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Merging Fargesia Dracocephala Into Fargesia Decurvata (Bambusoideae, Poaceae): Implications from Morphological and ITS Sequence Analyses

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Merging Fargesia Dracocephala Into Fargesia Decurvata (Bambusoideae, Poaceae): Implications from Morphological and ITS Sequence Analyses Merging Fargesia dracocephala into Fargesia decurvata (Bambusoideae, Poaceae): Implications from Morphological and ITS Sequence Analyses Yu-Qu Zhang1., Xu-Mei Wang2., A-Li Wu3, Yi Ren1* 1 College of Life Sciences, Shaanxi Normal University, Xi’an, China, 2 School of Pharmacy, Xi’an Jiaotong University, Xi’an, China, 3 Shaanxi Changqing National Nature Reserve, Huayang, China Abstract Aims: Fargesia decurvata is closely allied with F. dracocephala and differs in 5 major characters (i.e. the culm sheath blade base shape, the width of the culm sheath blade base, the auricle shape, and the lower surface of leaf blade) in Fargesia.Itis difficult to distinguish these two species because of existing of transitional statements of characters. The aims of this paper are to (i) investigate whether the variation of the characters is continuous or not; (ii) reveal whether the publishment of F. dracocephala was the result of discontinuous sampling of F. decurvata or not. Methods: Ten populations of F. decurvata and F. dracocephala were investigated in their entire distribution (including type localities). The statements of 5 major characters were measured from 693 annual and 693 perennial culms of 231 individuals in 10 populations, and analyzed at population, individual and culm levels. UPGMA cluster analysis was carried out based on 29 characters from 10 populations of F. decurvata and F. dracocephala and 2 populations of F. qinlingensis as outgroup. The ITS sequences were also sequenced and analyzed. Important Findings: Five major characters exhibited great variation not only at population level, but at individual level within a population, even the culm level within an individual and in different parts of the same culm. Cluster analyses showed that 10 populations of F. decurvata and F. dracocephala were not divided into two species, but they were well separated with outgroup. There was no difference in floral organ between F. decurvata and F. dracocephala. MP and NJ trees based on ITS sequences showed the same results with the cluster analysis on morphological characters. All the facts indicated that the publishment of F. dracocephala was the result of discontinuous sampling of F. decurvata, and F. dracocephala should be treated as the synonym of F. decurvata. Citation: Zhang Y-Q, Wang X-M, Wu A-L, Ren Y (2014) Merging Fargesia dracocephala into Fargesia decurvata (Bambusoideae, Poaceae): Implications from Morphological and ITS Sequence Analyses. PLoS ONE 9(7): e101362. doi:10.1371/journal.pone.0101362 Editor: Shuang-Quan Huang, Central China Normal University, China Received January 28, 2014; Accepted June 5, 2014; Published July 2, 2014 Copyright: ß 2014 Zhang et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Funding: Funding came from the Giant Panda International Cooperation Project of the State Forestry Administration of China (WH0630). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Competing Interests: The authors have declared that no competing interests exist. * Email: [email protected] . These authors contributed equally to this work. Introduction vegetative characters [5,6]. Each of the bamboo is distributed in a wide range and grows in diverse habitat, but the specimens can be The correct definition of species is the most basic work for collected in limited locations and habitat, many vegetative further researches. The type and other voucher specimens play an variations might be ignored due to the limited sampling. In important role in the process of defining a species. However, additional, only a small part of the culms, branches or rhizomes because the herbarium specimens are often collected from the can be collected as the specimens. As a result, the specimen-based limited location(s) and/or limited individual(s) or sometimes just description and definition of bamboo species would inevitably from parts of an individual, the continuity of variation of cause the continuous variations of the characters to be fragmented. character(s) within a species would be artificially isolated. The Fargesia is a large genus in Bambusoideae (Poaceae) with about artificial isolation of character(s) might lead to the result that a 90 species distributed mainly in the alpine areas at altitude of virtual species will be defined as different ones [1–4]. 1400–3800 m in south-west China, Vietnam and adjacent The problems are particularly prominent in the definition of Himalayas [7,8]. There are 78 species in China and 61 of them most of bamboo species. Because the bamboos typically have to go were nominated and published without the description of through over decades or even more than a century of vegetative reproductive organs and many of them are narrow distribution growth before flowering, and generally died off within a few years species [6–8]. In the field survey of Fargesia species as the main after flowering, thus the chance to get the flowers and caryopses is very little when the specimens were collected, the defining and food of giant panda, we found that some bamboo samples from identifying of the bamboo species have been mainly depended on this genus could not be accurately identified because of the existing PLOS ONE | www.plosone.org 1 July 2014 | Volume 9 | Issue 7 | e101362 Taxonomy of Fargesia decurvata and F. dracocephala of transitional or intermediate characters between or among the base. There is no description of the reproductive organs. species. Among these species, F. decurvata Lu (Fig. 1) and F. Thereafter Yi published F. aurita, which is based on specimens dracocephala Yi are a pair of species which have wider distribution collected from Fengdu County, Sichuan Province (Yi Tongpei and are most difficult to identify. Therefore, we considered that 75410, Type in Herb. Forestry School of Sichuan Province) [10], these two species might be the good materials to study the but the name was treated as a synonym of F. decurvata by Yi himself definition of bamboo species. [7]. Fargesia decurvata was described by Lu [9] based on specimen Fargesia dracocephala was described by Yi based on specimens collected from Hejiaping, Changyang County, Hubei Province (Lu collected from Guangwu Mountain, Nanjiang County, Sichuan Jionglin 78120, Type in Herb. Honan Col. Agr. and PE). He Province (Yi Tongpei 75540, Type in Herb. Forestry School of considered that this species differs from other Fargesia species in Sichuan Province), Qianjiaping Forest Farm, Pingli County, following aspects: the base of the sheath blade is as wide as the Shaanxi Province and Shennongjia, Hubei Province [11]. He sheath tongue or nearly so, the culm sheath tongue is promontory considered that this species differs from F. robusta in following as a bow and strongly decurrently on both sides, the culm sheath aspects: the internodes are nearly solid and the hollows are 1– has dense gray small bristles on the back and dense coarse wools at 2.7 mm in diameter, the culm sheath is nearly hairless or with Figure 1. Morphology of Fargesia decurvata. A. Bamboo clump. Bar = 30 cm. B. Middle parts of a bamboo shoot. Bar = 1 cm. C. Branch and leaves. Bar = 0.8 cm. D. Yong infructescence. Bar = 1.5 cm. E. Close up of a node. Bar = 1 cm. doi:10.1371/journal.pone.0101362.g001 PLOS ONE | www.plosone.org 2 July 2014 | Volume 9 | Issue 7 | e101362 Taxonomy of Fargesia decurvata and F. dracocephala Figure 2. A–C. Close up of upper part of sheathes and lower part of sheath blades in different of a same annual culm in Pop 4. Bar = 5 mm. A. In lower part, showing blade base decurrent (arrow) and as wide as sheath apex. B. In middle part, showing blade base not decurrent (arrow) and narrower than sheath apex. C. In upper part, showing blade base not decurrent (arrow) and much narrower than sheath apex. D–L. Close up of leaf auricles and oral setae, showing the shape of leaf auricles and position of oral serae. Bar = 1 mm. D–F. From Pop 1. D–E. From an annual culm. D. Long triangular auricle with serae at upper margins and apex. E. Curving long triangular auricle with setae at margins and apex. F. Short sickle-like with setae mostly at one margin and apex from a perennial culm. G–H. From Pop 3. G. Sickle-like auricle with setae mostly at one margin and apex from an annual culm. H. Very narrow sickle-like auricle with setae mostly at one margin and apex from a perennial clam. I–J. Culms from Pop PLOS ONE | www.plosone.org 3 July 2014 | Volume 9 | Issue 7 | e101362 Taxonomy of Fargesia decurvata and F. dracocephala 8. I. Banding auricle with setae at upper margins and apex from an annual culm. J. Short banding auricle with setae at upper margins and apex from a perennial culm. K–L. From Pop 10. K. very small auricle with setae at whole margin from an annual culm. L. Almost without auricle but with setae on upper margin of sheath from a perennial culm. C: culm. CB: culm sheath blade. CS: culm sheath. LA: leaf auricle. LS: leaf sheath. S: oral seta. doi:10.1371/journal.pone.0101362.g002 sallow thin bristles on the back, the leaf sheath is elliptic and with described for the first time [11], the auricles were described as setae at apex, the leaf blade has 3–4 pairs of secondary veins. He ‘‘auriculis foliorum ellipticis’’ (leaf auricle elliptical) and no described the morphology of flowers later [12]. description of the seta position, but it seems that the auricles are Fargesia decurvata and F.
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