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Taxonomic notes on Chinese Lilium L. (Liliaceae) with proposal of three nomenclatural revisions

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In this revision, we treat Lilium wenshanense L.J. Peng & F.X. Li, L. jinfushanense L.J. Peng & B.N. Wang and L. huidongense J.M. Xu from China as synonyms of L. brownii F.E. Brown ex Miellez, L. taliense Franchet and L. lijiangense L.J. Peng, respectively. We justify the synonymy by demonstrating the existence of continuous morphological variation, which we observed in the field and among herbarium specimens.
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Phytotaxa 172 (2): 101–108
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Copyright © 2014 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Jinshuang Ma: 30 Apr. 2014; published: 13 Jun. 2014
http://dx.doi.org/10.11646/phytotaxa.172.2.5
101
Taxonomic notes on Chinese Lilium L. (Liliaceae) with proposal of three
nomenclatural revisions
YUN-DONG GAO & XIN-FEN GAO*
Key Laboratory of Mountain Ecological Restoration and Bioresource Utilization & Ecological Restoration Biodiversity Conservation
Key Laboratory of Sichuan Province, Chengdu Institute of Biology, Chinese Academy of Sciences; email: xfgao@cib.ac.cn
*Author for corresponding
Abstract
In this revision, we treat Lilium wenshanense L.J. Peng & F.X. Li, L. jinfushanense L.J. Peng & B.N. Wang and L. huidongense
J.M. Xu from China as synonyms of L. brownii F.E. Brown ex Miellez, L. taliense Franchet and L. lijiangense L.J. Peng,
respectively. We justify the synonymy by demonstrating the existence of continuous morphological variation, which we
observed in the field and among herbarium specimens.
Key words: Hengduan Mountains, Lilium, taxonomic revision, synonym
Introduction
The genus Lilium Linnaeus (1753: 302) exhibits great diversity in China, especially in the Hengduan Mountains Region
(HDM hereafter) (Haw 1986; Gao et al. 2013a). Specifically, ca. 30 species of Lilium are distributed in southwestern
China, including the HDM, according to the Flora of China (Liang and Tamura 2000). Recent research (Gao et al.
2012, 2013a, b) shows that Lilium also includes ca. 8 species formerly referred to Nomocharis Franchet (1889: 113),
which is largely endemic to southwestern China, especially the HDM.
Lilium is typical of uncertain taxonomic resolution within HDM genera. On the one hand, two new species of
Lilium native to the HDM have been published in the past two years (Gao et al. 2012, 2013b), and one of the authors
of the present study (Gao Y-D) has proposed that many more new Lilium taxa may exist within the region. On the other
hand, several Lilium species that have been accepted by the authoritative Flora of China may not merit species rank.
In this paper we will provide a revised treatment of HDM Lilium based on several lines of evidence.
Materials and methods
Over the last few years, we completed more than ten expeditions in southwestern China to collect Lilium and its allied
genera with the aim to build a comprehensive molecular phylogeny of this genus. As we made progress on the project,
we recognized that some taxonomic questions needed to be clarified. In particular, we noted that several HDM endemic
species, which were published in 1980s, were documented only by their types or by a few collections. We realized
the importance of reevaluating the taxonomic ranks of species represented by rare collections. Thus, we rigorously
examined specimens in China, visited herbaria (CDBI, KUN, PE, and SZ), and used images in the Chinese Virtual
Herbarium repository (CVH, http://www.cvh.org.cn/). Additionally, we obtained images of specimens of Lilium from
P, K and E herbaria.
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Taxonomic considerations
Lilium brownii F.E. Brown ex Miellez in Spae (1845: 437)
Type:—CHINA, Hubei, Qiayuangu Shan near Xiang-yang, July 1904, Silvestri 199 (FI!).
= Lilium wenshanense Peng & Li (1990: 33) syn. nov. Type:—CHINA, Yunnan, Kunming, Heilongtan, via material cultivated in Kunming
Botanic Garden, 30 Jun 1989, Peng 89-1 (holotype, KUN!).
Description:—Bulb scales articulate or not (the rest of the description follows Liang & Tamura 2000).
Lilium wenshanense was segregated from L. formosanum Wallace (1891: 442) based on the former having
segmented bulb scales (Peng & Li 1990). According to the authors, L. wenshanense is endemic to south Yunnan
(southwest China). Based on the holotype specimen of Lilium wenshanense housed at KUN (Peng 89-1!), the species
resembles L. brownii. This seems reasonable because L. brownii has a wide range in China and a sympatric distribution
with L. wenshanense while L. formosanum is endemic to Taiwan (Liang & Tamura 2000; Gao et al. 2013a).
FIGURE 1. Bulbs of Lilium brownii from different populations showing variable segmentation of the scales. A-B, Materials representing
populations from Nanchuan, Chongqing (Gao G2010001, SZ) showing segmented (A) and unsegmented (B) scales, respectively; C,
Jiangyou, Sichuan (WJ10051401, SZ); D, Lushui, Yunnan (Gao G2009002, SZ).
TAXONOMIC NOTES ON CHINESE LILIUM L. Phytotaxa 172 (2) © 2014 Magnolia Press 103
The main reason to question the species rank of L. wenshanense is the key feature used to erect it. The articulated
bulb scales described in L. wenshanense can be found in many natural populations of L. brownii in China, including
within some populations that exhibit both articulated and non-articulated bulbs (Fig. 1). Thus, the bulb scale feature
cannot be used to distinguish L. wenshanense from L. brownii. Moreover, molecular phylogenetic analyses of the
internal transcribed spacer (ITS) (Gao et al. 2012, 2013a, b) also suggest that there is no boundary between L. brownii
and L. wenshanense. Based on molecular phylogeny and continuous variation of the bulb scale characteristic, we
propose that only one taxon should be recognized.
Another reason to question the status of L. wenshanense is the source of type specimen. Only the holotype
specimen exists, and it was prepared from an accession of unknown origin in cultivation at Kunming Botanical Garden.
Thus, the native habitat of L. wenshanense remains unknown, and the original authors did not investigate individuals
other than the type. Concerns over the type specimen may further support our assertions that L. wenshanense is not
deserving a recognition as a distinct species.
Lilium taliense Franchet (1893: 319)
Type:—CHINA, Yunnan, between Hokin (Heqin) and Tali (Dali), 24 Jul 1883, J.M. Delavay s.n. (syntype P00730930 P!); [China]
Yunnan, Hokin (Heqin), Kou-toui, on rocky slope, 17 Jul 1889, J.M. Delavay s.n. (syntype P00730931, P!).
= Lilium jinfushanense Peng & Wang (1986: 225) syn. nov. Type:—CHINA, Yunnan, Kunming, Heilongtan, via material cultivated in
Kunming Botanic Garden, 3 Aug 1984, Peng 84-23 (holotype, KUN!).
Description:—All tepals white or sometimes inner tepals adaxially yellow (the rest of the description follows Liang
& Tamura 2000).
Distribution and habitat:—Forests, grassy slopes; 2000–3600 m (based on annotated specimens). Sichuan,
Yunnan, Chongqing, Guizhou.
Lilium jinfushanense was described and typified from cultivated material at Kunming Botanical Garden (Peng
84-23!) derived from bulbs originally collected on Jinfo Mountain (Chongqing and Guizhou provinces). Based on
the published material, L. jinfushanense differs from L. taliense Franchet (1892: 319) by having more purple spots at
the throat of flower and yellowish tepals. However, we have observed purple spots and yellowish flowers in natural
populations of L. taliense, which occurs in northwestern Yunnan (Fig. 2A, B). Thus, the spots and tepal color appear
insufficient to justify the species status of L. jinfushanense. We more rigorously compared L. jinfushanense to L.
taliense by visiting the L. jinfushanense type locality at Jinfo Mountain. There, we observed three populations of L.
taliense with inner tepals having yellowish coloring on their adaxial surfaces (Fig. 2C) and, thus, bearing striking
resemblance to L. jinfushanense. We also noticed that the cultivated type specimen of L. jinfushanense exhibits narrower
leaves compared to the native populations. We speculate that the observed variation in leaf size may be indicative
of unpredictable phenotypic expression under cultivation. The available morphological data appear insufficient for
considering L. jinfushanense as a distinct species. Moreover, molecular phylogenetic analyses have resolved the
yellow-tepal taxon from the type locality within L. taliense (Gao et al. 2012, 2013a), and this supports our proposed
treatment. A more broadly circumscribed L. taliense, including L. jinfushanense, has yellow and white inner tepal
morphs and a wider geographic range in Yunnan, Sichuan, and Guizhou.
Lilium lijiangense Peng (1984: 189)
Type:—CHINA, Yunnan, Lijiang, Yulong Snow Range (Yu-Long Xueshan), in understory on rocky cliffs, 3300-3400 m, Sep 1981, Peng
81-26 (holotype KUN!).
= Lilium huidongense Xu (1985: 232) syn. nov. Type:—CHINA, Sichuan, Huidong, Leidaniu Village, on rocky cliff. S. K. Wu 1526
(holotype SZ!, isotype KUN!).
Distribution and habitat:—Forests, grassy slopes of cliffs, 2800–3400 m (based on annotated specimens). SW
Sichuan, NW and NE Yunnan.
Lilium huidongense was published in 1985 based on two specimens collected in June 1959 by S.K. Wu (S. K. Wu
1568, holotype SZ!, isotype KUN!) bearing buds rather than flowers. Unfortunately, the bud colors have faded over
time in all the available type material. Both the holotype and isotype bear a collection label indicating that the bud color
was ‘pink’. The pink color probably explains Professor Heng Li’s 1988 annotation of the isotype as L. bakerianum
Coll. et Hemsl. (1891: 138) var. rubrum Stern (1948: 4), which bears pink flowers and was known to occur near the
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104 Phytotaxa 172 (2) © 2014 Magnolia Press
FIGURE 2. Flowers of Lilium taliense Franchet from NW Yunnan showng continuous morphological variations in tepal spots andcolor
i.e., white to yellow). Materials representing populations from A, Wengshui, Yunnan (Kham Expedition 10-56, PE) showing variability
of spot tepal spots; B, Gezan, Yunnan (Kham Expedition 10-10, PE) showing variability of tepal color on a single individual; C, Jinfo
Mountain, the type locality of L. jinfushanense exhibiting yellow inner tepals.
TAXONOMIC NOTES ON CHINESE LILIUM L. Phytotaxa 172 (2) © 2014 Magnolia Press 105
L. huidongense type locality. However, our own examinations of both types reveal that L. huidongense appears identical
in all features to L. lijiangense, which has yellow flowers. We visited the type locality of L. huidongense in Huidong
County, Sichuan and observed that L. huidongense exhibits leaf morphology typical of L. lijiangense. In particular,
the leaves have5–7(–9) veins and clusters of curly hairs at the bases of leaves, which are rare features of wild Chinese
lilies (voucher: Sichuan, Leibo, Leidaniu Village, Gao 20090729 SZ!). We also observed L. bakerianum var. rubrum
nearby the type locality of L. huidongense (voucher: ibidem, Gao G09008, SZ!), and suspect that the pink descriptor
on both type labels may result from a mistake made during the field work. We conclude that the shared morphology of
L. huidongense and L. lijiangense are sufficient to synonymize them.
Additional misidentified specimens of L. lijiangense were revealed by our herbarium and literature research. In
particular, L. ningnanense Xu (1986: 68) was named and described in 1986 from material collected close to the type
locality of L. huidongense. L. ningnanense was subsequently merged with L. lijiangense in the Flora of China (Liang
& Tamura 2000) based on shared features including yellow tepals with purple spots and the leaf veins as well as the
leaf base hairs. Moreover, a specimen (Yunnan, Kunming, Dongchuan District, Shekuai Town, Jiulong Village, Peng
Hua et al. 8253, KUN!) identified as Nomocharis pardanthina Franchet (1889: 113), was also collected nearby the
type locality of L. huidongense and exhibits morphology characteristic of L. lijiangense. Thus, both L. huidongense
and L. ningnanense are synonyms of L. lijiangense and the N. pardanthina specimen of Peng Hua et al. (Yunnan,
Kunming, Dongchuan District, Shekuai Town, Jiulong Village, Peng Hua et al. 8253, KUN!) represents an additional
collection of L. lijiangense. The existing and new specimens of L. lijiangense that result from our research show
that the species has a geographic distribution wider than previously known. Previously, it was considered limited to
northwestern Yunnan (Peng 1984; Liang & Tamura 2000), but we now recognize that it also occurs in southwestern
Sichuan and northwestern Yunnan provinces.
Representative specimens examined: L. browniiCHINA. Anhui: Huanshan, 30 Jul 1973, X. Zhou & C. Pan
314 (PE!). Beijing: Fangshan, 23 Jul 1933, K.M. Liou 957 (PE!). Chongqing: Chengkou, 19 Jul 1959, P.Y. Li 5276
(PE!); ibidem, 18 Jul 1959, P.Y. Li 5388 (PE!); ibidem, 20 Jul 1959, P.Y. Li 6661 (PE!); ibidem, 26 Jul 1958, T.L. Dai
101487 (PE!); ibidem, 30 Aug 1958, T.L. Dai 102224 (PE!); ibidem, 15 Sep 1958, T.L. Dai 102751 (PE!); ibidem, 14
Oct 1958, Anonymous 103760 (PE!); ibidem, 3 Sep 1958, T.L. Dai 107086 (PE!); ibidem, 20 Jul 1958, Anonymous
105716 (PE!); Fengjie, 2 Jul 1958, Z.R. Zhang 25493 (PE!); ibidem, 28 Jun 1958, H.F. Zhou 26461 (PE!); ibidem, 24
Jun 1964, H.F. Zhou & H.Y. Su 108624 (PE!); Nanchuan, 28 Jun 1964, K.J. Guan et al. 964 (PE!); ibidem, 27 May 1928,
W.P. Fang 1038 (PE!); ibidem, 4 Jun 1935, Z.X. Qu 1100 (PE!); ibidem, 19 Jun 1935, Z.X. Qu 1465 (PE!); ibidem, 27
Jun 1957, G.F. Li 72387 (PE!); Wanyuan, 1 Jul 1959, P.Y. Li 4351 (WUK!); ibidem, 1 Aug 1959, P.Y. Li 6026 (WUK!);
ibidem, 5 Oct 1971, 263 Sichuan Expedition 1336 (PE!); Wushan, 1 Aug 1964, H.F. Zhou & H.Y. Su 109889 (WUK!);
ibidem, 25 Jul 1958, G.H. Huang 58953 (PE!); Wuxi, 30 Aug 1958, G.H. Huang 59443 (PE!); Youyang, 4 May 1959,
School of Biology of Southwest Normal University 2325 (PE!). Fujian: Changting, 6 May 1946, Anonymous 5192 (PE!);
Taining, 16 Jun 1978, G.L. Cai 481 (KUN!); ibidem, 16 Jun 1934, L.G. Lin 695 (PE!); Wuyishan, 5 Jul 1974, 236-6
Expedition 564 (PE!); ibidem, 30 Jul 1964, Z.P. Jian et al. 400819 (PE!); Huixian, 1 Jun 1956, B.Z. Guo 3143 (WUK!);
ibidem, 15 Aug 1965, Anonymons 5331 (PE!). Gansu: Kangxian, 1 Oct 1963, Z.Y. Zhang 17071 (WUK!); Qingshui, 1
Jun 1986, J.X. Yang 6967 (WUK!); Tianshui, 1 Jul 1951, Z.W. Zhang 90 (WUK!); ibidem, 1 Jun 1957, Y.W. Cui 10075
(WUK!); ibidem, 1 Aug 1951, J.M. Liu 10394 (WUK!); ibidem, 1 Aug 1960, K.J. Fu 13941 (WUK!); ibidem, 1 Aug
1960, K.J. Fu 13969 (WUK!); ibidem, 1 Jul 1964, K.J. Fu 15757 (WUK!); Wenxian, 1 Oct 1958, Z.P. Wei 2924 (WUK!);
ibidem, 1 Oct 1958, Z.P. Wei 3098 (WUK!); ibidem, 1 Jul 1959, Z.Y. Zhang 6507 (WUK!); ibidem, 1 Jul 1959, Z.Y.
Zhang 7556 (WUK!); ibidem, 1 Jul 1959, Z.Y. Zhang 8786 (WUK!); ibidem, 1 Sep 1959, Z.Y. Zhang 13714 (WUK!);
ibidem, 1 Oct 1974, J.Q. Xin 21869 (WUK!); Wudu, 1 May 1959, Z.Y. Zhang 2904 (WUK!); ibidem, 1 Jun 1959, Z.Y.
Zhang 5463 (WUK!); Zhangxian, 12 Jun 1956, Huanghe Expedition 4777 (PE!); Zhouqu, 1 Jul 1964, B.Z. Guo 5503
(WUK!); Boluo, 11 May 1930, N.J. Chen 40971 (PE!); Dapu, 12 Jul 1932, W.T. Tsang 21161 (PE!). Guangdong:
Heping, 12 Jun 1958, Z.F. Wei 120489 (KUN!); ibidem, 12 Jun 1958, Z.F. Wei 120490 (PE!); Huiyang, 16 Sep 1958, Z.F.
Wei 121714 (PE!); Nanxiong, 25 Jun 1958, L. Deng 6544 (KUN!); Pingyuan, 2 Jul 1968, X.G. Li 202033 (PE!); Qujiang,
6 Jul 1924, To & Tsang 12674 (PE!); Raoping, 28 May 1957, X.G. Li 200810 (PE!); ibidem, 12 Jun 1957, X.G. Li 200915
(PE!); Shixing, 8 Jul 1958, L. Deng 6658 (PE!); Xianggang, 20 Jun 1970, Shiu Ying Hu 10453 (PE!); Yangshan, 15 Jun
1956, L. Deng 1467 (KUN!); Fusui, 27 Apr 1957, S.Q. Chen 12096 (PE!); Jingxi, 23 Sep 1935, X.P. Gao 55816 (PE!).
Guangxi: Jinxiu, 29 May 1957, S.Q. Chen 12520 (KUN!); Lingyun, 8 Jun 1933, A.N. Steward & H.C. Cheo 611 (PE!);
Longzhou, 28 May 1935, X.P. Gao 55158 (PE!); Rongshui, 9 Jun 1957, D.Z. Chen 651 (KUN!); ibidem, 5 Jun 1959,
Q.H. Lue 2352 (PE!); ibidem, 30 Aug 1958, S.Q. Chen 15427 (PE!); Tinalin, 2 Jun 1936, H.H. Su 67544 (PE!); Anshun,
30 May 1935, S.W. Feng 482 (PE!); Huangping, 3 Jul 1959, Southern Guizhou Expedition 2519 (KUN!). Guizhou:
Jiangkou, 17 Jun 1959, T.P. Zhu 1239 (KUN!); Songtao, 10 Jul 1959, Northern Guizhou Expedition 1422 (PE!); Songtao,
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106 Phytotaxa 172 (2) © 2014 Magnolia Press
10 Jul 1959, T.P. Zhu 1432 (KUN!); Xifeng, 1 Jul 1936, S.W. Deng 90475 (PE!); Luanchuan, 1 Oct 1983, Loess Plateau
Expedition 1885 (WUK!); Lushi, 1 Sep 1958, J.Q. Fu 1441 (WUK!). Henan: Lushi, 1 Oct 1958, J.Q. Fu 2055 (WUK!);
ibidem, 13 Jul 1935, K.M. Liou 4726 (PE!); Songxian, 26 Aug 1960, Henan Expedition 2362 (PE!); ibidem, 16 Sep
1960, Guan & Dai 2636 (PE!); ibidem, 23 Aug 1959, Anonymous 35078 (PE!); ibidem, 23 Aug 1959, Anonymous 35078
(PE!); Xinyang, 18 Jul 1965, Y.L. Chen & T.L. Min 3 (PE!); Xishan, 13 Jul 1960, K.J. Guan & T.L. Dai 1108 (PE!);
Enshi, 25 Jun 1958, M.Y. Fang 24486 (PE!); Fangxian, 11 Aug 1938, K.M.Liou 9180 (PE!). Huibei: Junxian, 1 Jun
1973, S.Q. Zhong 134 (WUK!); ibidem, 1 Aug 1959, J.Q. Xin 10928 (WUK!); Laifeng, 13 Aug 1958, H.J. Li 4973 (PE!);
Xingshan, 17 Jun 1957, G.X. Fu & Z.S. Zhang 205 (PE!); ibidem, 10 Jul 1957, H.J. Li 974 (PE!); Xuanen, 1 Jul 1958,
H.J. Li 4763 (WUK!); ibidem, 11 Jul 1958, H.J. Li 3775 (PE!); ibidem, 13 Jul 1958, H.J. Li 4208 (PE!). Hunan:
Changning, 10 Jul 1936, C.S. Fan & Y.Y. Li 226 (PE!); Hengshan, 15 Aug 1953, K. J. Guan & B.M. Yang 223 (PE!);
ibidem, 14 Aug 1948, Y. Liu 406 (PE!); Sangzhi, 1 Jun 1958, L.H. Liu 9131 (WUK!); Xinning, 14 Sep 1984, Ziyunshan
Expedition 522 (PE!); Anfu, 1 Aug 1963, J.S. Yue et al. 3036 (WUK!); ibidem, 18 Aug 1959, S.K. Lai 1949 (PE!).
Jiangxi: Dayu, 7 Jul 1965, M.X. Nie et al. 9284 (KUN!); Guangcang, 16 Oct 1962, J.S. Yue et al. 2514 (PE!); Guixi, 26
Oct 1979, Shen & Huang 302 (LBG!); Huichang, 4 Jul 1958, Q.M. Hu 3066 (PE!); ibidem, 13 Jul 1958, Q.M. Hu 3486
(PE!); Jingangshan, 30 Jul 1965, S.K. Lai 4549 (KUN!); ibidem, 7 Sep 1963, M.X. Nie et al. 7867 (PE!); Lianhua, 1 Aug
1959, S.K. Lai 1629 (LBG!); Longnan, 27 Jun 1970, Mission 236 Team 1097(a) (PE!); ibidem, 27 Jun 1970, Mission
236 Team 1097(b) (PE!); Ningdu, 25 Oct 1958, Q.M. Hu 5637 (LBG!); Pingxiang, 20 Oct 1954, Jiangxi Expedition
2662 (PE!); Qianshan, 19 Aug 1958, S.K. Lai 4317 (KUN!); Shangrao, 12 Sep 1958, S.K. Lai et al. 4959 (KUN!);
Suichuan, 3 Oct 1963, J.S. Yue et al. 4567 (KUN!); Tongluo, 7 Jun 1959, J. Xiong 4327 (LBG!); Xunwu, 10 Aug 1962,
J.S. Yue et al. 1744 (KUN!); Xunwu, 22 Jul 1951, Y. Lin 15024 (LBG!); Yihuang, 5 Aug 1959, Q.H. Li 2000 (PE!);
Yongxin, 10 Jul 1959, S.K. Lai 1285 (PE!); Zixi, 10 Jul 1958, M.X. Nie 3260 (LBG!); Danfeng, 1 Aug 1958, B.Z. Guo
3708 (WUK!); Fengxian, 1 Jun 1960, K.J. Fu 12599 (WUK!). Shaanxi: Foping, 1 Sep 1958, X.M. Zhang 545 (WUK!);
ibidem, 30 Jun 1952, K.J. Fu 4830 (PE!); Hanzhong, 1 Oct 1960, P.Y. Li 10030 (WUK!); Huxian, 1 Jul 1951, B.Z. Guo
148 (WUK!); Lanao, 1 Oct 1957, Z.P. Wei & S.H. Luo 312 (WUK!); ibidem, 1 Jul 1959, P.Y. Li 8460 (WUK!); ibidem,
1 Jul 1959, P.Y. Li 8491 (WUK!); ibidem, 25 Jul 1959, P.Y. Li 7763 (KUN!); ibidem, 29 Jul 1959, P.Y. Li 7846 (KUN!);
Liuba, 1 Sep 1977, Z.X. Hu & Y.H. Guo 780 (WUK!); Meixian, 1 Oct 1958, X.M. Zhang 182 (WUK!); ibidem, 1 Sep
1962, K.J. Fu 14758 (WUK!); Mianxian, Oct 1942, K.J. Fu 4013 (WUK!); Nanzheng, 1 Aug 1972, Y.Q. He 7925
(WUK!); ibidem, 1 Sep 1957, K.J. Fu 10787 (WUK!); Ningshan, 1 Sep 1961, Z.W. Zhang 1012 (WUK!); ibidem, 1 Sep
1961, Z.W. Zhang 1209 (WUK!); ibidem, 1 Sep 1961, Z.Y. Zhang 1222 (WUK!); ibidem, 1 Jun 1959, J.Q. Xin 6697
(WUK!); ibidem, 1 Jul 1978, K.J. Fu 17614 (WUK!); ibidem, 1 Jul 1978, K.J. Fu 17781 (WUK!); Shiquan, 1 May 1959,
J.Q. Xin 3710 (WUK!); Taibai, 1 Sep 1958, S.H. Luo 230 (WUK!); ibidem, 1 May 1959, J.X. Yang 437 (WUK!); ibidem,
Jun 1958, Z.P. Wei 895 (WUK!); ibidem, Jul 1956, K.J. Fu 8266 (WUK!); ibidem, Jul 1957, K.J. Fu 9145 (WUK!);
ibidem, 9 Oct 1955, Zhou et al. 58 (PE!); Weinan, Jul 1952, Z.B. Wang 15801 (WUK!); Zhengping, Jul 1991, J.S. Ying
31 (WUK!); Zhouzhi, 1 Aug 1958, X.M. Zhang 219 (WUK!); Hengqu, 1 Jul 1984, Shanxi Expedition 2206 (WUK!);
Qinyuan, 15 Jul 1959, K.J. Guan & Y.L. Chen 896 (PE!). Shanxi: Yangchen, 31 Jul 1959, S.Y. Bao & S.J. Yan 2194
(PE!). Sichuan: Ebian, 8 Aug 1939, Z.W. Yao 4528 (PE!); Emeishan, 18 Jul 1973, Anonymous 11 (PE!); ibidem, 5 Aug
1927, T.Y. Zhou & G.J. Xu 26 (PE!); ibidem, 13 Jul 1971, 263 Sichuan Expedition 43 (PE!); ibidem, 28 Sep 1939, T.N.
Liou & C. Wang 944 (PE!); ibidem, 29 Oct 1938, T.N. Liou 12832 (PE!); ibidem, 6 Jul 1957, G.H. Yang 55713 (PE!);
Jiading, 3 Jul 1928, H.C. Zhou 68 (PE!); Jiulong, 1 May 1959, M.X. Wang 7520 (PE!); Leibo, 8 Jul 1960, S. Jiang et al.
7571 (PE!); Maoxian, 16 Jun 1959, Mao Wen Expedition 2732 (PE!); Meishan, 19 May 1959, Anonymous 5014 (PE!);
Muchuan, 7 May 1959, Muchuan Expedition 2120 (PE!); Pingshan, 24 Aug 1931, Wang & Tu 420 (PE!); Tianquan, 30
May 1959, Sichuan Angreculture School 438 (PE!); ibidem, 13 Sep 1963, K.J. Guan et al. 3418 (PE!); Zhaohua, 1 Nov
1958, Z.P. Wei 3602 (WUK!); Daguan, 16 Jun 1973, B.X. Sun 627 (PE!); Eryuan, 30 Jun 1929, R.C. Ching 23041 (PE!).
Yunnan: Wenshan, 29 May 1943, B.Q. Zhong & K.R. Kuang 441 (PE!); Fengqing, 17 Jun 1938, T.T. Yu 16339 (KUN!);
Fugong, 3 May 1978, Bijiang Expedition 289 (KUN!); ibidem, 21 May 1964, J.S. Yang 3423 (KUN!); ibidem, 29 May
1982, Qing Zang Expedition 6875 (KUN!); Funing, 24 May 1940, C.W. Wang 89569 (PE!); Haoming, 21 Sep 1957, B.Y.
Qiu 55141 (PE!); Hekou, 6 Apr 1941, S.E. Liu 18519 (PE!); ibidem, 9 Sep 1939, C.W. Wang 81657 (PE!); Jiangchuan,
9 Aug 1975, B.Y. Qiu 60637 (KUN!); Jinping, 22 May 1956, China Russian Expedition 2487 (PE!); ibidem, 21 May
1956, China Russian Expedition 3155 (PE!); Lingcang, 18 Sep 1939, H.C. Ching 30641 (PE!); Lushui, 27 Jul 1978,
Nujiang Expedition 1949 (KUN!); Maguan, 2 Jun 1959, Q.A. Wu 8211 (KUN!); Pingbian, 27 May 1934, H.T. Tsai 55462
(PE!); ibidem, 10 Jun 1934, H.T. Tsai 60151 (PE!); Xichou, 25 May 1964, S.Z. Wang 421 (KUN!); ibidem, 29 Apr 1959,
Q.A. Wu 7787 (KUN!); ibidem, 27 May 1984, D.Z. Fu 84306 (PE!); Yangbi, 16 Jun 1984, B. Bartholomew et al. 159
(KUN!); Zhenxiong, 10 Jun 1980, S.Y. Bao 196 (KUN!); Jiande, 8 Jul 1958, Anonymous 29539 (PE!). Zhejiang:
Longquan, Jul 1958, R.H. Shan 5505 (PE!).
TAXONOMIC NOTES ON CHINESE LILIUM L. Phytotaxa 172 (2) © 2014 Magnolia Press 107
L. talienseCHINA. Chongqing: Nanchuan, 2 Jul 1978, Geological survey team of plant 697 (CDBI!); ibidem,
1 Jul 1978, Geological survey team of plant 569 (CDBI!); ibidem, 2 Jun 1978, Geological survey team of plant 160
(CDBI!); ibidem, 21 Jul 1957, G.F. Li 63021 (PE!); ibidem, 11 Aug 1957, J.H. Xiong et al. 92596 (PE!); ibidem, 3 Jul
1964, K.J. Guan et al. 1058 (PE!). Guizhou: Tongren, 16 Jul 1939, B.Q. Zhong 886 (PE!). Sichuan: Muli, s.d., T.T.
Yu 7366 (KUN!); ibidem, 20 Aug 1937, T.T. Yu 14059 (KUN!); ibidem, 1 Aug 1978, Q.S. Zhao 8543 (CDBI!); ibidem,
2 Jul 1978, Q.S. Zhao et al. 6756 (CDBI!); ibidem, 2 Aug 1978, Q.S. Zhao et al. 7925 (CDBI!); ibidem, 20 Aug 1937,
T.T. Yu 14059 (PE!). Yunnan: Dali, 27 May 1964, Li 78 (KUN!); Eryuan, 17 Jul 1940, R.C. Ching 23272 (KUN!);
ibidem, 22 Aug 1963, Yunnan Institute for drug control 63-1883 (KUN!); ibidem,17 Jul 1940, R.C. Ching 23272
(PE!); Jianchuan, 8 September 1956, P.Y. Mao 246 (PE!); Jingdong, 1 Aug 1975, B.Y. Qiu 52903 (PE!); Kunming, s.d.,
S.H. Yuan 101118 (KUN!); ibidem, 1 Aug 1975, B.Y. Qiu 59128 (CDBI!); ibidem, 26 Oct 1941, S.E. Liu 19206 (PE!);
Langping, 28 Jul 1963, J.S. Yang 5030 (KUN!); Lijiang, 28 Jul 1959, Kunming Botanical Garden 22464 (KUN!);
ibidem, 19 Jul 1962, A.L. Zhang 100550 (KUN!); ibidem, 15 Jul 1939, Z.G. Zhao 30397 (KUN!); ibidem, 29 Jul 1940,
R.C. Ching 30929 (KUN!); ibidem, 25 Jul 1939, Z.G. Zhao 30451 (KUN!); ibidem, 24 Aug 1939, R.C. Ching 21451
(KUN!); ibidem, 2 Jul 1939, C.Y. Zhao 20959 (KUN!); ibidem, Aug 1963, Lijiang Botanical Garden 100550 (KUN!);
ibidem, 29 Aug 1937, T.T. Yu 15509 (PE!); ibidem, Jul 1922, J.F. Rock 5171 (PE!); ibidem, 15 Jul 1937, R.C. Ching
30397 (PE!); ibidem, s.d., R.C. Ching 21421 (PE!); ibidem, 24 Aug 1939, Y.Z. Zhao 21451 (PE!); ibidem, 18 Jul 1956,
P.Y. Mao 131 (PE!); ibidem, 1910, G. Forrest 6152 (PE!); ibidem, Aug 1922, G. Forrest 23221 (PE!); s.l., s.d., E.E.
Maire 4008 (PE!).
L. lijiangense—CHINA. Sichuan: Huidong, 29 Jul 2009, Y.D. Gao G20090729 (SZ!); ibidem, 26 Jun 1956, S. K. Wu
1568 (SZ!, KUN!); Ningnan, 12 Jul 1978, Ningnan Exped. 0321 (HSM!). Yunnan: Lijiang, Jul 1981, L.J. Peng 81-26
(KUN!); Kunming, 26 Jul 2008, Peng Hua et al. 8253 (KUN!).
Acknowledgements
The author appreciate the assistance of the staff in the herbaria of CDBI, E, HUH, K, KUN, P, PE, SZ and FI in the
study of specimens. We also give our special gratitude to AJ Harris from Oklahoma State University for her help
in English editing, and Dr. Li-Bing Zhang of Missouri Botanic Garden, Timothée Le Péchon from School of Life
Sciences, University of KwaZulu-Natal, South Africa for their help in literature survey and suggestions. This work
was supported by the General Financial Grant from the China Postdoctoral Science Foundation to Y. D. Gao (Grant
No. 2013M542296).
Reference
Collect, H. & Hemsley, W.B. (1891) On a collection of plants from Upper Burma and Shan States. Journal of the Linnean Society, Botany
28: 1–150.
Elwes, H.J (1880) A monograph of the genus Lilium. Taylor and Francis, London, 222 pp.
http://dx.doi.org/10.5962/bhl.title.536
Franchet, A.R. (1889) Nomocharis. Journal de Botanique (Morot) 3: 113–114.
Franchet, A.R. (1892) Les Lis de la Chine et du Thibet. Journal de Botanique (Morot) 6: 305–321.
Gao, Y.D., Hohenegger, M., Harris, A., Zhou, S.D., He, X.J. & Wan, J. (2012) A new species in the genus Nomocharis Franchet (Liliaceae):
evidence that brings the genus Nomocharis into Lilium. Plant Systematics and Evolution 298: 69–85.
http://dx.doi.org/10.1007/s00606-011-0524-1
Gao, Y.D., Harris, A., Zhou, S.D. & He, X.J. (2013a) Evolutionary events in Lilium (including Nomocharis, Liliaceae) are temporally
correlated with orogenies of the Q–T plateau and the HDM. Molecular Phylogenetics and Evolution 68: 443–460.
http://dx.doi.org/10.1016/j.ympev.2013.04.026
Gao, Y.D., Zhou, S.D. & He, X.J. (2013b) Lilium yapingense (Liliaceae), a new species from Yunnan, China, and its systematic significance
relative to Nomocharis. Annales Botanici Fennici 50: 187–194.
http://dx.doi.org/10.5735/085.050.0311
Haw, S.G. (1986) The Lilies of China. Timber Press, Portland, 152 pp.
GAO & GAO
108 Phytotaxa 172 (2) © 2014 Magnolia Press
Liang, S.Y. & Tamura, M. (2000) Lilium L.. In: Wu, Z.Y. & Raven, P.H. (eds.) Flora of China 24. Science Press, Beijing; & Missouri
Botanical Garden Press, St. Louis, pp. 135–159.
Linnaeus C. (1753) Species Plantarum. Laurentius Salvius, Stockholm, Sweden, 1200 pp.
Spae D. (1845) Lilium brownii. Annales de la société royale d’agriculture et de botanique de Gand 1: 437–438.
Peng, L.J. (1984) One new species of Lilium from Yunnan. Acta Botanica Yunnancia 6: 189–191(in Chinese).
Peng, L.J. & Li, X.F. (1990) Lilium wenshanense, one new species of Lilium. Acta Botanica Yunnanica, Suppl. 3: 33–34 (in Chinese).
Peng, L.J. & Wang, B.N. (1986) One new species of Lilium from Sichuan. Acta Botanica Yunnanica 8: 225–256 (in Chinese).
Stearn, W.T. (1948) The botanical name of some lilies. The Gardeners’ Chronicle: a weekly illustrated journal of horticulture and allied
subjects 3: 4–5.
Wallace, A.R. (1891) Dr. Wallace on Japanese lilies. Garden, an illustrated weekly journal of gardening in all its branches 40: 380–442.
Xu, J.M. (1985) New species of Liliaceae from Sichuan. Acta Phytotaxonomica Sinica 23: 232–235 (in Chinese).
Xu, J.M. (1986) Two new species of Lilieae from Sichuan. Bulletin of Botanical Research 6: 67–70 (in Chinese).
... Thus, based on morphological evidences, we believe that these collections of Father Ten should be assigned to Lilium lijiangense L.J. Peng (1984: 189) (Figs. 3 & 4). In prior studies, we have completely revised the circumscription of L. lijiangense (Gao & Gao 2014), in which the major distinguishing features were seven to nine veins on broad lanceolate leaves and white hair as we observed in the two collections of Father Ten. Moreover, Lilium lijiangense has a geographic range that includes the region of Father Ten's collections, based on two newly collected specimens from Dongchuan, Yunnan (Peng Hua et al. 8253,KUN!,Gao ...
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The name Lilium tenii H.Léveillé (1909: 263) was assigned by Hector Léveillé to a specimen collected by Father Siméon Ten on 15 August 1907 from Dongchuan (Tong-Tchouan) of Yunnan Province, China. The name L. tenii has long been treated as a synonym for Lilium primulinum var. ochraceum (Franch.) Stearn (1948: 13). The treatment was based on the monograph of Chinese lilies by William Wright Smith (1923). Smith (1923) mentioned Lilium tenii when he revised L. ochraceum Franchet (1892: 319), which was later reduced to a variety under L. primulinum Baker (1892: tab. 7227). However, Smith (1923) did not treat L. tenii as a synonym of L. ochraceum. Instead, Smith (1923) pointed the distinctness of L. tenii by saying: ‘This is a puzzling plant….. I can find nothing quite like the leaves of Tenii……some of the leaves having 9–11 distinct nerves. On the evidence before me I cannot justify the reduction of this species, at any rate meanwhile, to any previously described species. It is a near ally of ochraceum (p134–135)’. As part of his revision of the taxonomy of Léveillé, Smith (1923) regarded L. tenii as ‘possibly valid (p141)’ and did not synonymize it within any other species.
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We describe and illustrate Lilium yapingense sp. nova (Liliaceae) and show its position within the Lilium—Nomocharis complex (Liliaceae). It is similar in appearance to L. nanum but differs by (1) having no spots on the tepal bases, instead possessing symmetric stripes; (2) nectaries lacking fimbriate projections on the surfaces, but having two dark grooves; and (3) an orange-colored instead of a white bulb. Phylogenetic analyses using nuclear ITS showed that L. yapingense merits specific rank and that it is more closely related to Nomocharis than to Lilium. However, the morphological synapomorphies thought to distinguish Nomocharis from Lilium are absent from the new species. The morphology and phylogeny of L. yapingense support previous studies, which show that Nomocharis and Lilium have intergrading morphologies and that Lilium is paraphyletic with respect to Nomocharis.
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The Hengduan Mountains (H-D Mountains) in China flank the eastern edge of the Qinghai-Tibet Plateau (Q-T Plateau) and are a center of great temperate plant diversity. The geological history and complex topography of these mountains may have prompted the in situ evolution of many diverse and narrowly endemic species. Despite the importance of the H-D Mountains to biodiversity, many uncertainties remain regarding the timing and tempo of their uplift. One hypothesis is that the Q-T Plateau underwent a final, rapid phase of uplift 8-7 million years ago (Mya) and that the H-D Mountains orogeny was a separate event occurring 4-3 Mya. To evaluate this hypothesis, we performed phylogenetic, biogeographic, divergence time dating, and diversification rate analyses of the horticulturally important genus Lilium, including Nomocharis. The Lilium-Nomocharis complex is distributed throughout the temperate Northern Hemisphere but is most diverse within the H-D Mountains and Q-T Plateau. Our matK and ITS phylogenies support previous studies showing that Nomocharis is nested within Lilium. However, we detected incongruence between the two gene trees which may result from hybridization. Dating analyses performed using the ITS dataset showed that the evolution of major lineages within Lilium-Nomocharis may be temporally coincident with Q-T Plateau uplift occurring 8-7 Mya and H-D Mountains uplift approximately 4-3 Mya. Our analyses of diversification times and rates among Lilium-Nomocharis clades are less conclusive. However, these do suggest high extinction rates among H-D Mountains lineages.
Zhu 1432 (KUN!); Xifeng
Jul 1959, T.P. Zhu 1432 (KUN!); Xifeng, 1 Jul 1936, S.W. Deng 90475 (PE!); Luanchuan, 1 Oct 1983, Loess Plateau Expedition 1885 (WUK!);
  • Ibidem
ibidem, 13 Jul 1935, K.M. Liou 4726 (PE!);
Henan Expedition 2362 (PE!); ibidem
  • Songxian
Songxian, 26 Aug 1960, Henan Expedition 2362 (PE!); ibidem, 16 Sep 1960, Guan & Dai 2636 (PE!); ibidem, 23 Aug 1959, Anonymous 35078 (PE!); ibidem, 23 Aug 1959, Anonymous 35078
Fang 24486 (PE!); FangxianLiou 9180 (PE!) Huibei: Junxian
  • K M Enshi
Enshi, 25 Jun 1958, M.Y. Fang 24486 (PE!); Fangxian, 11 Aug 1938, K.M.Liou 9180 (PE!). Huibei: Junxian, 1 Jun 1973, S.Q. Zhong 134 (WUK!); ibidem, 1 Aug 1959, J.Q. Xin 10928 (WUK!); Laifeng, 13 Aug 1958, H.J. Li 4973 (PE!);
  • Xingshan
Xingshan, 17 Jun 1957, G.X. Fu & Z.S. Zhang 205 (PE!); ibidem, 10 Jul 1957, H.J. Li 974 (PE!);