Peonies of the World - Look inside - Royal Botanic Gardens, Kew

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CONTENTS 

SYNOPSIS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vi 

PREFACE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii 

ACKNOWLEDGEMENTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ix 

1. INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 

1.1 Course of the work contributing to this monograph . . . . . . . . . . . . . . . . . . . 1 

1.2 Methodology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 

1.3 Principles of taxonomic treatment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 

2. HISTORY OF TAXONOMIC STUDIES OF PAEONIA . . . . . . . . . . . . . . . . . . . 11 

2.1 Brief review of the taxonomic history of Paeonia since Linnaeus (1753) . . . . . 11 

2.2 Commentary on the taxonomic history of Paeonia . . . . . . . . . . . . . . . . . . . . 21 

3. CHARACTERS AND THEIR VARIATIONS . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 

4. TAXONOMIC SYNOPSIS OF THE GENUS PAEONIA . . . . . . . . . . . . . . . . . . . 49 

I. Paeonia sect. Moutan DC. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50 

Ia. Paeonia subsect. Delavayanae Stern (species 1, 2) . . . . . . . . . . . . . . . . . . . . . 52 

Ib. Paeonia subsect. Vaginatae Stern (species 3–8) . . . . . . . . . . . . . . . . . . . . . . . 53 

II. Paeonia sect. Onaepia Lindl. (species 9, 10) . . . . . . . . . . . . . . . . . . . . . . . . . . . 53 

III. Paeonia sect. Paeonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54 

Subdivisions of sect. Paeonia 

IIIa. Paeonia subsect. Albiflorae (Salm-Dyck) D. Y. Hong (species 11–14) . . . . . 57 

IIIb. Paeonia subsect. Foliolatae Stern (species 15–25) . . . . . . . . . . . . . . . . . . . . 57 

IIIc. Paeonia subsect. Paeonia (species 26–32) . . . . . . . . . . . . . . . . . . . . . . . . . . 58 

5. DESCRIPTIONS OF SPECIES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61 

BIBLIOGRAPHY . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245 

INDEX TO BOTANICAL NAMES IN PAEONIA . . . . . . . . . . . . . . . . . . . . . . . . . . 259 

AUTHORS OF BOTANICAL NAMES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 271 

HERBARIA VISITED OR WITH SPECIMENS ON LOAN . . . . . . . . . . . . . . . . . . 275 

HERBARIUM SPECIMENS EXAMINED AND THEIR 

SPECIES DESIGNATIONS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 277

vi 

PEONIES OF THE WORLD 

SYNOPSIS 

SYNOPSIS 

The present monograph presents a taxonomic revision of the genus Paeonia based on extensive field 

observations, population sampling, and statistical analysis of various characters. All specimens available 

from 65 herbaria were examined and identified, and thus there is a list of herbarium specimens 

examined and their species designations. The monograph recognises three sections, sect. moutan DC., 

sect. Onaepia Lindl. and sect. Paeonia. Two subsections are recognised in sect. Moutan: subsect. 

Delavayanae Stern and subsect. Vaginatae Stern. I recognise three subsections in sect. Paeonia: subsect. 

Paeonia, subsect. Albiflorae (Salm-Dyck) D. Y. Hong, and subsect. Foliolatae Stern. The subsection 

Albiflorae (Salm-Dyck) D. Y. Hong is a new status of sect. Albiflorae Salm-Dyck (1834). Thirty-two 

species and 26 subspecies of wild peonies are recognized in the monograph, including a new 

subspecies from Croatia, Paeonia daurica Andrews subsp. velebitensis D. Y. Hong. Each species is 

described, its distribution is mapped, and its relationships with other species are discussed in detail. 

The monograph reports 23 chromosome countings, including five new records: Paeonia intermedia C. 

A. Mey. (2n=10), P. qiui Y. L. Pei & D. Y. Hong (2n = 10), P. sterniana H. R. Fletcher (2n = 10), 

P. rockii subsp. rockii (2n = 10), and a new chromosome number for P. emodi Wall. ex Royle (2n = 

20). The monograph contains six colour plates, three black and white plates, 78 line drawings of 

plants, and 39 maps. Twelve lectotypes and 17 neotypes are newly designated, and more than 40 taxa 

are reduced as synonymy (syn. nov.).


PREFACE 

vii 

PREFACE 

My monograph on the genus Paeonia will comprise three parts. This book, covering taxonomy and 

phytogeography, is the first. The second will illustrate the polymorphism and diversity within the 

genus with colour photographs taken in the field, accompanied by short descriptions; and the third 

will deal with phylogeny and evolution. 

In the summer of 1984, DAI Lung-Kai, chief officer of the Research Department at the 

Institute of Botany, the Chinese Academy of Sciences, informed me that I should put forward a 

proposal for a project to be funded by a generous grant of 50,000 RMB from the Chinese Academy 

of Sciences. The grant was especially for scholars returned from overseas, and I had been a visiting 

scholar in Sweden from 1979 to 1981. 

In choosing my subject, I wished to reflect a combination of academic and public interests, and 

I had already been thinking about the genus Paeonia. My wife, PAN Kai-Yu, a botanist and author 

of Paeonia for Flora Reipublicae Popularis Sinicae (Vol. 27, 1979), encouraged me to study this 

interesting group. 

The genus Paeonia has been culturally significant in both the East and West for millennia. Tree 

peony has long been called ‘King of Flowers’ in China, and the herbaceous peony was called 

‘Queen of Herbs’ in ancient Greece. 

Systematically, the genus Paeonia is isolated from other groups of angiosperms in many respects; for 

example, its embryogenesis involves the existence of a free nuclei stage, the coenocytic proembryo, it 

contains unique chemical compounds, e.g. paeoniflorin, its anthroecia undergo centrifugal 

development, and chromosome behaviour at meiosis shows permanent structural hybridity. 

The origin of the cultivated tree peony, whose wild relatives are all endemic to China, also 

struck us as worthy of investigation. 

My work on Paeonia started in the spring of 1985, when I collected materials for cytotaxonomy 

in Sichuan, Shaanxi and Hebei provinces with assistance from ZHU Xian-Yun. This work was 

published in two papers: Hong et al. 1988 and Hong 1989. 

Three events that bolstered the Paeonia project took place shortly afterwards. First, in 1990, 

Peter Raven, president of the Missouri Botanical Garden, encouraged me to increase my focus on 

Paeonia when I mentioned it as one of several projects in our laboratory. 

Second, in 1991, Tao SANG (now at Michigan State University) sought my advice on whether 

he should take Paeonia as the subject of the dissertation project for his Ph.D (at Ohio State 

University). I suggested that we divide the work: he should work on molecular systematics and 

evolution while I continued to work on taxonomy, mainly based on morphology. Since then, we 

have been in successful collaboration. 

Third, Dr Wilhelm Sauer invited me and my wife to work at Tübingen University in Germany 

for three months, starting in November 1991. He kindly borrowed for us thousands of Paeonia 

specimens from a number of major European herbaria. 

Readers will readily note differences in the number of species recognised and their 

circumscriptions when comparing the present work with previous monographs or revisions of 

Paeonia. Baker (1884) recognised 22 species, of which only eight are identical or partly identical to 

those described in this monograph. Lynch (1890) described 25 species, but only six correspond with 

those described in this work. Huth (1891) recognised 14 species, of which again only six agree with 

my taxa/taxonomic view. Stern’s (1946) monograph of the genus Paeonia has been the most widely

viii 


PREFACE 

accepted work and still has a significant influence. He recognised 33 species, with only 19 

corresponding with the 32 species in this monograph in both name and circumscription. Halda 

(2004) recognised 25 species, of which only 12 agree with my work. 

The differences, as readers will realise from the first chapter of this book, arise from the 

principles and practices of taxonomy. A rational classification must be based on the biology of plant 

groups. To fully understand the variation of characters, their patterns and ranges, the population 

concept is crucial. With these principles in mind, our practices included: 

1) extensive field observations and population sampling; 

2) critical examination of all available herbarium specimens; 

3) statistical analysis; and 

4) examination of as many type specimens as possible. 

I have found that many of the characters emphasised by previous authors are continuously 

variable or polymorphic, and thus less significant in the taxonomy of Paeonia (Haw, 2001a, 2001b; 

see Chapter 3). I have also found that a number of characters neglected by previous authors — for 

example, the shape of roots, the shape of involucral bracts and sepals, and the number and pattern 

of leaflets or the segmentation of lower leaves — are valuable for taxonomy in Paeonia (see 

Chapter 3). 

Parts of this taxonomic revision of Paeonia have been reinforced by later workers. My treatment 

of tree peonies (Paeonia sect. Moutan DC.) (Hong & Pan, 1999a, 1999b) has been confirmed (Zou 

et al., 1999) or essentially confirmed (Zhao et al., 2004), and has been accepted by Haw (2001a, 

2001b) and Cheng (2005).


ACKNOWLEDGEMENTS 

ix 


This monograph would not have been possible without the help of others. 

I must first thank the National Geographic Society for five grants that were crucial to our work. 

My gratitude also goes to the Chinese Academy of Sciences for the financial support that allowed 

me to start this fascinating project. 

For more than 10 years, Dr P. Raven, president of the Missouri Botanical Garden, has 

encouraged and supported me, giving very valuable suggestions. 

Dr W. Sauer, professor at Tübingen University before his retirement, offered an unexpected 

opportunity to examine thousands of herbarium specimens borrowed from several large European 

herbaria in 1991. Dr Sauer also generously gave me three specimens of Paeonia collected by himself 

and his wife, Gerda Sauer, in the Balkans, which led to the description of a new species, P. saueri, 

which is named after them. 

The English of the first four chapters has been much improved by Miss P. Woodward and Dr J. 

W. Waddick. Dr I. Al-Shehbaz at the Missouri Botanical Garden helped me in a number of ways, 

with valuable suggestions and assistance in the publication of three articles. The late Dr G. H. Zhu 

helped me in searching for literature in the library of the Missouri Botanical Garden. Dr A. Brach 

of Harvard University also provided many valuable references. 

Dr YU Hong from Yunnan University organised both our expedition to NW Yunnan Province 

in 1997, for which we were joined by Prof. GU Hong-Ya from Peking University, and our 

expedition to NE Yunnan in 2004. Prof. LI Xue-Yu from Shihezi University in Xinjiang, organised 

our expedition to the Tianshan and Altai mountains in 1993, for which we were joined by Dr T. 

Sang from Ohio State University and Prof. ZHANG Zi-Yu from the Second Military Medical 

University in Shanghai. In our expeditions to Xizang (Tibet) in 1996 and 2006, Mr XU A-Shen 

made a great contribution, and he also helped me subsequently by supplying Paeonia materials. 

During our expedition to the Caucasus in 1999, particular help was given by Dr Sikharuildze 

and Dr Eristavi from the National Institute of Botany, Georgian Academy of Sciences at Tbilisi. 

We also obtained much help from the directors and staff of the Institute of Botany, Azerbaijan 

Academy of Sciences at Baku (particularly Dr O. V. Ibadov); the National Institute of Botany, 

Georgian Academy of Sciences at Tbilisi; the Batumi Botanical Garden, Georgian Academy of 

Sciences; the Stavropol Botanical Garden, Russia; and the Botanical Garden of Krasnodar 

University, Russia. 

On our expedition to the western Mediterranean in 2001, great assistance was received from 

Dr A. Fridlender of the University of Marseilles. Dr YUAN Yong-Ming at Neuchâtel University, 

Switzerland, led us to the Generoso population of P. officinalis in southern Switzerland. 

Dr T. Koruklu of Ankara University was key to the work in Turkey during our expedition to 

the eastern Mediterranean in 2002. In Greece, Prof. D. Tzanoudakis of the University of Patras 

kindly guided us to several important localities. 

In Japan, Dr M. Nakata, now at the Toyama Botanic Garden, assisted in our trips to Shikoku 

in 1990 and 1992, and Dr CHEN Zhi-Duan of our laboratory joined our trip to Hokkaido in 1998. 

Dr ZHU Xiang-Yun of our institute joined the expeditions to Sichuan, Shaanxi and Hebei 

provinces in 1985, and he again made an expedition to Sichuan and Chongqing (Mt Jinfoshan) 

with Mr SONG Shu-Yin in 1986. Dr QIU Jun-Zhuan, my former student, now in the USA, after 

whom Paeonia qiui Y. L. Pei & D. Y. Hong was named, made field trips to W Hubei, Henan and

x 



NW Sichuan in 1988 and 1989, and Dr PEI Yan-Long, now in Canada, made trips to Shanxi and 

W Hubei in 1991, 1992 and 1993. 

In our expedition to Henan in 1994, the late Prof. WANG Shui-Yi and Dr WANG Yin-Zhen 

formerly from Henan Agricultural University (now in our laboratory) were members of the team. 

Dr HE Yong-Hua from the Chengdu Institute of Biology, assisted our expedition to NW Sichuan 

in 1995 and Dr ZHANG Shu-Ren in our laboratory joined our expedition to Tibet in 1996. 

Prof. YE Yong-Zhong from Henan Agricultural University, Dr FENG Yu-Xin (my former Ph.D 

student now in the USA) and I made a field trip to W Hubei, W Henan and Shaanxi in 1997, which 

was assisted by Mr DAI Zhen-Neng at the Bureau of Forestry of Baokang County, Hubei. At the 

same time, PAN Kai-Yu and Dr XIE Zhong-Wen, now in the USA, made a field trip to Anhui 

Province. In 1998, Mr LOU Lu-Huan helped us in our trip to Mt Tianmu in Zhejiang, and Dr CAO 

Wei assisted us in our expedition to NE China. 

I did field work in Idaho in 1999 with the assistance of Dr Jenny Q. Y. Xian (now at North Carolina 

State University). In 2005, PAN Kai-Yu and I made a field trip to western N America. Assisted by Dr 

QIU Jun-Zhuan, we observed and sampled two populations of P. californica near Los Angeles. Miss 

P. Woodward organised a trip to the Blue Mountains in Oregon and Washington states, where we 

observed and sampled three populations of P. brownii. 

We made several field trips in 2004. Prof. REN Yi, from Shaanxi Normal University, assisted our 

expedition to Mt Taibaishan in Shaanxi. Dr ZHOU Zhi-Qin, of Southwest University, and Mr 

YANG Shi-Xuan joined our trip to Mt Shennongjia in W Hubei. Mr CAO Rui, at Inner Mongolia 

University, and Dr YANG Fu-Shen and Miss PENG Dan, of our laboratory, assisted our expeditions 

to Inner Mongolia. Mr ZHANG Hong-Yue and Mr CHEN Yan, of our laboratory, joined our 

several expeditions to NE China, Hebei, Henan and Inner Mongolia. 

Miss LI Er-Li made the pencil and ink drawings, whereas Miss CAI Shu-Qin and Mr SUN Yin- 

Bao drew the ink drawings for this monograph. All the scanning electron microscope photographs 

were taken by Mr XIAO Yin-Huo and developed by Mr YANG Xue-Jian. Miss LI Qiao-Ling, 

my former secretary, helped a great deal with office work and in the preparation of articles and 

applications for grants. 

To all the persons mentioned above, we are greatly indebted. In addition, our work was helped 

incalculably by many local farmers and villagers in China and elsewhere. We cannot mention them 

individually but we offer them all our sincere thanks. Without them, our work would not have 

been successful.

40 PEONIES OF THE WORLD 

CHARACTERS AND THEIR VARIATIONS 

Fig. 3.10. Polymorphism in indumentum on follicles in P. anomala subsp. veitchii in the population Hong et al. H95034 

(Lixian, Sichuan Province): a, glabrous; b, sparsely hairy. 

Fig. 3.11. Variation in colour of indumentum on carpels and of filaments within and between populations in P. tenuifolia: 

a, Hong & Zhou H99028 from Igoeti, Georgia; b–d, Hong & Zhou H99043 from Mukhrani, Georgia.


CHARACTERS AND THEIR VARIATIONS 

41 

at the apex. For all other species, i.e. most species in sect. Paeonia, the gradation is more or less 

interrupted. The sepals are mostly non-caudate and thus more or less distinct from bracts, though 

inconsistent in shape and size (Fig. 3.12e,f ). 

The bracts-sepals series is of certain taxonomic value. Because of their similar leaflets (or leaf 

segments), Paeonia intermedia and P. anomala had long been confused before Hong and Pan (2004) 

found that P. intermedia has most of its sepals rounded at the apex (Fig. 5.26B), whereas P. anomala 

has all or most of its sepals caudate at the apex (Fig. 5.14B). This character is closely correlated with 

root shape (Figs 5.14A, 5.26A). Using these two characters, the two species are readily 

distinguished. 

The bracts-sepals series also reflects the systematic position of a taxon within Paeonia; in 

general, a gradual gradation of the bracts-sepals series with most or all of the sepals caudate at the 

apex is correlated with primitive characters, such as woody habit, multi-flowered stems (shoots) 

and diploidy. 

3.9 NUMBER OF FLOWERS PER SHOOT 

This character is described relatively easily. In sect. Moutan, the shoots of Paeonia ludlowii and P. 

delavayi are constantly multi-flowered, whereas flowers in the other six species are solitary and 

terminal. In section Onaepia, the two species, P. brownii and P. californica, are both multi-flowered, 

though the flowers often look solitary and terminal. One or two undeveloped (sterile) flower buds 

can often be found in the axils of the upper leaves. For these two sections, there is no taxonomic 

problem caused by misunderstanding of this character. 

In sect. Paeonia, however, all the species except those mentioned below consistently have 

solitary flowers. Paeonia lactiflora and P. emodi are both described as multi-flowered and cause no 

taxonomic problems. Problems occur only in the P. anomala–P. veitchii group. Paeonia anomala was 

described as single-flowered (Schipczinsky, 1937; Pan, 1979), whereas P. veitchii was described as 

multi-flowered (Lynch, 1909; Pan, 1979). The demarcation between these two species was made 

on the basis of this single character. It is true that P. veitchii (= P. anomala subsp. veitchii) often has 

multi-flowered stems, but single-flowered stems with one or two undeveloped flower buds in axils 

of the upper leaves are also often found, and single-flowered stems occur very rarely without any 

additional undeveloped flower buds. Such individuals have been described as a variety, P. veitchii 

var. uniflora K. Y. Pan (1979). In P. anomala (= P. anomala subsp. anomala), flowers are nearly always 

solitary, but, besides the terminal flower, one or two undeveloped flower buds can sometimes be 

found in axils of the upper leaves, and very occasionally there are two flowers on a stem. Paeonia 

altaica K. M. Dai & T. H. Ying (1990) was based on such an individual. In our observation of 

specimens from the site in the Altai (Wuzkiliti, Habahe County) where the type was collected, 

three kinds of individuals coexisted: single-flowered without any additional undeveloped flower 

bud, single-flowered with one or two undeveloped flower buds, and two-flowered. Plainly, the 

character of flower number varies in the P. anomala–P. veitchii group. We therefore treated the 

group as a single species with two subspecies isolated by the Gobi. 

3.10 PETALS 

3.10.1 PETAL COLOUR 

This character has been frequently used in the taxonomy of Paeonia. Many taxa were described on 

the basis of petal colour, e.g. P. lutea, P. delavayi var. atropurpurea, P. delavayi var. alba and P. japonica 

(which is white whereas P. obovata is pink). An extreme case is Kemularia-Nathadze’s


1. PAEONIA LUDLOWII 

61 

5. DESCRIPTIONS OF SPECIES 

I – IIa. PAEONIA sect. MOUTAN subsect. DELAVAYANAE Stern 

(species 1 and 2) 

1. Paeonia ludlowii (Stern & G. Taylor) D. Y. Hong, Novon 7 (2): 157, figs 1, 2 (1997); Hong, 

Pan & Turland in Wu, Raven & Hong, Fl. China 6: 130 (2001); Haw, The New Plantsman 8: 

168 (2001). Basionym: Paeonia lutea Delavay ex Franch. var. ludlowii Stern & G. Taylor, J. Roy. 

Hort. Soc. 76: 217 (1951); Stern & Taylor, Bot. Mag. 169: tab. 209 (1953). Paeonia lutea Delavay 

ex Franch. subsp. ludlowii (Stern & G. Taylor) Halda, Acta Mus. Richnov., Sect. Nat. 6(3): 234 

(1999). Paeonia ludlowii (Stern & G. Taylor) J. J. Li & D. Z. Chen in Li et al., Bull. Bot. Res. 

Harbin 18(2): 154 (1998). TYPE: China, SE Xizang [Tibet]: “Kongbo Prov., Miling, Tsangpo 

Valley”, 28 May 1938, F. Ludlow, G. Sherriff & G. Taylor 4540 (holotype BM!); China, Xizang 

[Tibet], Charme, Char Chu, 28°26'N, 93°05'E, 3,320 m, 22 Oct. 1938, F. Ludlow, G. Sherriff 

& G. Taylor 6392 (paratype BM!); Lung-Chayue Chu, 2,900 m, 28 Apr. 1936, F. Ludlow & G. 

Sherriff 1376 (paratype BM!). 

Paeonia lutea auct. non Delavay ex Franch.: Stern, J. Roy. Hort. Soc. 72: 394, fig. 157 (1947). 

Deciduous and caespitose shrubs, up to 3.5 m tall, glabrous throughout. Roots attenuate downward, not 

fusiform. Stems grey, up to 4 cm in diameter. Shoots green, with 8–12 scales at the base. Lower leaves 

biternate, green above, pale glaucous beneath; petioles 8–18 cm long; leaflets 9, lateral 3 leaflets on each 

side with main petiolules 2–3.5 cm long, terminal 3 leaflets with main petiolules 5–11 cm long; leaflets 

nearly sessile, but not decurrent, 6–19 cm long, 5–15 cm wide, 3-segmented to halfway or nearly to the 

base; segments 4–12 cm long, 1.5–5.5 cm wide, mostly 3-lobed to the middle; lobes 2–5 cm long, 0.5–2.5 

cm wide, entire or with 1 or 2 teeth, segments, lobes, and teeth all acuminate at the apex. Flowers 3 or 4 

on each shoot, both terminal and axillary, with the terminal one blooming first, forming a cyme; pedicels 

slightly curved, 5–14 cm long, naked or with a leafy bract; involucrate bracts 4 or 5, green; sepals 3 or 4, 

grading into one another, all or all except one caudate at the apex; petals pure yellow, spreading, obovate 

rounded at the apex, 4–5.5 cm long, 2.5–3.5 cm wide; filaments yellow, 1.1–1.5 cm long, anthers yellow, 

c. 4 mm long; disk fleshy, 1 mm high, yellow, waved; carpels mostly single, very rarely 2; stigmas sessile, 

yellow. Follicles cylindrical, 4.7–7 cm long, 2–3.3 cm in diameter. Seeds kidney-shaped, dark brown, c. 1.5 

cm long, 1.2 cm in diameter. Figs 5.1A, 5.1B, 5.1C, 5.1D. 

phenology. Flowering from late May to early June and fruiting in September. 

chromosome number. 2n = 10 (Okada & Tamura, 1979; Li et al., 1998). 

habitat and distribution. Paeonia ludlowii was collected from sparse forests and thickets, on 

granites at altitudes of 2,870–3,450 m. This is a narrow endemic in southeastern Xizang (Tibet) and 

known from Nyingchi, Mailing, and Lhünzê counties at 28.4–29.9°N, 92.4–94.8°E. 

This species is also a medicinal plant and is often dug out by local people for its root bark. In 

three of the five populations studied (D. Y. Hong et al. H96007, H96014, H96030), hundreds of 

individuals were dug out by people from Gansu and Qinghai provinces (Hong, 1997a: fig. 2). This 

has caused a serious threat to the survival of this species. Effective measures must be taken to 

conserve this beautiful flower. Map 5.1. 

notes. In the description of Paeonia lutea var. ludlowii, Stern and Taylor (1951, 1953) indicated that 

the taxon was distinctly different from the variety lutea, distinguished by its long, commonly


1. PAEONIA LUDLOWII 

Map 5.1. Distribution of Paeonia ludlowii (Stern & G. Taylor) D. Y. Hong. 

unbranched stems to 8 feet (2.4 m) vs. 5 feet (1.5 m) in var. lutea, its larger and more open flowers, 

with up to 2 carpels that are twice as large as those of var. lutea. These differences have been 

confirmed upon examination of plants in five populations in Mailing and Nyingchi counties and 

five populations of var. lutea (= P. delavayi). As shown in Hong (1997a: figs 1 and 2), plants of P. 

ludlowii are tall from a caespitose base, and have relatively large, pure yellow flowers, yellow 

filaments, acuminate leaf segments and lobes, and typically one carpel per flower (more than 97% 

of the flowers examined had a single carpel and fewer than 3% had two). Furthermore, P. ludlowii 

produces very large follicles that contain the largest seeds in the genus. By contrast, plants of P. 

delavayi are not caespitose and have much shorter stems, acute leaf lobes and segments, smaller 

flowers, yellow petals that are nearly always red-blotched at the base, purple-red filaments, and 

three or four, rarely two, much smaller carpels. These differences clearly support the recognition of 

the variety ludlowii as a distinct taxon and species. 

Paeonia ludlowii is a tall shrub that often forms large and dense clumps with dozens of stems. A 

single individual can have up to 105 flowers (Hong, 1997a: fig. 2). All of the five populations studied 

were small in area, and the largest population was about 200 m in diameter. Except for the Quenima 

Village population (D. Y. Hong et al. H96020), which had only four individuals, all the populations 

observed consisted of many rather densely packed individuals, and the species was a dominant element 

in the community. Two factors may explain the small population areas that contain a large number of 

individuals. First, this species has a high seed-set, and its seeds appear to have a high germination rate. 

Nearly 100 seedlings were found in an area of a square meter under a large individual in the Nanyigou 

population (D. Y. Hong et al. H96030). Second, the seeds of P. ludlowii are large (ca. 1.2 cm diameter) 

and are not adapted to long-distance dispersal; perhaps they are mostly moved by rats. The species is 

Fig. 5.1A (opposite). Paeonia ludlowii (Stern & G. Taylor) D. Y. Hong: a, a shoot with a cyme and single carpels; b, a 

seedling, showing hypogeal seed germination. Drawn by Miss LI Ai-Li.


12. PAEONIA EMODI 

1 cm 

1 cm 

a 

1 cm 

c 

b 

d


11. PAEONIA EMODI 

121 

1 cm 

Fig. 5.12B. Paeonia emodi Wall. ex Royle: involucrate bracts and sepals, based on S. L. Zhou H01031 (PE). 

Drawn by Miss LI Ai-Li. 

P. sterniana in having nearly always multiple flowers per stem (rather than nearly always solitary) and 

the carpels mostly tomentose (rather than always glabrous). 

Saunders and Stebbins (1938) crossed Paeonia anomala and P. emodi, obtaining a very low seed 

set and sterile hybrids. This supported their observation that the two species are not only 

morphologically distinct but also reproductively isolated. 

additional specimens examined. CHINA, Xizang [Tibet], Gyirong County: Jiangcun Village, 

W slope, 28°18'N, 85°20'E, 2,480 m, in shrubs, 2 Aug. 2001, S. L. Zhou H01031 (A, BM, CAS, K, 

MO, P, PE); loc. eodem, in bushes, 2,500 m, 20 June 1975, Acad. Sin. Nat. Resources Exped. 75–247 

(PE); loc. eodem, E slope, 2,350 m, 6 June 1972, Xizang Chinese Traditional Medicinal Plant Exped. 413 

(PE). XINJIANG, Kashkaria, Brumhal Pass, May 1870, Henderson, Yarkand Exped. 1870 (LE). INDIA, 

Uttarakhand: Kumaon, Pithoragarh Bogdiar, 3,000 m, 29 Apr. 1965, N. C. Nair 35613 (CAL); loc. 

eodem, 2,000 m, 2 May 1966, N. C. Nair 36295 (CAL); loc. eodem, 3 May 1965, N. C. Nair 35634 

(CAL); Kumaon, Dwali, 2,438 m, 16 May 1848, R. Strachey & J. E. Winterbottom s.n. (CAL); loc. 

eodem, Naga Hills, 2,438 m, 18 June 1882, H. Collett 7 (CAL); Kumaon, Khati, 2,740 m, 10 June 

1919, N. Gill 829 (CAL); Kumaon, China Pahar, 2,286 m, May 1912, N. Gill 452 (CAL); Kumaon, 

China Peak, 2,286 m, 19 June 1913, N. Gill 531 (CAL); Kumaon, Pondur, 2,286 m, 16 May 1848, 

R. Strachey & J. E. Winterbottom s.n. (CAL); Kumaon, Rathi-Bogdwar, 2,000–3,000 m, 13 May 1958, 

T. A. Rao 6755 (CAL); Kumaon, 2,300 m, 1843, Strachey & Winterbottom 1 (P); Kumaon, W Almora, 

Pindari River, Biskam 2330 (E); Baba Pum, Reeckee, 9 June 1847, Winterbottom 180 (P); Garhwal 

Himalayas, Gangani, Uttarkashi, 1,800 m, 3 June 1982, B. S. Aswal 11983 (F, G); Garhwal, Sitapur, 

1,600 m, 21 May 1972, B. D. Naithani 47920 (CAL, G); Garhwal, Ramssi forest, 2,500 m, 10 June 

1959, M. A. Rau 10099 (CAL); Garhwal, Kanara to Mohankhal, 2,100 m, 27 Apr. 1963, C. L. 

Malhotra 27233 (CAL); Tehri-Garhwal, 2,100 m, May 1893, I. S. G. s.n. (G); Tehri-Garhwal, 

Ghuttu-Area, 1,800 m, 10 June 1972, B. D. Naithani 48250 (CAL, G); Tehri-Garhwal, Shanchatti, 

Fig. 5.12A (opposite). Paeonia emodi Wall. ex Royle: a, the lower part of the plant; b, stem with a lower leaf; c, carpels 

and disk; d, the upper surface of a leaf, showing bristles along veins. Drawn by Miss LI Ai-Li.

Peonies of the World - Look inside - Royal Botanic Gardens, Kew

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