et al., 2004;Faried et al., 2018). In some cases, the very closely species had been reduced to a subspecies due to high degree of sequence similarity such as E. dahurica to E. sinica subsp. ...
... In some cases, the very closely species had been reduced to a subspecies due to high degree of sequence similarity such as E. dahurica to E. sinica subsp. dahurica (Kakiuchi et al., 2011) and recently E. dahurica had been reduced to E. sinica (Kakiuchi, 2017). In other cases, different species had been separated from species complex such as E. ...
The descriptions of dubious new species that lack robust taxonomic rigor create more confusion rather than bridging the Linnean shortfall in biodiversity. In an era of biodiversity crisis, it becomes urgent to undertake integrative taxonomic revisions to resolve taxonomic confusions in several plant taxa, even if that leads to drastic decline in the species number. Here I resolve the taxonomic conundrum of Ephedra in India by adopting an integrative taxonomic approach and using comprehensive set of characters from multiple lines of evidence (morphology, anatomy, palynology, seed micromorphology and molecular data). I reduce the number of Ephedra species in India from the currently known 16 to only 4 well-defined species: E. foliata, E. gerardiana, E. intermedia and E. regeliana. I provide proper species delimitations, detailed descriptions, taxonomic keys, photoplates of diagnostic characters, regional distributions and phylogenetic relationships of Ephedra in India, validated by robust empirical evidence. Our studies reveal that the previously reported three species: E. nebrodensis, E. pachyclada and E. przewalskii do not occur in India. The recently described four species (E. sumlingensis, E. pangiensis, E. khurikensis and E. yangthangensis) are synonymized with E. intermedia, and another E. kardangensis synonymized with E. gerardiana. Five recent designations viz. E. sheyensis, E. yurtungensis, E. yurtungensis var. lutea, E. lamayuruensis and E. khardongensis are recognized as nomen nodums due to lack of descriptions, diagnosis and type specimens. Our study provides a robust and reliable set of 16 morphological characters, validated by significant statistical support, which can prove useful for species delimitation in Ephedra. I also recorded few novel characters of evolutionary significance in Ephedra, which merit further investigation in future. Looking ahead, I believe that the methodological and data analytical learnings from this study can guide the future research direction in designing integrative taxonomic studies on such complex plant taxa elsewhere in the world.
... The cultivation of this plant was reported in few studies, but many fields contain other species of Ephedra. Genetic studies with ITS (internal transcribing sequence of nuclear ribosomal DNA) on Chinese Ephedra fields have shown that crops grown in six fields have been identified as Ephedra sinica and E. intermedia (Kakiuchi et al., 2011). In North African regions (native origin of this species), Louhaichi et al. (2011) mentioned the establishment of two field genebanks containing E. alata in arid zone of Libya. ...
Desert plants are a potentially sustainable source of bioactive molecules. This paper investigates phenolic profile, total polyphenols, flavonoids and condensed tannins in a collection of desert shrubs of Tunisian flora (Pituranthos tortuosus; Retama raetam; Ephedra alata; Ziziphus lotus; Capparis spinoza; Calligonum comosum). The collection was build according to people traditional tendency uses. The chemical composition was estimated in ethanol extracts of aerial parts together with a comparison of antioxidant activities. The results were significantly dependent on plant material source. Phenolic contents ranged from 15.52 to 37.5 mg of gallic acid equivalents (GAE)/g of Dry Weight (DW) of plant, and flavonoids ranged from 2.92 to 9.14 mg of rutin equivalents (RE)/g DW. The phenolic profile was composed of 23 compounds identified through LC-ESI-MS analysis. Five phenolic acids (quinic acid, gallic acid, syringic acid, p-coumaric acid, and trans-Ferulic acid) and five flavonoids (catechin, epicatechin, rutin, quercetrin and naringenin) were predominant and common between plants. Based on all data visualized within the heatmap, plants were clustered into two groups. The tow species Z. lotus and E. alata were selected for their highest antioxidant potential. These results allowed us to select the combination of three phenolic acids (rutin, qurectrin, quinic acid) that seems determinant in the strong antioxidant capacity. In addition, the possible domestication and cultivation of the two selected plants are discussed.
... It is partially because gross morphology does not reveal considerable interspecies variation, particularly in Ephedra as well in as some Gnetum lineages, e.g., Chinese lianoid Gnetum. In addition, intraspecific variation is also present in some species that possess a broad geographic distribution, e.g., E. distachya (Kakiuchi et al. 2011) and G. parvifolium (Huang et al. 2010), which further complicates taxonomic choices and assessments of species delimitation. In addition, vegetative parts of the Gnetales may exhibit plasticity as a response to the environment. ...
... A revision of the New World species is currently in preparation by the first author, and the Asian species present the greatest taxonomic challenge. New species of Ephedra continue to be described, mostly from India and China, although they appear morphologically close to E. intermedia and E. saxatilis (Yang et al., 2003;Yang, 2005;Sharma & Uniyal, 2009;Sharma et al., 2010;Sharma & Singh, 2015 (Freitag & Maier-Stolte, 1993, 1994Kakiuchi et al., 2011). Rydin et al., 2004;Huang et al., 2005;Rydin & Korall, 2009;Loera et al., 2015). ...
Ephedra, Gnetum, and Welwitschia constitute the gymnosperm order Gnetales of still unclear phylogenetic relationships within seed plants. Here we review progress over the past 10 years in our understanding of their species diversity, morphology, reproductive biology, chromosome numbers, and genome sizes, highlighting the unevenness in the sampling of species even for traits that can be studied in preserved material, such as pollen morphology. We include distribution maps and original illustrations of key features, and specify which species groups or geographic areas are under sampled.
... Previous phylogenetic studies on Ephedra have identifi ed low levels of molecular sequence divergence, suggesting a recent origin (Ickert-Bond and Wojciechowski 2004, Rydin et al. 2004, Huang et al. 2005, Wang et al. 2005, Rydin and Korall 2009, Kakiuchi et al. 2011, Qin et al. 2013. In this study, we sequenced additional molecular markers to estimate the divergence time of NW Ephedra species, generating an alignment of 7703 bp, improving resolution and providing a more suitable framework for the objectives proposed in this study. ...
In this study we selected the New World species of Ephedra to understand the ecological consequences of different dispersal syndromes. The twenty-three species of Ephedra in the New World have a disjunct distribution in North and South American arid and semi-arid habitats, exhibiting three dispersal syndromes related to dispersal by birds, wind and rodents. Using DNA sequence data we inferred phylogenetic relationships and lineage divergence times, and used these estimates to test different ecological assumptions. Using comparative methods we tested for correlations between dispersal syndromes and a set of ecological variables (niche breadth, niche evolution, distributional ranges and niche position). We found that speciation events in the New World coincided with the expansion of arid habitats in this region. We suggest that the bird dispersal syndrome is related with higher rates of climatic niche evolution for all variables used, including aridity index, mean annual temperature and mean annual precipitation. Distribution ranges were correlated with niche breadth, they were however not significantly different between dispersal syndromes. Species inhabiting the extremely arid regions on niche axes had narrower niche breadths. We conclude that species whose seeds are dispersed by birds have colonized a broader set of habitats and that those with wind and rodent dispersal syndromes might have promoted the colonization of more arid environments.
Anatomically modern humans are now believed to have arrived in Eurasia as much as 80,000- 120,000 years ago (e.g., see Callaway 2015), and in bands of hunters and gatherers these early people moved east and west progressively across this massive landmass. In the process of spreading out over this vast continental region, they developed very ancient and long-lasting paleoethnobotanical relationships with numerous species; the more significant of these prehistoric relationships in Eurasia included medicinally and psychoactively important genera such as Cannabis, Papaver, and Ephedra (for Cannabis, see chapter 1 in this volume; for
Papaver somniferum L., the opium poppy, see chapter 2). In the case of Ephedra, species in this genus appear to have been used very early on in some regions of both the Old and New World beginning many thousands of years ago.
Although Kampo medicine is now fully integrated into the modern Japanese healthcare system, most Kampo formulations depend on imported crude drugs from limited foreign areas. To prepare for possible shortages of crude drugs in the future, a wider scope for the supply of medicinal plants is necessary. We conducted field research and collaborated with international laboratories for phylogenic analysis and evaluation of medicinal plant resources. Our research on ephedra plants from a wide region of Eurasia has, for example, confirmed their phylogenic structure: based on DNA sequencing analysis of nuclear ribosomal internal transcribed spacer region 1 (ITS1) as well as the chloroplast intergenic spacer between trnL and trnF (trnL-F), the 8 major Chinese species and related plants grown on the continent could be divided into 3 groups. Additionally, Ephredra sinica was found to be synonymous with Ephredra dahurica and was reduced to a subspecies of Ephredra distachya. Furthermore, Ephredra likiangensis and Ephredra gerardiana, which are grouped in separate phylogenic trees, would be good candidates for medicinal material. Aconites from Hokkaido, as an example of domestic plants reviewed, were collected for phylogenic and aconitine alkaloid content analysis. The phylogenic analysis of nr ITSs revealed that the majority of specimens were genetically similar. However, the aconitine alkaloid content of the tuberous roots demonstrated that specimens from different habitats had varying alkaloid profiles. Environmental pressure of each habitat is presumed to have caused the morphology and aconitine alkaloid profiles of these genetically similar specimens to diversify.
Ephedrae Herba (ma-huang in Chinese), the herbal stems of Ephedra sinica Stapf, which belongs to the Ephedraceae family, is an important crude drug in traditional Chinese medicine. Some researchers think that this species is identical to E. distachya L., which mainly grows in Europe. Thus, in this study, we carried out anatomical, chemical, and molecular genetic studies on E. distachya plants collected in Switzerland, France, and Turkey to consider whether E. distachya could be used to produce Ephedrae Herba. As a result, we found that the morphology of the subepidermal fiber bundles in cross sections of the herbal stems of E. distachya and the frequencies of ephedrine alkaloids in E. distachya were quite different from those observed in E. sinica and that the DNA sequence of the ITS1 region of E. distachya contains 11 base changes compared with that of E. sinica. Based on these facts, we concluded that E. distachya and E. sinica should be treated as different taxa and that, according to the prescriptions of the current Japanese and Chinese pharmacopoeias, E. distachya is not suitable for producing Ephedrae Herba.
Classification of Ephedra has long been a matter of debate, as Ephedra plants have few morphological characteristics to aid classification. In particular, the distinction between E. equisetina and E. major subsp. procera is uncertain. Thus, in this study, an attempt was made to clarify their relationship by molecular analysis and content of ephedrine alkaloids. Molecular analysis revealed that E. equisetina and E. major subsp. procera have a close relationship with differing matrilineage, and chemical analysis showed that they have the same characteristic of varied alkaloid composition ratio by collection site. We suggest that on the basis of molecular phylogeny that E. major subsp. procera should be treated as a synonym or subspecies of E. equisetina instead of a subspecies of E. major.
