A Monograph of the genus Microtoena (Lamiaceae) 9782759825295

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Current Natural Sciences

Qiang WANG

A Monograph of the Genus Microtoena (Lamiaceae)

This book was originally published by Science Press, © Science Press, 2018.

Qiang WANG State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Beijing Responsible Editors: Jing WANG and Haiguang WANG

Printed in France

EDP Sciences – ISBN(print): 978-2-7598-2528-8 – ISBN(ebook): 978-2-7598-2529-5 All rights relative to translation, adaptation and reproduction by any means whatsoever are reserved, worldwide. In accordance with the terms of paragraphs 2 and 3 of Article 41 of the French Act dated March 11, 1957, “copies or reproductions reserved strictly for private use and not intended for collective use” and, on the other hand, analyses and short quotations for example or illustrative purposes, are allowed. Otherwise, “any representation or reproduction – whether in full or in part – without the consent of the author or of his successors or assigns, is unlawful” (Article 40, paragraph 1). Any representation or reproduction, by any means whatsoever, will therefore be deemed an infringement of copyright punishable under Articles 425 and following of the French Penal Code. The printed edition is not for sale in mainland China. Customers in mainland China please order the print book from Science Press. ISBN of the China edition: Science Press ISBN: 978-7-03-056235-7 ©

Science Press, EDP Sciences, 2020

DEDICATION

Dedicated to the ‘father’ of Microtoena and ex-director (1905–1922) of the Royal Botanic Gardens, Kew: Dr. David Prain

PREFACE The genus Microtoena is an enigmatic member of the mint family. Most species of this genus are rare and poorly collected in the field. Although this rare plant is not widely cultivated in most gardens, the lovely flowers of it are certainly attractive, both to the taxonomists and to the general public. Recent studies on chemical components of Microtoena reveal its huge potential in Medicine. Obviously, for a genus of such horticultural and medicinal importance, it is essential to have a scientifically sound taxonomy. In the taxonomic history of Microtoena, more than thirty species and varieties have been described and reported. However, most of these taxa are poorly known, and the relationship between species was still unclear. The only taxonomic system of Microtoena was proposed about fifty years ago. Unfortunately, this taxonomic system with six series is not reliable, because it is mainly based on a few quantitative characters which are found to be unstable. I have worked on this enigmatic genus for more than ten years and have carefully investigated nearly all the distribution areas of Microtoena. Based on extensive field observations, population sampling, critical examination of herbarium specimens, scanning electron microscopy (SEM) observations, and statistical analysis of all relevant characters, as well as molecular phylogenetic analysis, two sections with nineteen species are recognized here for Microtoena. In addition, twenty- two names of species and varieties are reduced to synonyms, and one species is excluded from Microtoena. Based on molecular phylogenetic analyses and reconstructions of ancestral states of diagnostic characters as well as distribution patterns, I traced evolutionary trends of a number of characters of taxonomic significance, and also inferred the biogeographical history of Microtoena. It is to be hoped that the present monograph will stimulate the intensive study and conservation of these lovely plants. It is necessary for us to know such a rare, enigmatic, and important genus better, before we lose it. Qiang WANG Oct. 10, 2017

ACKNOWLEDGEMENTS I am indebted to numerous persons for their great help to my research. First, I especially send my sincere gratitude to Prof. De-Yuan HONG, Prof. Kai-Yu PAN, and Prof. Song GE of Institute of Botany, Chinese Academy of Sciences. I could not have completed this work without their kind encouragement, great suggestions, and financial support. I am also extremely grateful to my dear friend, Mike Gilbert of Royal Botanic Gardens, Kew, for his great suggestions and help in Nomenclature and English writing. I should send my special gratitude to Prof. Hong-Ya GU and Prof. Guang-Yuan RAO of Peking University, and to Prof. An-Min LU, Prof. Shi-Liang ZHOU, Prof. Xian-Chun ZHANG, and Prof. Xiang-Yun ZHU of Institute of Botany, Chinese Academy of Sciences, for their great suggestions on methodology, taxonomic treatments, and Nomenclature of the present monograph. My special thanks are due to Dr. Alan Paton and Dr. Gemma Bramley of Royal Botanic Gardens, Kew, for their helpful suggestions on my research. Grateful thanks to Dr. Tim Utteridge, the head of Identification & Naming of Royal Botanic Gardens, Kew, for hosting me to work in Kew Garden for one year. A special word of thanks to Dr. Chun-Lei XIANG of Kunming Institute of Botany, Chinese Academy of Sciences for providing the terrific photos of Microtoena robusta, and to Dr. You-Sheng CHEN and Dr. Bin LIU of Institute of Botany, Chinese Academy of Sciences, for providing the terrific photos of Microtoena delavayi and Microtoena wardii respectively. Special thanks to Mr. Yun-Xi ZHU for preparing all the lovely illustrations, which are scientifically accurate and aesthetically excellent. I am very grateful to Dr. Yan YU of Sichuan University in data processing, and to Mr. Xin-Tang MA for his great help in my field work. Thanks and acknowledgement are also due to my best friends and great backings in Institute of Botany, Chinese Academy of Sciences and Royal Botanic Gardens, Kew: Ai-Li LI, Andre Schuiteman, Clare Drinkell, Da-Ming ZHANG, Fu-Sheng YANG, Hong-Yao ZHANG, Hui-Yuan Liu, Laura Pearce, Laura Jennings, Lesley Walsingham, Li-Min LU, Li-Na ZHAO, Liang-Qian LI, Maria Alvarez, Min FENG, Min LI, Ning-Ning YU, Qin BAN, Qing-Hua WANG, Qing-Yin ZENG, Ran WEI, Ruth Clarke, Ren SA, Sally Dawson, Shu-Rong CHEN, Tian-Gang GAO, Wen-Li CHEN, Xiao-Guo XIANG, Xiao-Hua JIN, Xiao-Ling GONG, Xiao-Quan WANG, Yan CHEN, You-Sheng CHEN, Yu JIA, Yu-Fen DU, Zhe-Ping XU, Zhen-Yu LI, Zhi-Duan CHEN, Zhi-Rong YANG. My sincere thanks are due to directors, curators, and collection managers of the herbaria A, BM, CDBI, E, GH, HGAS, HITBC, HX, IBK, IBSC, K, KUN, LE, NAS, P, PE,

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A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

QTPMB, SM, SZ, W, WU and WUK for their permission to examine the specimens. Last, I am indebted to my wife Li-Ping ZHANG for her continuous support to my work. This research was financially supported by the National Natural Science Foundation of China (Grant Nos. 31200157, 31110103911, J1310002), and the National Science Infrastructure Platform Foundation of China (Grant No. 2013FY112100).

CONTENTS PREFACE ACKNOWLEDGEMENTS CHAPTER I. TAXONOMIC HISTORY AND QUESTIONS TO BE ADDRESSED ................................................................................................................. 1 TAXONOMIC HISTORY ......................................................................................... 1 CIRCUMSCRIPTION AND PHYLOGENETICAL POSITION .............................. 5 SUBDIVISION........................................................................................................... 6 SPECIES DELIMITATION ....................................................................................... 7 QUESTIONS TO BE ADDRESSED ......................................................................... 8 CHAPTER II. MORPHOLOGY ................................................................................. 9 ROOTS ....................................................................................................................... 9 STEMS ....................................................................................................................... 9 LEAVES ................................................................................................................... 11 LEAF LENGTH .......................................................................................................... 11 LEAF SHAPE ............................................................................................................ 12 LEAF MARGIN .......................................................................................................... 14 LEAF BASE .............................................................................................................. 15 LEAF SURFACE......................................................................................................... 16 INFLORESCENCE .................................................................................................. 26 BRACTS................................................................................................................... 27 BRACT LENGTH ........................................................................................................ 27 BRACT SHAPE .......................................................................................................... 29 CALYX .................................................................................................................... 30 CALYX LENGTH........................................................................................................ 31 CALYX SPLIT RATIO .................................................................................................. 33 CALYX TOOTH RATIO ............................................................................................... 34 COROLLA ............................................................................................................... 36 COROLLA COLOUR................................................................................................... 36 COROLLA LENGTH ................................................................................................... 38

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A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE) COROLLA TUBE RATIO .............................................................................................. 40 APEX OF THE UPPER LIP OF THE COROLLA .................................................................. 41 HAIRS OF THE UPPER LIP OF THE COROLLA ................................................................. 42 MIDDLE LOBE OF THE LOWER LIP OF THE COROLLA ..................................................... 43

STAMENS ............................................................................................................... 44 ANTHERS ................................................................................................................ 44 HAIRS ON THE FILAMENTS ........................................................................................ 44 STYLE ...................................................................................................................... 45 POLLEN ................................................................................................................... 45 FRUITS .................................................................................................................... 47 CHAPTER III. MOLECULAR PHYLOGENY, BIOGEOGRAPHY AND CHARACTER EVOLUTION .................................................................................... 48 PHYLOGENETICAL POSITION AND SUBDIVISION ....................................... 48 ORIGIN AND DIVERSIFICATION ....................................................................... 50 CHARACTER EVOLUTION .................................................................................. 51 CHAPTER IV. TAXONOMIC REVISION OF MICROTOENA ............................ 54 MICROTOENA PRAIN ............................................................................................ 54 KEY TO SECTIONS AND SPECIES OF MICROTOENA .................................... 55 SECTION 1. Microtoena...........................................................................................57

1. Microtoena esquirolii....................................................................................57 2. Microtoena insuavis.............................................................................................. 63 3. Microtoena patchoulii ...................................................................................68 4. Microtoena mollis .........................................................................................73 SECTION. 2. Delavayana .........................................................................................77

5. Microtoena delavayi......................................................................................77 6. Microtoena wardii.........................................................................................83 7. Microtoena urticifolia ...................................................................................86 8. Microtoena albescens....................................................................................88 9. Microtoena stenocalyx ..................................................................................90 10. Microtoena miyiensis ..................................................................................93 11. Microtoena megacalyx ................................................................................94 12. Microtoena moupinensis .............................................................................96 13. Microtoena prainiana ...............................................................................102 14. Microtoena muliensis ................................................................................105 15. Microtoena omeiensis ...............................................................................107

CONTENTS

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16. Microtoena brevipedunculata ...................................................................110 17. Microtoena nepalensis ..............................................................................112 18. Microtoena robusta ...................................................................................112 19. Microtoena vanchingshanensis............................................................. 116 EXCLUDED NAME .............................................................................................. 118 BIBLIOGRAPHY ..................................................................................................... 119 INDEX TO SPECIMENS CITED ........................................................................... 121 INDEX TO BOTANICAL NAMES AND SYNONYMS ....................................... 126 DATA OF STANDARD DEVIATION ANALYSIS............................................... 128

CHAPTER I. TAXONOMIC HISTORY AND QUESTIONS TO BE ADDRESSED TAXONOMIC HISTORY Before the establishment of Microtoena, some taxa of this genus had attracted the attention of taxonomists. The first species described was by Henry Fletcher Hance (1884) from Guangdong Province (China), as Gomphostemma insuave Hance. This was transferred to Microtoena by John Isaac Briquet (1895). Sir Joseph Dalton Hooker (1885) described Plectranthus patchouli C.B. Clarke ex Hook. f. from Assam (India). However, he was suspicious about the position of this new species. Charles Baron Clarke (1889) commented that P. patchouli might be a species of Cymaria or a close relative. David Prain (1889) renamed it as Microtoena cymosa Prain, but this name is illegitimate. Finally, this species was correctly transferred to Microtoena by Shwe Jye Hsuan (1965), as M. patchoulii (C.B. Clarke ex Hook. f.) C.Y. Wu & S.J. Hsuan. Adrien René Franchet (1887) described Clerodendrum moupinense Franch. from Sichuan Province (China). This species was later transferred to Microtoena by D. Prain (1895), as M. moupinensis (Franch.) Prain. The establishment of Microtoena was in 1889. Two years before 1889, David Prain, later director (1905-1922) of the Royal Botanic Gardens, Kew, visited India to investigate commercial plants. The ‘true Patchouli plant’ of Clarke attracted Prain’s attention, and he didn’t agree with Hooker and Clarke, and thought that it was unjustifiable to place this species in Plectranthus or Cymaria. In his opinion, this plant should belong to a new genus. In 1889, D. Prain founded his new genus Microtoena, and transferred P. patchouli to Microtoena. In accordance with the ‘Kew Rule’ about the priority of epithets being limited to their use only within a particular genus, which permitted changing an epithet when a species was transferred for the first time from one genus to another, D. Prain rejected the epithet ‘patchouli’ and renamed P. patchouli as M. cymosa, the first species name in Microtoena. The ‘Kew Rule’ has been rejected, and Prain’s renaming is now illegitimate. S.J. Hsuan (1965) transferred P. patchouli into Microtoena as M. patchoulii, and reduced the name M. cymosa to a synonym. William Botting Hemsley (1890) described two new species of Microtoena from

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A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

Hubei Province (China), based on the collections of Augustine Henry: M. robusta Hemsl. with robust stems and M. urticifolia Hemsl. with Urtica-like leaves. At the same year, D. Prain (1890) described M. griffithii Prain as new, based on two specimens collected from Assam (India). The lectotype of M. griffithii designated in the present study, Griffith 4059 [preserved in K (K 000928194)], might be the oldest collection of a Microtoena. It was collected by William Griffith in April 1863, approximately twenty-six years before the establishment of Microtoena. Five years later, D. Prain (1895) described another new species: M. delavayi Prain with two varieties, var. vera Prain and var. grandiflora Prain. The name M. delavayi var. vera is invalid (Art. 24. 3), and the autonym M. delavayi var. delavayi should be accepted. In addition, D. Prain transferred Clerodendrum moupinense to Microtoena, as M. moupinensis (Franch.) Prain. In the same year, J.I. Briquet (1895) regarded Gomphostemma insuave Hance as conspecific with M. cymosa. Therefore, he made the new combination, M. insuavis (Hance) Prain ex Briq., and reduced M. cymosa to synonymy. Friedrich Ludwig Emil Diels (1900) described a new species from China, M. prainiana Diels. This species was collected from Jinfo Mountain, Nanchuan county, Chongqing City (China). This distinct species has glomerate cymes in ovoid panicles. It has also been found in other provinces of SW China, such as Sichuan, Yunnan, and Guizhou. Augustine Abel Hector Léveillé (1911) described three new species: M. mollis H. Lév., M. esquirolii H. Lév., and M. coreana H. Lév.. Both M. mollis and M. esquirolii were collected from Guizhou Province (China), but M. coreana from Seoul City (South Korea) was far from SW China and SE Asia, the main distribution area of Microtoena. A.A.H. Léveillé was suspicious about this remote species, and marked it as a doubtful member of the genus. Unaware of the combination of J.I. Briquet (1895) for Gomphostemma insuave, Stephen Troyte Dunn (1913) transferred G. insuave to Microtoena again, and combined it as M. insuavis (Hance) Prain ex Dunn, which is certainly illegitimate. In addition, S.T. Dunn regarded M. cymosa, M. mollis, and M. esquirolii as conspecific with M. insuavis, and reduced these names to synonyms. Heinrich Raphael Eduard von Handel-Mazzetti (1936) described a new species and a new variety from China: M. maireana Hand.-Mazz. narrowly distributed in Huize County of Yunnan Province and M. urticifolia var. subedentata Hand.-Mazz., a poorly known taxon endemic to Nu Jiang region of Yunnan Province. Cheng Yi Wu (1959) made the first taxonomic revision of Microtoena in China. In his revision, nine species and two varieties were recognized. C.Y. Wu followed S.T. Dunn (1913) and reduced M. cymosa, M. mollis, and M. esquirolii to synonyms of M. insuavis, and the invalid name M. delavayi var. vera to a synonym of M. delavayi. In addition, C.Y. Wu described two new species of Microtoena, M. tenuiflora C.Y. Wu

CHAPTER I. TAXONOMIC HISTORY AND QUESTIONS TO BE ADDRESSED

3

with slim flowers and M. megacalyx C.Y. Wu with a conspicuously dilated calyx. Following the revision of C.Y. Wu, S.J. Hsuan (1965) made an updated taxonomic revision of Microtoena for China. In this revision, S.J. Hsuan added nine new species and four new varieties as follows: M. pauciflora C.Y. Wu, M. subspicata C.Y. Wu, M. subspicata var. intermedia C.Y. Wu & S.J. Hsuan, M. delavayi var. lutea C.Y. Wu & S.J. Hsuan, M. delavayi var. amblyodon C.Y. Wu & S.J. Hsuan, M. affinis C.Y. Wu & S.J. Hsuan, M. albescens C.Y. Wu & S.J. Hsuan, M. muliensis C.Y. Wu & S.J. Hsuan, M. stenocalyx C.Y. Wu & S.J. Hsuan, M. omeiensis C.Y. Wu & S.J. Hsuan, M. vanchingshanensis C.Y. Wu & S.J. Hsuan, M. urticifolia var. brevipedunculata C.Y. Wu & S.J. Hsuan, and M. longisepala C.Y. Wu. In addition, S.J. Hsuan made a new combination, M. patchouli (C.B. Clarke ex Hook. f.) C.Y. Wu & S.J. Hsuan (formerly Plectranthus patchouli and Microtoena cymosa), and separated it from M. insuavis as an independent species. Unfortunately, S.J. Hsuan was still unaware of Briquet’s combination for G. insuave (J.I. Briquet, 1895), and took up the later illegitimate combination made by S.T. Dunn (1913), M. insuavis (Hance) Prain ex Dunn. S.J. Hsuan did not agree with S.T. Dunn on the reduction of M. mollis to a synonym of M. insuavis, and restored it as an independent species. She reduced M. tenuiflora to a synonym of M. delavayi. In Hsuan’s revision, a taxonomic system with five series for Chinese Microtoena was proposed, which was mainly based on the morphology of the calyx. Hsi Wen Li and C.Y. Wu (1977) described a monospecific series, ser. Miyienses C.Y. Wu & H.W. Li based on their new species M. miyiensis C.Y. Wu & H.W. Li. In addition, C.Y. Wu and H.W. Li separated M. maireana from ser. Stenocalyces C.Y. Wu & S.J. Hsuan, and transferred it into their new series. William Thomas Stearn (1982) described a new species from Nepal, Microtoena nepalensis Stearn, which has extremely tiny bracts. One year later, W.T. Stearn (1983) described three further new species of Microtoena from East Himalayan: M. bhutanica Stearn endemic to Bhutan, M. siamica Stearn endemic to Thailand, and M. wardii Stearn endemic to S Tibet. In the Flora of China vol. 17, H.W. Li and Ian Charleson Hedge (1994) accepted 20 species and 5 varieties of Microtoena in China, mainly based on the revision of Hsuan (1965). In addition, they accepted the legitimate combination name M. insuavis (Hance) Prain ex Briq. for G. insuave. However, they improperly included its homonym M. insuavis (Hance) Prain ex Dunn as a synonym of M. patchoulii. Based on specimen examination, extensive field observation, and careful analysis of a number of characters, De Yuan Hong and I (Wang & Hong, 2011) made a taxonomic revision on the most complicated group of Microtoena, the M. insuavis complex. In this revision, we confirmed the separation of M. patchoulii and M. insuavis, and restored M. esquirolii as independent species. In addition, we reduced M. subspicata and M. subspicata var. intermedia to synonyms of M. esquirolii, M.

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pauciflora to a synonym of M. patchoulii, and M. siamica to a synonym of M. insuavis respectively. In 2012, D.Y. Hong & I dealt with the identity of M. affinis, a species which has been extremely doubtful (Li & Hedge, 1994). Based on field investigations of the type localities of M. affinis and allied species, specimen examination, and statistical analysis of characters, it was reduced to a synonym of M. delavayi (Wang & Hong, 2012). In 2015, I expounded the identity of M. coreana (Wang, 2015), a species which has long been extremely doubtful. A.A.H. Léveillé (1911), who described this species, was suspicious about its position. C.Y. Wu (1959) and S.J. Hsuan (1965) both thought that M. coreana should be excluded from Microtoena. My results showed that M. coreana was actually an element of Tripora divaricata (Maxim.) P.D. Cantino (formerly Caryopteris divaricata Maxim.), a species which was long placed in the family Verbenaceae. As a result, I excluded M. coreana from the genus Microtoena, and reduced it to a synonym of T. divaricata. All the species and variety names of Microtoena referred above are summarized in the Table I-1. Table I-1. Chronology of the genus Microtoena. Authors (year)

new taxa recognized

Accepted name (the present study)

Hance (1884)

Gomphostemma insuave Hance

M. insuavis (Hance) Prain ex Briq.

Hooker (1885)

Plectranthus patchouli C.B. Clarke ex Hook. f. M. patchoulii (C.B. Clarke ex Hook. f.) C.Y. Wu & S.J. Hsuan

Franchet (1887)

Clerodendrum moupinense Franch.

M. moupinensis (Franch.) Prain

Prain (1889)

Microtoena cymosa Prain nom. illeg.

M. patchoulii (C.B. Clarke ex Hook. f.) C.Y. Wu & S.J. Hsuan

Hemsley (1890)

M. robusta Hemsl. M. urticifolia Hemsl.

Accepted Accepted

Prain (1890)

M. griffithii Prain

M. moupinensis (Franch.) Prain

Prain (1895)

M. moupinensis (Franch.) Prain M. delavayi Prain M. delavayi var. vera Prain nom. inval. M. delavayi var. grandiflora Prain

Accepted Accepted M. delavayi Prain M. delavayi Prain

Briquet (1895)

M. insuavis (Hance) Prain ex Briq.

Accepted

Diels (1900)

M. prainiana Diels

Accepted

Léveillé (1911)

M. mollis H. Lév. M. esquirolii H. Lév. M. coreana H. Lév.

Accepted Accepted Tripora divaricata (Maxim.) P.D. Cantino

Dunn (1913)

M. insuavis (Hance) Prain ex Dunn. nom. illeg. M. insuavis (Hance) Prain ex Briq.

Handel-Mazzetti (1936)

M. maireana Hand.-Mazz. M. urticifolia var. subedentata Hand.-Mazz.

M. moupinensis (Franch.) Prain M. wardii Stearn

Wu (1959)

M. tenuiflora C.Y. Wu M. megacalyx C.Y. Wu

M. delavayi Prain Accepted

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continued Authors (year)

new taxa recognized

Hsuan (1965)

M. pauciflora C.Y. Wu ex S.J. Hsuan

Accepted name (the present study)

Wu & Li (1977)

M. miyiensis C.Y. Wu & H.W. Li

Accepted

Stearn (1982)

M. nepalensis Stearn

Accepted

Stearn (1983)

M. bhutanica Stearn M. siamica Stearn M. wardii Stearn

M. delavayi Prain M. insuavis (Hance) Prain ex Briq. Accepted

M. patchoulii (C.B. Clarke ex Hook. f.) C.Y. Wu & S.J. Hsuan M. subspicata C.Y. Wu ex S.J. Hsuan M. esquirolii H. Lév. M. subspicata var. intermedia C.Y. Wu ex S.J. M. esquirolii H. Lév. Hsuan M. delavayi var. lutea C.Y. Wu & S.J. Hsuan M. wardii Stearn M. delavayi var. amblyodon C.Y. Wu & S.J. M. wardii Stearn Hsuan M. affinis C.Y. Wu & S.J. Hsuan M. delavayi Prain M. albescens C.Y. Wu & S.J. Hsuan Accepted M. stenocalyx C.Y. Wu & S.J. Hsuan Accepted M. muliensis C.Y. Wu ex S.J. Hsuan Accepted M. omeiensis C.Y. Wu & S.J. Hsuan Accepted M. vanchingshanensis C.Y. Wu & S.J. Hsuan Accepted M. urticifolia var. brevipedunculata C.Y. Wu & M. brevipedunculata (C.Y. Wu & S.J. Hsuan) Q. S.J. Hsuan Wang M. longisepala C.Y. Wu ex S.J. Hsuan M. moupinensis (Franch.) Prain M. patchoulii (C.B. Clarke ex Hook. f.) C.Y. Accepted Wu & S.J. Hsuan

CIRCUMSCRIPTION AND PHYLOGENETICAL POSITION In the taxonomic history of Microtoena, only four generic names have been involved in its circumscription, including Clerodendrum, Gomphostemma, Plectranthus, and Tripora. Before the establishment of Microtoena, three species of Microtoena had been described under the genera Clerodendrum, Gomphostemma, and Plectranthus respectively. These misplaced species were transferred to Microtoena soon after its establishment. Since then, no species of Microtoena has been described under any other genus. The last misplaced species, Microtoena coreana, was incorrectly included within Microtoena, and has been reduced to a synonym of Tripora divaricata. Obviously, in the taxonomic history, the circumscription of Microtoena is stable and without problem. In practice, Microtoena is a quite distinct genus. It can be easily recognized in the field by its dichotomous cymes and the remarkable upper lip of the corolla, which is laterally compressed galeate. When founding the genus Microtoena, D. Prain (1889) thought his new genus was closely related to the genus Craniotome, so he transposed Craniotome into Microtoena as the name for this new genus. Therefore, the name Microtoena is actually a meaningless anagram, which was strongly criticized by Richard Thomas Lowe (1868). D. Prain (1889) placed his Microtoena in tribe Stachydeae, the fifth tribe in George Bentham’s eight-tribe taxonomic system for Lamiaceae (Bentham, 1876). J.I. Briquet

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(1897) proposed a widely accepted taxonomic system of Lamiaceae comprising eight subfamilies, and placed Microtoena and Craniotome in the subtribe Lamiinae, tribe Stachydeae of his sixth subfamily Stachyoideae. In the ‘The Families and Genera of Vascular Plants-VII Lamiales’ (Kadereit, 2004), Joachim Walter Kadereit placed Microtoena in subfamily Lamioideae (Harley, 2003). Recent molecular phylogenetic studies (Scheen et al., 2010; Bendiksby et al., 2011) presented new evidence for the circumscription and phylogenetical position of Microtoena. Anne Cathrine Scheen et al. (2010) sampled only one species (M. patchoulii) from Microtoena. The results of their analysis based on three chloroplast DNA fragments (trnL, trnL-F, rps16) showed that Microtoena formed a clade with Craniotome, Anisomeles and Pogostemon, which nested in the tribe Pogostemoneae, subfamily Lamioideae. Based on the study of A.C. Scheen et al., Mika Bendiksby et al. (2011) added another chloroplast marker (matK) in their analysis. Besides M. patchoulii, M. Bendiksby sampled another species, M. delavayi. The result confirmed the relationship of Microtoena, Craniotome, Anisomeles and Pogostemon, and the position of Microtoena in Lamiaceae. Unfortunately, the results of M. Bendiksby et al. showed that the two species of Microtoena might be paraphyletic, although their resolution was weakly supported (JK < 50) and unclear. For the first time, the circumscription of this genus was challenged.

SUBDIVISION The genus Microtoena is a poorly known group, most species of which are extremely rare in the field. The infrequence of Microtoena has lead to poor taxonomic study of Microtoena and no integrated subdivision has been proposed. S.J. Hsuan was the first taxonomist to subdivide Microtoena. In her taxonomic revision of Microtoena in China (Hsuan, 1965), She recognized five series for the Chinese species of Microtoena: ser. Cymosae; ser. Robustae; ser. Stenocalyces; ser. Urticifoliae, and ser. Vanchingshanenses. This system was supplemented by C.Y. Wu and H.W. Li (1977), who added ser. Miyienses to make a total of six series (Table I-2). The six-series system is mainly based on quantitative parameters of the calyx, such as the length of the calyx, and the length and width of the calyx teeth. However, the calyx and calyx teeth undergo continuous enlargement period after anthesis. There is a great risk to base a taxonomic system on these unstable quantitative characters. Table I-2. The six-series taxonomic system of Microtoena. Series name

Taxa included

ser. Cymosae (incl. 5 species & 1 variety)

M. patchoulii, M. pauciflora, M. mollis, M. subspicata, M. subspicata var. intermedia, M. insuavis

ser. Miyienses (monospecific series)

M. miyiensis

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continued Series name

Taxa included

ser. Robustae (monospecific series)

M. robusta

ser. Stenocalyces (incl. 7 species & 3 variety)

M. delavayi, M. delavayi var. grandiflora, M. delavayi var. lutea, M. delavayi var. amblyodon, M. muliensis, M. omeiensis, M. stenocalyx, M. affinis, M. albescens, M. maireana

ser. Urticifoliae (incl. 5 species & 1 variety)

M. urticifolia, M. urticifolia var. brevipedunculata, M. moupinensis, M. prainiana, M. longisepala, M. megacalyx

ser. Vanchingshanenses (monospecific series)

M. vanchingshanensis

SPECIES DELIMITATION Species delimitation is an eternal topic in taxonomy. For Microtoena, because of the poor collections and insufficient field observation, the variation ranges of characters were poorly understood in the existing taxonomic studies. Some quantitative characters, such as corolla length, flower number, and leaf length, are excessively emphasized as diagnostic in species delimitation. As a result, many species of Microtoena are questionable and need to be rechecked. Two problematic species are mentioned here as examples. The first one is M. pauciflora, which was described by C.Y. Wu and S.J. Hsuan based on only one collection (Hsuan, 1965) from Yunnan Province (China). S.J. Hsuan (1965) took the view that it was closely related to M. patchoulii, another sympatric species. H.W. Li and I.C. Hedge (1994) accepted this species in the Flora of China, but made the remark: “This species may only be a variety of Microtoena patchoulii”. Based on field investigations, examination of specimens, and statistical analysis of characters, this species is actually conspecific with M. patchoulii (Wang & Hong, 2011). The other problematic species is M. affinis. This species was also described by C.Y. Wu and S.J. Hsuan (1965) based on only one collection, which is from Xundian County of Yunnan Province (China), within the distribution range of M. delavayi. C.Y. Wu remarked on the slip of the holotype that “this species is similar to M. delavayi, and its diagnostic characters are uniflorous axillary cymes, small leaf blades, and remotely crenate leaf margin”. Unfortunately, after checking the holotype carefully, we found that the “uniflorous axillary cyme” of M. affinis was actually five-flowered (Wang & Hong, 2012). H.W. Li and I.C. Hedge (1994) accepted this species in Flora of China, but indicated that: “This species may be a form of Microtoena delavayi”. This problematic species has also been reduced to a synonym of M. delavayi based on our careful clarification (Wang & Hong, 2012). In fact, the two examples above are just the tip of the iceberg, and more problematic species of Microtoena should be rechecked.

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A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

QUESTIONS TO BE ADDRESSED In summary, Microtoena is a poorly known genus with a number of taxonomic problems. Three major questions about this genus are as follows: 1. Is Microtoena monophyletic? 2. What is the intrageneric structure of Microtoena? 3. How many species could be recognized in Microtoena? The aim of the present study is to address these questions of this lovely genus, based on extensive field observation, specimen examination, statistical analysis of characters, and molecular phylogeny.

CHAPTER II. MORPHOLOGY ROOTS No special trait has been observed in the root system of Microtoena, such as tubercle or branch thickening which is common in other genera (Stachys, Salvia, Phlomis et al.) of Lamiaceae. The root of Microtoena is straight and simple. In the present study, adventitious roots of Microtoena are observed for the first time. For most species of Microtoena, adventitious roots (Figure II-1A) and buds (Figure II-1B) occur on the nodes or internodes of lodged stems, which may grow up to new individuals after the decomposition of the old lodged stem.

Figure II-1. The adventitious root and bud of M. esquirolii (Guangxi Province, China): A. a new shoot with adventitious roots on a lodged stem, scale bar = 2 cm; B. an adventitious bud on a lodged stem, scale bar =5 mm.

STEMS The stem of Microtoena is usually erect and woody at base, and the height is from 0.5 m to 2 m. A prostrate stem base can be observed in some species, such as M. moupinensis, M. stenocalyx, and M. vanchingshanensis. The stem is usually green, but sometimes

10

A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

light purple or green with purple spots. In addition, old perennial stems are sometimes lodged with adventitious roots and buds, as mentioned above. The indumentum on stems is diverse in Microtoena. Four types can be recognized as follow: 1. sparsely puberulent type with short fine hairs (Figure II-2A); 2. densely hirsute type with spreading (sometimes appressed) short hairs (Figure II-2B); 3. densely hispid type with long spreading stiff hairs (Figure II-2C); and 4. rather densely hispid type with long and short spreading stiff hairs (Figure II-2D).

Figure II-2. The stem indumentum of Microtoena: A, sparsely puberulent type in M. esquirolii (by Q. Wang in Longlin County, Guangxi Province, China); B, densely hirsute in M. insuavis (by Q. Wang in Gengma County, Yunnan Province, China); C, densely hispid in M. mollis (by Q. Wang in Mile County, Yunnan Province, China); D, rather densely hispid in M. patchoulii (by Q. Wang in Tengchong County, Yunnan Province, China). Scale bar = 5 mm.

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11

LEAVES LEAF LENGTH The quantitative characters of Microtoena have never been statistically evaluated in the previous taxonomic studies. For the leaf length, which was emphasized as diagnostic for M. affinis, M. pauciflora, and M. albescens (Wu, 1959; Hsuan, 1965), the variation range of each species is still unclear. In the present study, the variation range of all the quantitative characters involved are evaluated based on statistical analysis, both at population and at species level. All the quantitative characters evaluated are photographed with a scale bar, and measured by beta version of NucType developed by Yan Yu of the College of Life Sciences, Sichuan University. The data is accurate to two decimal places, and the average and standard deviation are calculated and accurate to three decimal places. For the leaf length, fourteen populations of five species are sampled for analysis at population level (Figure II-3). For each species sampled, the result shows that the

Figure II-3. The variation of leaf length in sampled populations of Microtoena: 1. M. esquirolii, D. Y. Hong & Q. Wang, H&W08006 (Longlin, Guangxi, China); 2. M. esquirolii, D.Y. Hong & Q. Wang, H&W08007 (Longlin, Guangxi, China); 3. M. insuavis, D. Y. Hong & Q. Wang, H&W09001 (Gengma, Yunnan, China); 4. M. insuavis, D. Y. Hong & Q. Wang, H&W09002 (Gengma, Yunnan, China); 5. M. mollis, D. Y. Hong & Q. Wang, H&W08004 (Mile, Yunnan, China); 6. M. mollis, D. Y. Hong & Q. Wang, H&W08005 (Mile, Yunnan, China); 7. M. mollis, D. Y. Hong & Q. Wang, H&W09003 (Mengla, Yunnan, China); 8. M. mollis, D. Y. Hong & Q. Wang, H&W09004 (Mengla, Yunnan, China); 9. M. mollis, D. Y. Hong & Q. Wang, H&W09006 (Mile, Yunnan, China); 10. M. moupinensis, D. Y. Hong & Q. Wang, H&W09007 (Baoxing, Sichuan, China); 11. M. moupinensis, D. Y. Hong & Q. Wang, H&W09008 (Baoxing, Sichuan, China); 12. M. patchoulii, D. Y. Hong & Q. Wang, H&W08009 (Tengchong, Yunnan, China); 13. M. patchoulii, D. Y. Hong & Q. Wang, H&W09010 (Jingdong, Yunnan, China); 14. M. patchoulii, D. Y. Hong & Q. Wang, H&W09011 (Jingdong, Yunnan, China).

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A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

variation range of leaf length is continuous between different populations. The leaf length is highly variable in some population such as population H&W09004 of M. mollis, but the variation range of different populations overlaps each other within the same species. The result of analysis in species level shows a great plasticity of leaf length in most species of Microtoena. In most cases, it is inappropriate to base a new species on such an unstable character. For example, M. pauciflora and M. affinis, the short leaves of which were emphasized as diagnostic (Hsuan, 1965), have been reduced to a synonym of M. patchoulii (Wang & Hong, 2011) and M. delavayi (Wang & Hong, 2012) respectively. The present statistical analysis (Figure II-4) reveals the wide variation ranges of leaf length of M. insuavis, M. patchoulii, M. mollis, M. delavayi, and M. moupinensis. In addition, the averages from the holotypes are listed here for two species (M. albescens & M. miyiensis) without other collections. Further field investigations are badly needed to establish the variation range of the two species.

Figure II-4. The variation of leaf length of Microtoena in species level: 1. M. esquirolii; 2. M. insuavis; 3. M. patchoulii; 4. M. mollis; 5. M. delavayi; 6. M. wardii; 7. M. urticifolia; 8. M. albescens; 9. M. stenocalyx; 10. M. miyiensis; 11. M. megacalyx; 12. M. moupinensis; 13. M. prainiana; 14. M. muliensis; 15. M. omeiensis; 16. M. brevipedunculata; 17. M. nepalensis; 18. M. robusta; 19. M. vanchingshanensis.

LEAF SHAPE The leaf shape of Microtoena is usually described as oblong-ovate, ovate and broadly ovate, which are correct, but the line of demarcation of each state is very unclear. In the

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13

present study, the index LR (leaf ratio) is introduced to evaluate this character statistically. The formula to calculate the leaf ratio is as follows: LR (leaf ratio) = LW (leaf width) / LL (leaf length). The statistical analysis of leaf shape of sampled populations shows that the variation range of leaf shape is continuous within species (Figure II-5). The leaf shape of the two populations of M. moupinensis are highly variable, with leaf ratios from 0.52 to 0.84 and high standard deviation (more than 0.60). The leaf shape of the two populations of M. esquirolii is relatively stable. The population H&W08006 has leaf ratio of 0.634 in average with standard deviation of 0.019, and the population H&W08007 has leaf ratio of 0.663 in average with standard deviation of 0.018. For populations of M. mollis, the results show that the variation range of three populations (H&W08004, H&W08005, H&W09006) from Mile County is wider than that of the two populations (H&W09003, H&W09004) from Mengla County, but the variation range is still continuous within species.

Figure II-5. The variation of leaf shape in sampled populations of Microtoena: 1. M. esquirolii, D.Y. Hong & Q. Wang, H&W08006 (Longlin, Guangxi, China); 2. M. esquirolii, D.Y. Hong & Q. Wang, H&W08007 (Longlin, Guangxi, China); 3. M. insuavis, D.Y. Hong & Q. Wang, H&W09001 (Gengma, Yunnan, China); 4. M. insuavis, D.Y. Hong & Q. Wang, H&W09002 (Gengma, Yunnan, China); 5. M. mollis, D.Y. Hong & Q. Wang, H&W08004 (Mile, Yunnan, China); 6. M. mollis, D.Y. Hong & Q. Wang, H&W08005 (Mile, Yunnan, China); 7. M. mollis, D.Y. Hong & Q. Wang, H&W09003 (Mengla, Yunnan, China); 8. M. mollis, D.Y. Hong & Q. Wang, H&W09004 (Mengla, Yunnan, China); 9. M. mollis, D.Y. Hong & Q. Wang, H&W09006 (Mile, Yunnan, China); 10, M. moupinensis. D.Y. Hong & Q. Wang, H&W09007 (Baoxing, Sichuan, China); 11, M. moupinensis. D.Y. Hong & Q. Wang, H&W09008 (Baoxing, Sichuan, China); 12. M. patchoulii, D.Y. Hong & Q. Wang, H&W08009 (Tengchong, Yunnan, China); 13. M. patchoulii, D.Y. Hong & Q. Wang, H&W09010 (Jingdong, Yunnan, China); 14. M. patchoulii, D.Y. Hong & Q. Wang, H&W09011 (Jingdong, Yunnan, China).

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A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

The analysis in species level (Figure II-6) reveals that the leaf ratio of Microtoena is almost around 0.6, which is corresponding to the common description ‘ovate’ for most species of Microtoena. The lowest leaf ratio is observed in M. muliensis (0.476 in average with the standard deviation of 0.067), the leaves of which are usually described as oblong-ovate. The widest variation range is observed in M. moupinensis, the leaf ratio of which is from 0.52 to 0.89. According to Flora of China vol. 17 (Li & Hedge, 1994), the leaf shape of M. moupinensis is from ovate-cordate to triangular-ovate. The present analysis reveals a high standard deviation of 0.084 for M. moupinensis, which supports the instability in its leaf shape. Again, the averages from the holotypes are listed here for M. albescens (LR=0.78) and M. miyiensis (LR=0.68), the leaf shape of which is usually described as broadly ovate and rhombic-ovate respectively. More collections are badly needed to establish the variation range of the two species.

Figure II-6. The variation of leaf shape of Microtoena in species level: 1. M. esquirolii; 2. M. insuavis; 3. M. patchoulii; 4. M. mollis; 5. M. delavayi; 6. M. wardii; 7. M. urticifolia; 8. M. albescens; 9. M. stenocalyx; 10. M. miyiensis; 11. M. megacalyx; 12. M. moupinensis; 13. M. prainiana; 14. M. muliensis; 15. M. omeiensis; 16. M. brevipedunculata; 17. M. nepalensis; 18. M. robusta; 19. M. vanchingshanensis.

LEAF MARGIN Three types of leaf margin can be clearly recognized for Microtoena: dentate without any mucro (Figure II-7A); triangular-ovate serrate with mucrones (Figure II-7B); crenate with distinct mucrones (Figure II-7C). The ‘dentate’ type can usually be seen in M. esquirolii and M. prainiana; the ‘serrate’ type can be seen in M. patchoulii and M.

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15

insuavis, and the ‘crenate’ type can be seen in M. mollis and M. delavayi.

Figure II-7. The types of leaf margin of Microtoena, scale bar = 2 mm: A. dentate margin without any mucro; B. triangular-ovate serrate margin with mucrones; C crenate margin with distinct mucrones.

In addition, for some species, such as M. delavayi, the mature leaf margin is distinctly different from that of young leaves from new shoots (Figure II-8). However, for M. esquirolii and M. mollis, the leaf margin of young leaves is similar to the mature one.

Figure II-8. Variation of leaves within M. delavayi, scale bar = 1 cm: A. a small and remotely crenate leaf from a new shoot; B. the large and densely serrate leaf from a main stem.

LEAF BASE Three types of leaf base can be well recognized in Microtoena. The most common type

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A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

is the truncate-subcordate base (Figure II-9A), which is represented by most species, such as M. delavayi, M. wardii and M. omeiensis. The second type, the cuneate base (Figure II-9B), can be observed in some species, such as M. esquirolii, M. miyiensis and M. muliensis. The third type is the typical ‘cordate’ base (Figure II-9C) which can be found in M. mollis and M. vanchingshanensis.

Figure II-9. The leaf base of Microtoena, scale bar = 1cm: A. truncate-subcordate base of M. delavayi; B. cuneate base of M. esquirolii; C. cordate base of M. mollis.

Cultivation experiments (in the greenhouse of State Key Laboratory of Systematic and Evolutionary Botany (LSEB), Beijing, China) reveal that the leaf base is stable in different developing phases. In addition, no significant difference between wild and cultivated individuals was observed (Figure II-10). The species M. moupinensis should be mentioned here, which is special in having all three types of leaf base, even within the same individual.

Figure II-10. The leaf base of cultivated young plants: A. the truncate-subcordate base of M. delavayi, seeds from D.Y. Hong & Q. Wang, H&W08008 (Tengchong, Yunnan, China); B. the cuneate base of M. esquirolii, seeds from D.Y. Hong & Q. Wang, H&W08006 (Longlin, Guangxi, China).

LEAF SURFACE In the present study, micro-morphological characters of the leaf surfaces were observed

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17

under a Hitachi-S-4800 scanning electron microscope. Under the microscope, the main attention was paid to the hair density, hair type, hair ornamentation, and stomata of the sampled species, for both leaf surfaces. The results show that the hair density (Figures II-11–12), hair type (Figures II-13–14), hair ornamentation (Figures II-15–16), and stomata (Figures II-17–18) are diverse in Microtoena. For the sampled species, sparse hairs can be observed in M. esquirolii, on both adaxial (Figure II-11H–I) and abaxial surfaces (Figure II-12H–I). For M. delavayi, sparse hairs are observed on the adaxial surface (Figure II-11A–B), but dense hairs can be observed on the abaxial surface (Figure II-12A–B). For the other three species sampled, the hairs on both surfaces are always dense, but the density on the abaxial surface (Figure II-12C–G) is obviously higher than that on the adaxial surface (Figure II-11C–G). In addition, no significant difference in hair density was observed between populations of the same species. For the hair type, hispid and glandular hairs are common in the sampled species, but the number of cells of each hair is slightly different. In addition, villous hairs are observed in M. insuavis (Figure II-11C & Figure II-12C). Two types of hair ornamentation are observed in the sampled species. The first type, granular surface (Figure II-15H–I & Figure II-16H–I), is represented by the two samples of M. esquirolii. The second one is the smooth surface, which is common in other species sampled. The stomata index [= stomatal apparatuses per unit area / (stomatal apparatuses per unit area + ordinary cells per unit area)] is used here to reflect the stomata density. This index is less than 20% (Table II-1) in all the species sampled, and the index of M. delavayi and M. insuavis is obviously lower than that of the other three species (Figure II-17). The stomata of Microtoena is anomocytic without subsidiary cells (Figure II-18), and its guard cells are usually elliptical or subcircular. Table II-1. Stomata analysis of sampled species of Microtoena. Taxa

Stomata size (μm)

Stomatal index (%)

M. delavayi (817)

20.81×15.65

9.59

M. delavayi (H&W08008)

21.63×14.38

11.61

M. insuavis (H&W09001)

19.47×15.37

9.09

M. mollis (H&W08004)

18.65×17.48

19.16

M. mollis (H&W09003)

18.53×16.24

19.37

M. patchoulii (H&W09010)

23.58×16.63

16.75

M. patchoulii (580981)

21.13×15.76

17.55

M. esquirolii (H&W08006)

21.36×17.43

15.79

M. esquirolii (368)

20.42×17.54

16.24

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A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

Figure II-11. The hair density of the adaxial leaf surface of sampled species, scale bar = 500 μm: A. M. delavayi, Y. B. Zhang 817 (Xundian, Yunnan, China, holotype of M. affinis = M. delavayi); B. M. delavayi, Q. Wang H&W08008 (Tengchong, Yunnan, China); C. M. insuavis, Q. Wang H&W09001 (Gengma, Yunnan, China); D. M. mollis, Q. Wang H&W08004 (Mile, Yunnan, China); E. M. mollis, Q. Wang H&W09003 (Mengla, Yunnan, China); F. M. patchoulii, Q. Wang H&W09010 (Jingdong, Yunnan, China); G. M. patchoulii, R. L. Xiong & S. K. Wen 580981 (Tengchong, Yunnan, China, holotype of M. pauciflora = M. patchoulii); H. M. esquirolii, Q. Wang H&W08006 (Longlin, Guangxi, China); I. M. esquirolii, S. G. Wu 368 (Xinping, Yunnan, China, lectotype of M. subspicata var. intermedia = M. esquirolii).

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Figure II-12. The hair density of the abaxial leaf surface of sampled species, scale bar = 500 μm: A. M. delavayi, Y. B. Zhang 817 (Xundian, Yunnan, China, holotype of M. affinis = M. delavayi); B. M. delavayi, Q. Wang H&W08008 (Tengchong, Yunnan, China); C. M. insuavis, Q. Wang H&W09001 (Gengma, Yunnan, China); D. M. mollis, Q. Wang H&W08004 (Mile, Yunnan, China); E. M. mollis, Q. Wang H&W09003 (Mengla, Yunnan, China); F. M. patchoulii, Q. Wang H&W09010 (Jingdong, Yunnan, China); G. M. patchoulii, R. L. Xiong & S. K. Wen 580981 (Tengchong, Yunnan, China, holotype of M. pauciflora = M. patchoulii); H. M. esquirolii, Q. Wang H&W08006 (Longlin, Guangxi, China); I. M. esquirolii, S. G. Wu 368 (Xinping, Yunnan, China, lectotype of M. subspicata var. intermedia = M. esquirolii).

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Figure II-13. The hairs on the adaxial leaf surface of sampled species, scale bar = 100 μm: A. M. delavayi, Y. B. Zhang 817 (Xundian, Yunnan, China, holotype of M. affinis = M. delavayi); B. M. delavayi, Q. Wang H&W08008 (Tengchong, Yunnan, China); C. M. insuavis, Q. Wang H&W09001 (Gengma, Yunnan, China); D. M. mollis, Q. Wang H&W08004 (Mile, Yunnan, China); E. M. mollis, Q. Wang H&W09003 (Mengla, Yunnan, China); F. M. patchoulii, Q. Wang H&W09010 (Jingdong, Yunnan, China); G. M. patchoulii, R. L. Xiong & S. K. Wen 580981 (Tengchong, Yunnan, China, holotype of M. pauciflora = M. patchoulii); H. M. esquirolii, Q. Wang H&W08006 (Longlin, Guangxi, China); I. M. esquirolii, S. G. Wu 368 (Xinping, Yunnan, China, lectotype of M. subspicata var. intermedia = M. esquirolii).

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Figure II-14. The hairs on the abaxial leaf surface of sampled species, scale bar = 100 μm: A. M. delavayi, Y. B. Zhang 817 (Xundian, Yunnan, China, holotype of M. affinis = M. delavayi); B. M. delavayi, Q. Wang H&W08008 (Tengchong, Yunnan, China); C. M. insuavis, Q. Wang H&W09001 (Gengma, Yunnan, China); D. M. mollis, Q. Wang H&W08004 (Mile, Yunnan, China); E. M. mollis, Q. Wang H&W09003 (Mengla, Yunnan, China); F. M. patchoulii, Q. Wang H&W09010 (Jingdong, Yunnan, China); G. M. patchoulii, R. L. Xiong & S. K. Wen 580981 (Tengchong, Yunnan, China, holotype of M. pauciflora = M. patchoulii); H. M. esquirolii, Q. Wang H&W08006 (Longlin, Guangxi, China); I. M. esquirolii, S. G. Wu 368 (Xinping, Yunnan, China, lectotype of M. subspicata var. intermedia = M. esquirolii).

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Figure II-15. The hair ornamentation on the adaxial leaf surface of sampled species, scale bar = 10 μm: A. M. delavayi, Y. B. Zhang 817 (Xundian, Yunnan, China, holotype of M. affinis = M. delavayi); B. M. delavayi, Q. Wang H&W08008 (Tengchong, Yunnan, China); C. M. insuavis, Q. Wang H&W09001 (Gengma, Yunnan, China); D. M. mollis, Q. Wang H&W08004 (Mile, Yunnan, China); E. M. mollis, Q. Wang H&W09003 (Mengla, Yunnan, China); F. M. patchoulii, Q. Wang H&W09010 (Jingdong, Yunnan, China); G. M. patchoulii, R. L. Xiong & S. K. Wen 580981 (Tengchong, Yunnan, China, holotype of M. pauciflora = M. patchoulii); H. M. esquirolii, Q. Wang H&W08006 (Longlin, Guangxi, China); I. M. esquirolii, S. G. Wu 368 (Xinping, Yunnan, China, lectotype of M. subspicata var. intermedia = M. esquirolii).

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23

Figure II-16. The hair ornamentation on the abaxial leaf surface of sampled species, scale bar = 10 μm: A. M. delavayi, Y. B. Zhang 817 (Xundian, Yunnan, China, holotype of M. affinis = M. delavayi); B. M. delavayi, Q. Wang H&W08008 (Tengchong, Yunnan, China); C. M. insuavis, Q. Wang H&W09001 (Gengma, Yunnan, China); D. M. mollis, Q. Wang H&W08004 (Mile, Yunnan, China); E. M. mollis, Q. Wang H&W09003 (Mengla, Yunnan, China); F. M. patchoulii, Q. Wang H&W09010 (Jingdong, Yunnan, China); G. M. patchoulii, R. L. Xiong & S. K. Wen 580981 (Tengchong, Yunnan, China, holotype of M. pauciflora = M. patchoulii); H. M. esquirolii, Q. Wang H&W08006 (Longlin, Guangxi, China); I. M. esquirolii, S. G. Wu 368 (Xinping, Yunnan, China, lectotype of M. subspicata var. intermedia = M. esquirolii).

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A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

Figure II-17. The stomata density on the abaxial leaf surface of sampled species, scale bar = 50 μm: A. M. delavayi, Y. B. Zhang 817 (Xundian, Yunnan, China, holotype of M. affinis = M. delavayi); B. M. delavayi, Q. Wang H&W08008 (Tengchong, Yunnan, China); C. M. insuavis, Q. Wang H&W09001 (Gengma, Yunnan, China); D. M. mollis, Q. Wang H&W08004 (Mile, Yunnan, China); E. M. mollis, Q. Wang H&W09003 (Mengla, Yunnan, China); F. M. patchoulii, Q. Wang H&W09010 (Jingdong, Yunnan, China); G. M. patchoulii, R. L. Xiong & S. K. Wen 580981 (Tengchong, Yunnan, China, holotype of M. pauciflora = M. patchoulii); H. M. esquirolii, Q. Wang H&W08006 (Longlin, Guangxi, China); I. M. esquirolii, S. G. Wu 368 (Xinping, Yunnan, China, lectotype of M. subspicata var. intermedia = M. esquirolii).

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25

Figure II-18. The stomata morphology of sampled species, scale bar = 5 μm: A. M. delavayi, Y. B. Zhang 817 (Xundian, Yunnan, China, holotype of M. affinis = M. delavayi); B. M. delavayi, Q. Wang H&W08008 (Tengchong, Yunnan, China); C. M. insuavis, Q. Wang H&W09001 (Gengma, Yunnan, China); D. M. mollis, Q. Wang H&W08004 (Mile, Yunnan, China); E. M. mollis, Q. Wang H&W09003 (Mengla, Yunnan, China); F. M. patchoulii, Q. Wang H&W09010 (Jingdong, Yunnan, China); G. M. patchoulii, R. L. Xiong & S. K. Wen 580981 (Tengchong, Yunnan, China, holotype of M. pauciflora = M. patchoulii); H. M. esquirolii, Q. Wang H&W08006 (Longlin, Guangxi, China); I. M. esquirolii, S. G. Wu 368 (Xinping, Yunnan, China, lectotype of M. subspicata var. intermedia = M. esquirolii).

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A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

INFLORESCENCE Cymes are common in Microtoena, which are usually dichotomous, many flowered, axillary or in terminal panicles. The flower number per cyme was emphasized as diagnostic for M. affinis: 1-flowered, and M. pauciflora: 2–3-flowered (Hsuan, 1965). However, dissection of the inflorescence of the holotype of M. affinis reveals it actually has a cyme with five flowers (Wang & Hong, 2012). The statistical analysis (Wang & Hong, 2011) of populations of M. pauciflora also negates the former judgement (Hsuan, 1965) on its flower number. For the arrangement of inflorescence of Microtoena, two types can be easily recognized. The first type is lax with more or less 1-side branching (Figure II-19A), which is found in most species, such as M. patchoulii, M. insuavis, M. delavayi, and M. muliensis. The second one is compact, spikelike, or shortly ovoid (Figure II-19B). The spikelike panicles can be observed in M. esquirolii and M. stenocalyx, and the shortly ovoid panicles are represented by M. miyiensis and M. prainiana. The species M. moupinensis is a special case for inflorescence type. The inflorescence is highly variable from lax, more or less compact, to spikelike panicles, even within the same individual.

Figure II-19. The inflorescences of Microtoena, scale bar =1cm: A. the lax cymes of M. mollis, Q. Wang H&W08004 (Mile, Yunnan, China); B. the compact cymes of M. esquirolii, Q. Wang H&W08006 (Longlin, Guangxi, China).

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BRACTS The bracts of Microtoena are usually linear, lanceolate or ovate, minute, sometimes early deciduous. The linear bract is common in Microtoena and found in most species. The lanceolate and ovate ones are only found in some species, such as M. esquirolii, M. stenocalyx, M. miyiensis and M. prainiana. To reveal the variation range of bracts in population and species level, the bract length and shape are statistically analyzed as follows:

BRACT LENGTH The analysis at population level (Figure II-20) shows that the variation range of bract length is continuous within species. The shortest bract is observed in the two populations (H&W09001, H&W09002) of M. insuavis, which is distinct from all the other populations sampled. The result shows that the bract length of M. moupinensis is

Figure II-20. The variation of bract length in sampled populations of Microtoena: 1. M. esquirolii, D.Y. Hong & Q. Wang, H&W08006 (Longlin, Guangxi, China); 2. M. esquirolii, D.Y. Hong & Q. Wang, H&W08007 (Longlin, Guangxi, China); 3. M. insuavis, D. Y. Hong & Q. Wang, H&W09001 (Gengma, Yunnan, China); 4. M. insuavis, D. Y. Hong & Q. Wang, H&W09002 (Gengma, Yunnan, China); 5. M. mollis, D. Y. Hong & Q. Wang, H&W08004 (Mile, Yunnan, China); 6. M. mollis, D. Y. Hong & Q. Wang, H&W08005 (Mile, Yunnan, China); 7. M. mollis, D. Y. Hong & Q. Wang, H&W09003 (Mengla, Yunnan, China); 8. M. mollis, D. Y. Hong & Q. Wang, H&W09004 (Mengla, Yunnan, China); 9. M. mollis, D. Y. Hong & Q. Wang, H&W09006 (Mile, Yunnan, China); 10. M. moupinensis, D. Y. Hong & Q. Wang, H&W09007 (Baoxing, Sichuan, China); 11. M. moupinensis, D. Y. Hong & Q. Wang, H&W09008 (Baoxing, Sichuan, China); 12. M. patchoulii, D. Y. Hong & Q. Wang, H&W08009 (Tengchong, Yunnan, China); 13. M. patchoulii, D. Y. Hong & Q. Wang, H&W09010 (Jingdong, Yunnan, China); 14. M. patchoulii, D. Y. Hong & Q. Wang, H&W09011 (Jingdong, Yunnan, China).

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highly variable with a wide variation range (0.37–0.73 cm for H&W09007, 0.38–0.58 cm for H&W09008) and high standard deviation (0.095 cm for H&W09007, 0.56 cm for H&W09008). The five populations of M. mollis overlap each other in variation range, but the variation range of populations (H&W08004, H&W08005, H&W09006) from Mile County is wider than that of populations (H&W09003, H&W09004) from Mengla County. The analysis in species level (Figure II-21) reveals that Microtoena is diverse in bract length. Seven species, including M. insuavis, M. patchoulii, M. urticifolia, M. muliensis, M. omeiensis, M. nepalensis and M. robusta, have short bracts (the average < 0.4 cm), which have already been described as ‘tiny’ or ‘minute’ (Stearn, 1983; Li & Hedge, 1994). The shortest bract is represented by M. nepalensis, with the variation range of 0.16–0.21 cm, the average of 0.183 cm, and the standard deviation of 0.017 cm. Based on the analysis, the bract length of M. insuavis (0.21–0.51 cm), M. mollis (0.33–0.64 cm), M. moupinensis (0.32–0.73 cm) is highly variable, and the high standard deviation (0.051 cm for M. insuavis; 0.082 cm for M. mollis; 0.071 cm for M. moupinensis) of these species also show its instability of bract length. The averages from the holotypes are listed here for M. albescens (0.53 cm) and M. miyiensis (0.42 cm), which are typical for the genus Microtoena. Unfortunately, since no other collections are available, the variation range and stability for these two species are still unclear.

Figure II-21. The variation of bract length of Microtoena in species level: 1. M. esquirolii; 2. M. insuavis; 3. M. patchoulii; 4. M. mollis; 5. M. delavayi; 6. M. wardii; 7. M. urticifolia; 8. M. albescens; 9. M. stenocalyx; 10. M. miyiensis; 11. M. megacalyx; 12. M. moupinensis; 13. M. prainiana; 14. M. muliensis; 15. M. omeiensis; 16. M. brevipedunculata; 17. M. nepalensis; 18. M. robusta; 19. M. vanchingshanensis.

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BRACT SHAPE In the present study, the index BR (bract ratio) is used to evaluate this character statistically. The formula to calculate the bract ratio is as follows: BR (bract ratio) = BW (bract width) / BL (bract length). The statistical analysis of bract shape in the sampled populations shows that the variation range of bract shape is continuous within species (Figure II-22). Two groups can be well recognized in figure II-22. One with higher bract ratio (around 0.29) is represented by the two populations (H&W08006, H&W08007) of M. esquirolii. The other twelve populations of four species constitute the other group with a lower bract ratio (around 0.10). The statistical analysis reveals the distinct difference between the two groups in bract shape.

Figure II-22. The variation of bract shape in sampled populations of Microtoena: 1. M. esquirolii, D.Y. Hong & Q. Wang, H&W08006 (Longlin, Guangxi, China); 2. M. esquirolii, D.Y. Hong & Q. Wang, H&W08007 (Longlin, Guangxi, China); 3. M. insuavis, D. Y. Hong & Q. Wang, H&W09001 (Gengma, Yunnan, China); 4. M. insuavis, D. Y. Hong & Q. Wang, H&W09002 (Gengma, Yunnan, China); 5. M. mollis, D. Y. Hong & Q. Wang, H&W08004 (Mile, Yunnan, China); 6. M. mollis, D. Y. Hong & Q. Wang, H&W08005 (Mile, Yunnan, China); 7. M. mollis, D. Y. Hong & Q. Wang, H&W09003 (Mengla, Yunnan, China); 8. M. mollis, D. Y. Hong & Q. Wang, H&W09004 (Mengla, Yunnan, China); 9. M. mollis, D. Y. Hong & Q. Wang, H&W09006 (Mile, Yunnan, China); 10. M. moupinensis, D. Y. Hong & Q. Wang, H&W09007 (Baoxing, Sichuan, China); 11. M. moupinensis, D. Y. Hong & Q. Wang, H&W09008 (Baoxing, Sichuan, China); 12. M. patchoulii, D. Y. Hong & Q. Wang, H&W08009 (Tengchong, Yunnan, China); 13. M. patchoulii, D. Y. Hong & Q. Wang, H&W09010 (Jingdong, Yunnan, China); 14. M. patchoulii, D. Y. Hong & Q. Wang, H&W09011 (Jingdong, Yunnan, China).

The statistical analysis at species level (Figure II-23) is consistent with that of population level. The lower bract ratio (around 0.10) corresponds to the common

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A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

‘linear’ bract, which is represented by most species of Microtoena. The wider bract, which is usually described as ‘lanceolate’, ‘ovate’, or ‘leaflike’, can be observed in six species, including M. esquirolii (average of BR = 0.323), M. stenocalyx (average of BR = 0.348), M. miyiensis (BR of the holotype = 0.26), M. megacalyx (average of BR = 0.256), M. prainiana (average of BR = 0.447), and M. vanchingshanensis (average of BR = 0.448).

Figure II-23. The variation of bract shape of Microtoena in species level: 1. M. esquirolii; 2. M. insuavis; 3. M. patchoulii; 4. M. mollis; 5. M. delavayi; 6. M. wardii; 7. M. urticifolia; 8. M. albescens; 9. M. stenocalyx; 10. M. miyiensis; 11. M. megacalyx; 12. M. moupinensis; 13. M. prainiana; 14. M. muliensis; 15. M. omeiensis; 16. M. brevipedunculata; 17. M. nepalensis; 18. M. robusta; 19. M. vanchingshanensis.

CALYX The size of the calyx and calyx tooth has been used to subdivide Microtoena (Hsuan, 1965; Wu & Li, 1977). As mentioned in the Chapter I, the calyx and calyx teeth of Microtoena undergo a period of continuing enlargement after anthesis. This makes it very risky to use simple calyx and calyx tooth size in classification. To evaluate these unstable characters reasonably, three indexes are used in the present statistical analysis: CLA (calyx length in anthesis); SR (split ratio) = OTL (average of ordinary tooth length of calyx) / CL (calyx length), and TR (tooth ratio) = LLT (length of the longest tooth) / OTL (average of ordinary tooth length of calyx) (Figure II-24).

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Figure II-24. Illustrations of calyx parameters: A. a calyx of M. megacalyx, the ‘OTL’ is calculated based on the five ordinary teeth numbered from 1 to 5; B. a calyx of M. miyiensis, the ‘LLT’ is represented by the extremely elongated tooth numbered as 3, and the ‘OTL’ is calculated based on the four ordinary teeth numbered as 1, 2, 4 and 5.

CALYX LENGTH The analysis in population level (Figure II-25) shows that the variation range of calyx

Figure II-25. The variation of calyx length in sampled populations of Microtoena: 1. M. esquirolii, D.Y. Hong & Q. Wang, H&W08006 (Longlin, Guangxi, China); 2. M. esquirolii, D.Y. Hong & Q. Wang, H&W08007 (Longlin, Guangxi, China); 3. M. insuavis, D. Y. Hong & Q. Wang, H&W09001 (Gengma, Yunnan, China); 4. M. insuavis, D. Y. Hong & Q. Wang, H&W09002 (Gengma, Yunnan, China); 5. M. mollis, D. Y. Hong & Q. Wang, H&W08004 (Mile, Yunnan, China); 6. M. mollis, D. Y. Hong & Q. Wang, H&W08005 (Mile, Yunnan, China); 7. M. mollis, D. Y. Hong & Q. Wang, H&W09003 (Mengla, Yunnan, China); 8. M. mollis, D. Y. Hong & Q. Wang, H&W09004 (Mengla, Yunnan, China); 9. M. mollis, D. Y. Hong & Q. Wang, H&W09006 (Mile, Yunnan, China); 10. M. moupinensis, D. Y. Hong & Q. Wang, H&W09007 (Baoxing, Sichuan, China); 11. M. moupinensis, D. Y. Hong & Q. Wang, H&W09008 (Baoxing, Sichuan, China); 12. M. patchoulii, D. Y. Hong & Q. Wang, H&W08009 (Tengchong, Yunnan, China); 13. M. patchoulii, D. Y. Hong & Q. Wang, H&W09010 (Jingdong, Yunnan, China); 14. M. patchoulii, D. Y. Hong & Q. Wang, H&W09011 (Jingdong, Yunnan, China).

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A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

length is continuous within species. The results show that M. insuavis has the shortest calyx, the average calyx length is 0.290 cm (H&W09001) and 0.296 cm (H&W09002) respectively. The calyx length of M. esquirolii is similar to that of M. insuavis, with average of 0.306 cm for both of the two populations (H&W08006, H&W08007). The result shows that the calyx length of M. moupinensis is highly variable with wide variation range (0.34–0.69 cm for H&W09007, 0.39–0.56 cm for H&W09008) and high standard deviation (0.087 cm for H&W09007, 0.45 cm for H&W09008). The five populations of M. mollis still overlap each other in variation range. The analysis at species level (Figure II-26) reveals that most species of Microtoena are similar to each other in calyx length. The common length is around 0.4 cm. The shortest calyx is still represented by M. insuavis, which is 0.247 cm in average. The results also reveal the instability of calyx length in three species, including M. delavayi (variation range 0.38–0.61 cm; standard deviation 0.058 cm), M. moupinensis (variation range 0.32–0.69 cm; standard deviation 0.070 cm), and M. muliensis (variation range 0.30–0.50 cm; standard deviation 0.053 cm). For M. albescens and M. miyiensis without other collections available, the calyx length of the holotype is 0.48 cm and 0.35 cm respectively, which are typical sizes in Microtoena.

Figure II-26. The variation of calyx length of Microtoena in species level: 1. M. esquirolii; 2. M. insuavis; 3. M. patchoulii; 4. M. mollis; 5. M. delavayi; 6. M. wardii; 7. M. urticifolia; 8. M. albescens; 9. M. stenocalyx; 10. M. miyiensis; 11. M. megacalyx; 12. M. moupinensis; 13. M. prainiana; 14. M. muliensis; 15. M. omeiensis; 16. M. brevipedunculata; 17. M. nepalensis; 18. M. robusta; 19. M. vanchingshanensis.

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CALYX SPLIT RATIO The analysis in population level (Figure II-27) shows that the calyx split ratio of most populations is around 0.40. The calyx of the two populations (H&W08006, H&W08007) of M. esquirolii is more shallowly divided with a ratio of about 0.29, which is distinctly different from that of the other populations. The variation range of split ratio is continuous within species. In addition, for populations of some species such as M. moupinensis, the variation range of the split ratio is much wider than that of other species.

Figure II-27. The variation of calyx split in sampled populations of Microtoena: 1. M. esquirolii, D.Y. Hong & Q. Wang, H&W08006 (Longlin, Guangxi, China); 2. M. esquirolii, D.Y. Hong & Q. Wang, H&W08007 (Longlin, Guangxi, China); 3. M. insuavis, D. Y. Hong & Q. Wang, H&W09001 (Gengma, Yunnan, China); 4. M. insuavis, D. Y. Hong & Q. Wang, H&W09002 (Gengma, Yunnan, China); 5. M. mollis, D. Y. Hong & Q. Wang, H&W08004 (Mile, Yunnan, China); 6. M. mollis, D. Y. Hong & Q. Wang, H&W08005 (Mile, Yunnan, China); 7. M. mollis, D. Y. Hong & Q. Wang, H&W09003 (Mengla, Yunnan, China); 8. M. mollis, D. Y. Hong & Q. Wang, H&W09004 (Mengla, Yunnan, China); 9. M. mollis, D. Y. Hong & Q. Wang, H&W09006 (Mile, Yunnan, China); 10. M. moupinensis, D. Y. Hong & Q. Wang, H&W09007 (Baoxing, Sichuan, China); 11. M. moupinensis, D. Y. Hong & Q. Wang, H&W09008 (Baoxing, Sichuan, China); 12. M. patchoulii, D. Y. Hong & Q. Wang, H&W08009 (Tengchong, Yunnan, China); 13. M. patchoulii, D. Y. Hong & Q. Wang, H&W09010 (Jingdong, Yunnan, China); 14. M. patchoulii, D. Y. Hong & Q. Wang, H&W09011 (Jingdong, Yunnan, China).

The analysis at species level (Figure II-28) shows that the calyx split ratio of most species of Microtoena is around 0.40. A lower split ratio (average < 0.30) is represented by four species, including M. esquirolii, M. urticifolia, M. robusta, and M. vanchingshanensis. The split ratio of four species (including M. insuavis, M. delavayi, M. moupinensis, and M. muliensis) is highly variable with the variation range much

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A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

wider than that of the other species. In addition, M. albescens (SR = 0.44) and M. miyiensis (SR = 0.37) still have the ‘common’ calyx split of Microtoena, based only on the holotypes, the only material available.

Figure II-28. The variation of calyx split of Microtoena in species level: 1. M. esquirolii; 2. M. insuavis; 3. M. patchoulii; 4. M. mollis; 5. M. delavayi; 6. M. wardii; 7. M. urticifolia; 8. M. albescens; 9. M. stenocalyx; 10. M. miyiensis; 11. M. megacalyx; 12. M. moupinensis; 13. M. prainiana; 14. M. muliensis; 15. M. omeiensis; 16. M. brevipedunculata; 17. M. nepalensis; 18. M. robusta; 19. M. vanchingshanensis.

CALYX TOOTH RATIO The calyx of Microtoena has five calyx teeth. For most species, the five calyx teeth are subequal, such as in M. esquirolii, M. insuavis, and M. patchoulii. In some species such as M. moupinensis, M. omeiensis, and M. brevipedunculata, the five calyx teeth are unequal with one tooth slightly longer than the other four ordinary teeth. In a few species such as M. miyiensis and M. delavayi, the elongated tooth is extremely conspicuous. The series Miyienses of Microtoena is even mainly based on such an extremely elongated tooth (Wu & Li, 1977). In the present study, the index TR will reasonably reveal the relationship between of the elongated tooth and the ordinary teeth, and show the variation range of this character in all the species of Microtoena. The analysis at population level (Figure II-29) shows the average of tooth ratio in all the populations sampled is slightly above 1.0. The two populations of M. moupinensis have relatively higher tooth ratio (1.264 for H&W09007, 1.237 for H&W09008), which reveals that the five calyx teeth are unequal with one tooth slightly extended. In addition, the variation range of the tooth ratio is continuous within species. The tooth

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35

ratios of the five populations of M. mollis are similar to each other, and is stable with low standard deviations.

Figure II-29. The variation of tooth ratio in sampled populations of Microtoena: 1. M. esquirolii, D.Y. Hong & Q. Wang, H&W08006 (Longlin, Guangxi, China); 2. M. esquirolii, D.Y. Hong & Q. Wang, H&W08007 (Longlin, Guangxi, China); 3. M. insuavis, D. Y. Hong & Q. Wang, H&W09001 (Gengma, Yunnan, China); 4. M. insuavis, D. Y. Hong & Q. Wang, H&W09002 (Gengma, Yunnan, China); 5. M. mollis, D. Y. Hong & Q. Wang, H&W08004 (Mile, Yunnan, China); 6. M. mollis, D. Y. Hong & Q. Wang, H&W08005 (Mile, Yunnan, China); 7. M. mollis, D. Y. Hong & Q. Wang, H&W09003 (Mengla, Yunnan, China); 8. M. mollis, D. Y. Hong & Q. Wang, H&W09004 (Mengla, Yunnan, China); 9. M. mollis, D. Y. Hong & Q. Wang, H&W09006 (Mile, Yunnan, China); 10. M. moupinensis, D. Y. Hong & Q. Wang, H&W09007 (Baoxing, Sichuan, China); 11. M. moupinensis, D. Y. Hong & Q. Wang, H&W09008 (Baoxing, Sichuan, China); 12. M. patchoulii, D. Y. Hong & Q. Wang, H&W08009 (Tengchong, Yunnan, China); 13. M. patchoulii, D. Y. Hong & Q. Wang, H&W09010 (Jingdong, Yunnan, China); 14. M. patchoulii, D. Y. Hong & Q. Wang, H&W09011 (Jingdong, Yunnan, China).

The analysis at species level (Figure II-30) shows that the tooth ratio is usually around 1.1 in most species of Microtoena, such as M. esquirolii, M. megacalyx, and M. robusta, which reveals that most species of Microtoena have five subequal calyx teeth. Calyces with one slightly elongated tooth (average tooth ratio from 1.2 to 1.5) are seen in M. moupinensis, M. omeiensis, M. brevipedunculata, and M. nepalensis. Calyces with a conspicuously elongated tooth (TR > 1.5) are seen in M. albescens, M. delavayi, M. miyiensis, M. stenocalyx, M. urticifolia, and M. wardii, of which M. miyiensis has the highest tooth ratio (TR = 2.87). In addition, the tooth ratio is highly variable and unstable in five species, including M. delavayi (variation range of 1.68–2.31; standard deviation of 0.160), M. wardii (variation range of 1.47–2.02; standard deviation of 0.183), M. urticifolia (variation range of 1.83–2.24; standard deviation of 0.152), M. moupinensis (variation range of 1.03–1.49; standard deviation of 0.092), M. nepalensis (variation range of 1.38–1.67; standard deviation of 0.106).

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A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

Figure II-30. The variation of tooth ratio of Microtoena in species level: 1. M. esquirolii; 2. M. insuavis; 3. M. patchoulii; 4. M. mollis; 5. M. delavayi; 6. M. wardii; 7. M. urticifolia; 8. M. albescens; 9. M. stenocalyx; 10. M. miyiensis; 11. M. megacalyx; 12. M. moupinensis; 13. M. prainiana; 14. M. muliensis; 15. M. omeiensis; 16. M. brevipedunculata; 17. M. nepalensis; 18. M. robusta; 19. M. vanchingshanensis.

COROLLA The corolla of Microtoena is highly diverse in the colour, corolla length, corolla tube ratio (= corolla tube length / corolla length), shape of middle lobe of lower corolla lip, and apex of upper corolla lip. The taxonomical significance of these characters is evaluated as follows:

COROLLA COLOUR The corolla of Microtoena is mostly yellow, light yellow or white (Figure II-31). Three types of the upper lip colour can be easily recognized (Figure II-32). For the first type, the color is uniform over the whole corolla, including the upper lip (Figure II-32A), which is seen in most species, such as M. esquirolii, M. moupinensis, and M. prainiana. The second type is only observed in M. mollis, where the upper lip color is unstable (Figure II-32B). Here the upper lip can be purely yellow as in the rest of the corolla, with varying amounts of red spots, or purely red, even within the same individual. The third type is the purely red or dark red upper lip (Figure II-32C), which is represented by species such as M. delavayi, M. patchoulii, and M. robusta.

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Figure II-31. The corolla color of Microtoena: A. M. esquirolii, light yellow corolla (photographed by Q. Wang in Guangxi, China); B. M. patchoulii, yellow corolla with the red upper lip (photographed by Q. Wang in Yunnan, China); C. M. mollis, yellow corolla with more or less red spots on the upper lip (photographed by Q. Wang in Yunnan, China); D. M. delavayi, light yellow corolla with the dark red upper lip (photographed by Dr. Y.S. Chen in Yunnan, China); E. M. wardii, light yellow corolla (photographed by Dr. B. Liu in Tibet, China); F. M. stenocalyx, light yellow corolla (photographed by Q. Wang in Yunnan, China); G. M. moupinensis, yellow corolla (photographed by Q. Wang in Sichuan, China); H. M. robusta, white corolla with the dark red upper lip (photographed by Dr. C.L. Xiang in Hubei, China).

Figure II-32. The color of upper corolla lip of Microtoena in population level, scale bar = 1 cm: A. M. esquirolii, purely light-yellow corolla without any red spot on the upper lip (photographed by Q. Wang in Guangxi, China); B. M. mollis, yellow corolla with more or less red spots on the upper lip (photographed by Q. Wang in Yunnan, China); C. M. patchoulii, yellow corolla with the purely red upper lip (photographed by Q. Wang in Yunnan, China).

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A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

COROLLA LENGTH The corolla length was emphasized as diagnostic for some taxa of Microtoena, such as M. delavayi var. grandiflora (Hsuan, 1965) and M. siamica (Stearn, 1983). The present study evaluates this character statistically for all the species recognized in Microtoena, and the result of standard deviation analysis reveals notable differentiation in the corolla length of Microtoena. The analysis at population level (Figure II-33) shows that the variation range of calyx length is continuous within species, and two groups of corolla length can be well recognized. The first group with the longer corolla (corolla length > 2.0 cm) is represented by two populations of M. moupinensis (average of corolla length = 3.016 cm for H&W09007, and 3.004 cm for H&W09008). The other populations form the second group with a corolla length of about 1.5 cm. The results also show that the five populations of M. mollis still overlap each other in variation range, but one population

Figure II-33. The variation of corolla length in sampled populations of Microtoena: 1. M. esquirolii, D.Y. Hong & Q. Wang, H&W08006 (Longlin, Guangxi, China); 2. M. esquirolii, D.Y. Hong & Q. Wang, H&W08007 (Longlin, Guangxi, China); 3. M. insuavis, D. Y. Hong & Q. Wang, H&W09001 (Gengma, Yunnan, China); 4. M. insuavis, D. Y. Hong & Q. Wang, H&W09002 (Gengma, Yunnan, China); 5. M. mollis, D. Y. Hong & Q. Wang, H&W08004 (Mile, Yunnan, China); 6. M. mollis, D. Y. Hong & Q. Wang, H&W08005 (Mile, Yunnan, China); 7. M. mollis, D. Y. Hong & Q. Wang, H&W09003 (Mengla, Yunnan, China); 8. M. mollis, D. Y. Hong & Q. Wang, H&W09004 (Mengla, Yunnan, China); 9. M. mollis, D. Y. Hong & Q. Wang, H&W09006 (Mile, Yunnan, China); 10. M. moupinensis, D. Y. Hong & Q. Wang, H&W09007 (Baoxing, Sichuan, China); 11. M. moupinensis, D. Y. Hong & Q. Wang, H&W09008 (Baoxing, Sichuan, China); 12. M. patchoulii, D. Y. Hong & Q. Wang, H&W08009 (Tengchong, Yunnan, China); 13. M. patchoulii, D. Y. Hong & Q. Wang, H&W09010 (Jingdong, Yunnan, China); 14. M. patchoulii, D. Y. Hong & Q. Wang, H&W09011 (Jingdong, Yunnan, China).

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(H&W09004) is more unstable with the standard deviation of 0.119 cm. In addition, the corolla length is highly variable and unstable in the two populations of M. moupinensis (variation range of 2.75–3.67 cm with standard deviation of 0.280 in H&W09007; variation range of 2.70–3.44 cm with standard deviation of 0.253 in H&W09008). The analysis at species level (Figure II-34) supports the differentiation in corolla length of Microtoena. The longer corolla is seen in most species, such as M. delavayi, M. stenocalyx, and M. moupinensis. The typical corolla length is around 2.0 cm, and the longest corolla was observed in M. moupinensis (average of corolla length = 2.829 cm). The shorter corolla (about 1.5 cm) is represented by four species, including M. esquirolii, M. insuavis, M. patchoulii, and M. mollis, with the corolla of M. insuavis the shortest (average of corolla length = 1.191 cm; variation range of 0.78–1.40 cm). In addition, the results reveal that the corolla length is highly variable and unstable in four species: M. urticifolia (variation range of 1.62–2.64 cm; standard deviation of 0.422 cm), M. megacalyx (variation range of 1.73–2.70; standard deviation of 0.370 cm), M. moupinensis (variation range of 1.64–3.67; standard deviation of 0.420 cm), and M. omeiensis (variation range of 1.67–2.75; standard deviation of 0.356 cm).

Figure II-34. The variation of corolla length of Microtoena in species level: 1. M. esquirolii; 2. M. insuavis; 3. M. patchoulii; 4. M. mollis; 5. M. delavayi; 6. M. wardii; 7. M. urticifolia; 8. M. albescens; 9. M. stenocalyx; 10. M. miyiensis; 11. M. megacalyx; 12. M. moupinensis; 13. M. prainiana; 14. M. muliensis; 15. M. omeiensis; 16. M. brevipedunculata; 17. M. nepalensis; 18. M. robusta; 19. M. vanchingshanensis.

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COROLLA TUBE RATIO The corolla tube of Microtoena can be generally divided into ‘short tube’ and ‘long tube’ (Figure II-35). In the present study, a new index ‘tube ratio’ is introduced to show the relationship between the lengths of the tube and the overall corolla. The corolla tube ratio (CTR) = the corolla tube length (CTL) / the corolla length (CL). The corolla tube is statistically evaluated for all the species of Microtoena, based on the standard deviation analysis of the CTR, both in population and species level.

Figure II-35. The differentiation in the corolla tube of Microtoena, scale bar = 1 cm: A. M. patchoulii, a flower with the ‘short’ corolla tube; B. M. moupinensis, a flower with the ‘long’ corolla tube.

The analysis at population level (Figure II-36) shows that the variation range of the tube ratio is continuous within species, and two groups of corolla length can be clearly recognized. The first group with the higher tube ratio (CTR > 0.6) is represented by the two populations of M. moupinensis (average tube ratio = 0.683 for H&W09007, and 0.674 for H&W09008). The other populations form the second group with a tube ratio of around 0.2. The results also show that the variation ranges of the five populations of M. mollis overlap each other. In addition, all the populations sampled are similar to each other in the stability of tube ratio, with the standard deviation around 0.1.

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Figure II-36. The variation of the corolla tube ratio in sampled populations of Microtoena: 1. M. esquirolii, D. Y. Hong & Q. Wang, H&W08006 (Longlin, Guangxi, China); 2. M. esquirolii, D. Y. Hong & Q. Wang, H&W08007 (Longlin, Guangxi, China); 3. M. insuavis, D. Y. Hong & Q. Wang, H&W09001 (Gengma, Yunnan, China); 4. M. insuavis, D. Y. Hong & Q. Wang, H&W09002 (Gengma, Yunnan, China); 5. M. mollis, D. Y. Hong & Q. Wang, H&W08004 (Mile, Yunnan, China); 6. M. mollis, D. Y. Hong & Q. Wang, H&W08005 (Mile, Yunnan, China); 7. M. mollis, D. Y. Hong & Q. Wang, H&W09003 (Mengla, Yunnan, China); 8. M. mollis, D. Y. Hong & Q. Wang, H&W09004 (Mengla, Yunnan, China); 9. M. mollis, D. Y. Hong & Q. Wang, H&W09006 (Mile, Yunnan, China); 10. M. moupinensis, D. Y. Hong & Q. Wang, H&W09007 (Baoxing, Sichuan, China); 11. M. moupinensis, D. Y. Hong & Q. Wang, H&W09008 (Baoxing, Sichuan, China); 12. M. patchoulii, D. Y. Hong & Q. Wang, H&W08009 (Tengchong, Yunnan, China); 13. M. patchoulii, D. Y. Hong & Q. Wang, H&W09010 (Jingdong, Yunnan, China); 14. M. patchoulii, D. Y. Hong & Q. Wang, H&W09011 (Jingdong, Yunnan, China).

The analysis at species level (Figure II-37) supports differentiation by tube ratio in Microtoena. The lower tube ratio is represented by four species: M. esquirolii (CTR = 0.183), M. insuavis (CTR = 0.128), M. patchoulii (CTR = 0.220) and M. mollis (CTR = 0.128). The other fifteen species all have higher tube ratios above 0.5, of which the highest tube ratio is observed in M. moupinensis (CTR = 0.677). In addition, the result shows that the tube ratio of M. delavayi is relatively variable and unstable, with a variation range of 0.51–0.64 and a standard deviation of 0.031.

APEX OF THE UPPER LIP OF THE COROLLA The upper lip of the corolla of Microtoena is usually straight, galeate, and laterally compressed. Two types (Figure II-38) can be easily recognized for the apex of the

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Figure II-37. The variation of the corolla tube ratio of Microtoena in species level: 1. M. esquirolii; 2. M. insuavis; 3. M. patchoulii; 4. M. mollis; 5. M. delavayi; 6. M. wardii; 7. M. urticifolia; 8. M. albescens; 9. M. stenocalyx; 10. M. miyiensis; 11. M. megacalyx; 12. M. moupinensis; 13. M. prainiana; 14. M. muliensis; 15. M. omeiensis; 16. M. brevipedunculata; 17. M. nepalensis; 18. M. robusta; 19. M. vanchingshanensis.

Figure II-38. The apex of upper corolla lip of Microtoena, scale bar = 1 mm: A. M. delavayi, the entire apex; B. M. mollis, the emarginate apex.

upper lip. One is entire, which is represented by species such as M. delavayi, M. muliensis, and M. omeiensis. The other is emarginate, which is represented by most species, such as M. esquirolii, M. patchoulii, and M. miyiensis.

HAIRS OF THE UPPER LIP OF THE COROLLA Most species of Microtoena have hairs on the external surface of the upper corolla lip, which are usually stiff hairs with 2–3 nodes under SEM (Figure II-39).

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Figure II-39. The hairs on the external surface of upper corolla lip of Microtoena, scale bar = 50 μm: A. M. insuavis, a 2-nodes stiff hair; B. M. mollis, a 2-nodes stiff hair; C. M. esquirolii, a 3-nodes stiff hair.

MIDDLE LOBE OF THE LOWER LIP OF THE COROLLA The lower lip of the corolla of Microtoena is usually spreading, 3-lobed, and relatively longer than the upper one. The middle lobe of the lower corolla lip is diverse in Microtoena, and three types can be clearly recognized (Figure II-40). In the first type, the middle lobe is sub-circular and the largest of the three lower lobes (Figure II-40A). This is seen in species such as M. delavayi, M. wardii, and M. moupinensis. In the other two types, the middle lobe is smaller than the two lateral lobes. In the second type, the middle lobe is oblong-ovate (Figure II-40B), which is seen in species such as M. esquirolii, M. mollis, and M. vanchingshanensis. In the third type, it is ligulate (Figure II-40C), which is seen in two species, M. insuavis and M. patchoulii.

Figure II-40. The middle lobe of lower corolla lip of Microtoena, scale bar = 1 mm: A. M. delavayi, the subcircular middle lobe; B. M. esquirolii, the oblong-ovate middle lobe; C. M. insuavis, the ligulate middle lobe.

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STAMENS Species of Microtoena have four stamens, which are subequal and included in the upper corolla lip or slightly exserted.

ANTHERS Each stamen of Microtoena has two anther cells, which are apically divaricate, and becoming confluent before pollination (Figure II-41A). The anther cells dehisce along the long axis (Figure II-41B).

Figure II-41. The anther cells of Microtoena patchoulii (D.Y. Hong & Q. Wang, H&W08009, Tengchong, Yunnan, China), scale bar = 0.5 mm: A. the confluent two anther cells; B. the longitudinal dehiscing anther cells with pollen grains.

HAIRS ON THE FILAMENTS The filament of Microtoena is usually barbate at the base (Figure II-42A). The hairs are tiny (Figure II-42B) and with a smooth surface (Figure II-42C).

Figure II-42. The hairs on filaments of Microtoena patchoulii (D.Y. Hong & Q. Wang, H&W08009, Tengchong, Yunnan, China): A. hairs on the basal part of a filament, scale bar = 50μm; B. the dense hairs on the filament, scale bar = 50μm; C. the smooth surface of a hair, scale bar = 10μm.

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STYLE The style of Microtoena is almost as long as the stamens and included in the upper corolla lip or slightly exserted. The apex of style splits into two extremely unequal lobes (Figure II-43). The anterior lobe is much longer and subulate. The posterior lobe is extremely minute and even barely visible in some species, such as M. delavayi (Figure II-43A) and M. insuavis (Figure II-43B).

Figure II-43. The style lobes of Microtoena, scale bar = 500 μm: A. M. delavayi, Y. B. Zhang 817 (Xundian, Yunnan, China, holotype of M. affinis = M. delavayi): B. M. insuavis, Q. Wang H&W09001 (Gengma, Yunnan, China); C. M. mollis, Q. Wang H&W09003 (Mengla, Yunnan, China); D. M. patchoulii, Q. Wang H&W09010 (Jingdong, Yunnan, China); E. M. esquirolii, Q. Wang H&W08006 (Longlin, Guangxi, China); F. M. esquirolii, S. G. Wu 368 (Xinping, Yunnan, China, lectotype of M. subspicata var. intermedia = M. esquirolii).

POLLEN The pollen grains of Microtoena are tricolpate, ellipsoid in equatorial view (Figure II-44A), and sub-spheroidal in polar view (Figure II-44B). The extine usually has multi-reticulate sculpturing (Figure II-45).

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Figure II-44. The pollen grains of Microtoena, scale bar = 10 μm: A. an ellipsoid pollen grain with elongated apertures in equatorial view; B. a sub-spheroidal pollen grain in polar view.

Figure II-45. The ornamentation of pollen extine of Microtoena, scale bar = 1 μm: A. M. delavayi, Y. B. Zhang 817 (Xundian, Yunnan, China, the holotype of M. affinis = M. delavayi); B. M. insuavis, Q. Wang H&W09001 (Gengma, Yunnan, China); C. M. mollis, Q. Wang H&W09003 (Mengla, Yunnan, China); D. M. patchoulii, Q. Wang H&W09010 (Jingdong, Yunnan, China); E. M. patchoulii, R. L. Hsiung & S. K. Wen 580981 (Tengchong, Yunnan, China, the holotype of M. pauciflora = M. patchoulii); F. M. esquirolii, Q. Wang H&W08006 (Longlin, Guangxi, China).

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FRUITS The nutlets of Microtoena are usually ovoid, subglobose, or triquetrous, tiny, about 2 mm in diameter. The colour of the mature nutlet is usually from brown to black. The surface of the nutlet is usually smooth for most species, but slightly corrugate in M. insuavis.

CHAPTER III. MOLECULAR PHYLOGENY, BIOGEOGRAPHY AND CHARACTER EVOLUTION PHYLOGENETICAL POSITION AND SUBDIVISION Before the molecular phylogenetical studies on Microtoena and allied genera (Scheen et al., 2010; Bendiksby et al., 2011), the phylogenetical position of this genus had been estimated by many taxonomists. D. Prain (1889), who founded this genus, placed his new genus in the tribe Stachydeae, closely related to the genus Craniotome. J.I. Briquet (1897) placed both Microtoena and Craniotome in the subtribe Lamiinae of tribe Stachydeae of the subfamily Stachyoideae. J.W. Kadereit (2004) placed it in the subfamily Lamioideae. Recently molecular phylogenetical analysis sheds new light on the position of this genus. Scheen et al. (2010) and Bendiksby et al. (2011) reveal that Microtoena belongs to the tribe Pogostemoneae in the subfamily Lamioideae, and is sister to the clade consisting of Anisomeles and Pogostemon, with Craniotome situated at the base. However, both these studies are at a large scale (covering the entire subfamily Lamioideae), and only a few species of Microtoena are sampled (only one species in Scheen’s study; two species in Bendiksby study). In Bendiksby’s study, Microtoena with two species sampled (M. delavayi and M. patchoulii) is even non-monophyletic with week support (JK < 50). Therefore, further study on circumscription and intra-generic structure of Microtoena was badly needed. In the present phylogenetic analysis, three chloroplast DNA markers (matK, trnL intron and trnL–F spacer) used by Scheen et al. (2010) and Bendiksby et al. (2011) are selected. To investigate the intra-generic structure, eleven species of Microtoena are sampled for molecular phylogenetic analysis. Allied genera of Microtoena were also sampled based on the existing phylogenetical studies (Scheen et al., 2010; Bendiksby et al., 2011). The result (Figure III-1) shows that all the sampled species of Microtoena form a monophyletic clade, which is sister to the clade consisting of Pogostemon and Anisomeles. Within Microtoena, two sub-clades are well supported and strongly correlated to the morphology. One clade (MP / BI = 100 / 1) consists of the four species with short corolla tubes: M. esquirolii, M. patchoulii, M. insuavis and M. mollis. The

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other seven species sampled form the other clade (MP / BI = 71 / 1), in which all the species have a conspicuously elongated corolla tube.

Figure III-1. The molecular phylogenetic tree derived from the bayesian analysis (the parsimony jackknife support and bayesian posterior probability value are indicated above branches (MP / BI)) shows that the genus Microtoena splits into two groups with different corolla tube lengths.

In its taxonomic history, twenty-nine specific names have been published for Microtoena, and the accepted number of species is around twenty. Unfortunately, the intra-generic structure is still unclear for this genus. Although S.J. Hsuan (1965) proposed a five-series system for Chinese Microtoena, which was expanded to six series by C.Y. Wu & H.W. Li (1977), this system is mainly based on unstable quantitative characters, such as calyx size and calyx tooth length, which are unreliable, and might not reflect the true phylogenetic relationship. The present molecular phylogenetical analysis does not support the six-series system of S.J. Hsuan (1965) and C.Y. Wu & H.W. Li (1977). In practice, the molecular tree supports two well defined groups within Microtoena which might correspond to two sections. In addition, some taxonomic treatments based on character analysis (Wang & Hong, 2011) are supported by the present phylogenetic analysis. For instance, M. esquirolii is reduced to a synonym of M. insuavis by S.T. Dunn (1913), which is followed by C.Y. Wu (1959), S.J. Hsuan (1965), C.Y. Wu & H.W. Li (1977), and H.W. Li & I.C. Hedge (1994). In 2011, based on comprehensive analysis of characters, we made a revision for the M. insuavis complex, and separated M. esquirolii from the M. insuavis complex as an independent species. The present study reveals that M. insuavis is sister to M. patchoulii, and M. esquirolii is situated at the base of the ‘M. insuavis’ clade, which confirms our former treatment.

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ORIGIN AND DIVERSIFICATION Most species of Microtoena are endemic to China, and are mainly distributed in Yunnan, Sichuan and Guizhou provinces of SW China. Some species, such as M. patchoulii and M. insuavis, are widely distributed in SE Asia. A.A.H. Léveillé (1911) did reported a species M. coreana from Soul City of South Korea, which is far from the mainly distribution area of Microtoena. Based on an extensive character analysis, I have excluded this species from Microtoena (Wang, 2015), and transferred it to the genus Tripora as a synonym of T. divaricata. In the present study, to reveal the origin and subsequent diversification of Microtoena, RASP (Reconstruct Ancestral State in Phylogenies) version 2.1 beta is used. The ancestral state in distribution and biogeographic events of Microtoena and allied genera are analyzed based on the Bayesian Binary method. The result shows that the ancestor of Microtoena (node 7) is distributed in SW China with a probability of 91.16% (Figure III-2), which reveals that SW China might be the centre of origin. In addition, Microtoena is highly diversified in this area, with some species such as M. patchoulii, M. insuavis and M. robusta spreading to adjacent areas, including central China, SE Asia, Himalaya region and S Asia, which matches the dispersal events of the node 1 and node 4 (Figure III-2). For the allied genera of Microtoena, vicariance and dispersal events (node 11) are detected

Figure III-2. Reconstruction of ancestral state in phylogenies (the bayesian posterior probability value are indicated above branches, and the biogeographic events are marked above the nodes): A. Central China; B. SW China; C. SE Asia; D. Himalaya region; E. S Asia.

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in the ancestor of Pogostemon and Anisomeles. Dispersal events (nodes 8, 9) are also found within the genus Pogostemon.

CHARACTER EVOLUTION Many characters of Microtoena have been used in its species delimitation and subdivision (Hsuan, 1965; Wu & Li, 1977) within the genus, such as the inflorescence, bract, calyx, calyx tooth and corolla. In chapter II, the variation range and stability of these characters have been evaluated both at population and species levels. However, the evolutionary patterns of these character are still unclear. In the present study, RASP is used to re-establish the ancestral state of eight characters with taxonomic significance. For the bract shape, the ‘linear’ type is common in Craniotome, Anisomeles, and most species of Microtoena. The ‘lanceolate’ or ‘ovate’ bract can be found in Pogostemon and some species of Microtoena. The result (Figure III-3A) shows that the linear bract

Figure III-3. Reconstruction of ancestral state in phylogenies of diagnostic characters of Microtoena and allied genera (the bayesian posterior probability value are indicated above branches, and the pie charts represent the relative probability of ancestral state): A. the evolutionary pattern of bract shape; B. the evolutionary pattern of calyx split; C. the evolutionary pattern of calyx teeth; D. the evolutionary pattern of inflorescence.

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should be primary with high probability (95.99%) for Microtoena and allied genera, while the ovate bract is derived independently in Microtoena and Pogostemon. In addition, the ovate bract is derived twice in the genus Microtoena, for the two sub-clades respectively. The analysis of the depth of the calyx split (Figure III-3B) shows that shallow calyx splitting (less than one third) is primary with the probability of 52.67% for Microtoena and allied genera. The derived deep calyx splitting (up to half) is common in Microtoena, Pogostemon and Anisomeles, but it is reversed to the primary state in a few species of Microtoena, such as in M. esquirolii and M. robusta. The result shows that the equal or subequal calyx teeth are primary for Microtoena and allied genera, with the probability of 99.85% (Figure III-3C). The unequal types with one elongated tooth (1.2 < TR < 1.5) or extremely elongated tooth (TR > 1.5) are just derived independently in certain species of Microtoena. Although the cymes of most species of Microtoena are lax, the result shows the compact inflorescence is primary for Microtoena and allied genera, with the probability of 99.72% (Figure III-3D). The ancestral state of Microtoena is also compact with the probability of 90.04%. The lax cymes are derived independently within the two sub-clades of Microtoena. The result shows that the ‘small’ flower is primary for Microtoena and allied genera, with the probability of 99.76% (Figure III-4A). The ‘large’ flower is derived within one sub-clade, which includes most species of Microtoena. Species of the other sub-clade of Microtoena still retain the primary state in flower size. None of the existing taxonomic studies (Wu, 1959; Hsuan, 1965; Wu & Li, 1977, Stearn, 1983) has paid attention to the taxonomic significance of corolla tube elongation in Microtoena. Based on the character analysis in chapter II, Microtoena can be subdivided into two groups with lower corolla tube ratio (ca. 0.2) and higher corolla tube ratio (CTR > 0.5) respectively. However, the ‘elongated corolla tube’ in Microtoena is not a derived character. In fact, the ancestral state reconstruction shows that the elongated corolla tube is common and primary in Microtoena and allied genera (Figure III-4B). Therefore, the ‘reduced’ corolla tube should be used here for the derived state in one sub-clade of Microtoena. The result shows that the ‘wider’ middle lobe of lower corolla lip is primary in Microtoena and allied genera, with the probability of 99.8%(Figure III-4C). The ‘oblong-ovate’ and ‘ligulate’ middle lobes which are narrower than the lateral lobes are both derived within one sub-clade of Microtoena. Finally, the result shows that the ‘entire’ apex of the upper corolla lip is primary in the genera analyzed, with the probability of 99.79% (Figure III-4C). The ‘emarginate’ apex of the upper corolla lip is derived within one sub-clade of Microtoena.

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Figure III-4. Reconstruction of ancestral state in phylogenies of diagnostic characters of Microtoena and allied genera (the bayesian posterior probability value are indicated above branches, and the pie charts represent the relative probability of ancestral state): A. the evolutionary pattern of flower length; B. the evolutionary pattern of corolla tube; C. the evolutionary pattern of middle lobe shape of lower corolla lip; D. the evolutionary pattern of apex of upper corolla lip.

CHAPTER IV. TAXONOMIC REVISION OF

MICROTOENA MICROTOENA PRAIN Prain, Hook. Icon. Pl. 19: t. 1872. 1889; Briquet in Engl. u. Prantl, Nat. Pflanzenfam. 4, 3a: 268. 1897; Wu, Acta Phytotax. Sin. 8(1): 44–49. 1959; Hsuan, Acta Phytotax. Sin. 10(1): 41–56. 1965; Hsuan in Wu & Li, Fl. Reip. Pop. Sin. 66: 46. 1977; Stearn, Journ. Jap. Bot. 58(1): 1–13. 1983; Li & Hedge in Wu, Raven & Hong, Fl. China. 17: 50. 1994. Type: Microtoena patchoulii (C.B. Clarke ex Hook. f.) C.Y. Wu & S.J. Hsuan (= Plectranthus patchouli C.B. Clarke ex Hook. f. = Microtoena cymosa Prain nom. illeg.) Herbs perennial. Roots straight, simple. Stems erect, puberulent, hirsute, or hispid, base sometimes woody. Leaf blade papery, oblong-ovate, triangular-ovate, ovate to broadly ovate, adaxially puberulent or hirsutulous, abaxially hispid, villous and minutely glandular; base cuneate, truncate, subcordate to cordate; margin serrate or crenate, sometimes mucronate; apex acute, acuminate, or caudate-acuminate. Cymes axillary or terminal, dichotomous, sometimes with 1-sided branches, or in compact as spikelike or ovoid panicles. Bracts linear or lanceolate, sometimes caducous. Calyx 5-toothed, dilated after anthesis; teeth triangular, lanceolate to subulate, subequal or one tooth conspicuously elongated. Corolla white to yellow, sometimes marked with red or dark red on upper lip, sparsely hirsutulous outside; corolla tube conspicuous or reduced and inconspicuous; upper lip laterally compressed, apex entire or emarginate; lower lips 3-lobed with middle lobe ligulate, oblong-ovate, or subcircular, lateral lobes ovate. Stamens 4, subequal, included in upper lip or slightly exserted; filaments complanate, base barbate; anthers 2-cells, apically divaricate, confluent in anthesis, recondite under the upper lip. Stigma 2-lobed, one lobe extremely short to inconspicuous. Pollen grains tricolpate, ellipsoid; colpi long; extine sculpture multi-reticulate. Nutlets dark brown to black, smooth, sometimes slightly corrugate. Phenology: flowering from July to late March, and fruiting from November to May. Habitats: forests, forest margins, ravines, grassy slopes and thickets; from 100 m in De Lang-Son, Vietnam (M. insuavis) to 3650 m on Yulong Mountain, Lijiang, Yunnan Province, China (M. delavayi). Distribution: C, S and SW China, Bhutan, Indonesia, Laos, Myanmar, Nepal, North India, Thailand, and Vietnam (Map IV-1).

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Map IV-1. The distribution pattern of the genus Microtoena Prain.

KEY TO SECTIONS AND SPECIES OF MICROTOENA 1a. Corolla tube reduced and inconspicuous; middle lobe of lower corolla lip oblong- ovate or ligulate, narrower than lateral lobes (sect. I. Microtoena: spp. 1–4). 2a. Cymes compact, in spikelike panicles; bracts lanceolate. .................................1. M. esquirolii 2b. Cymes lax, branches sometimes 1-sided; bracts linear. 3a. Leaf base cuneate or truncate; leaf margin serrate; upper corolla lip uniformly red or dark red. 4a. Stems hirsute; leaf base cuneate; abaxial leaf surface densely villous ........................... ................................................................................................................. 2. M. insuavis

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4b. Stems hispid; leaf base truncate; abaxial leaf surface densely hispid.... 3. M. patchoulii 3b. Leaf base cordate; leaf margin mucronate crenate; upper corolla lip with scattered red spots................................................................................................................. 4. M. mollis 1b. Corolla tube conspicuous; middle lobe of lower corolla lip subcircular, ovate or obovate, wider than lateral lobes (sect. II. Delavayana: spp. 5–19). 5a. Stems hispid to densely hispid, hirsute; calyx teeth unequal, one tooth conspicuously elongated. 6a. Cymes lax, dichotomous, sometimes with 1-sided branches; bracts linear. 7a. Cymes dichotomous; corolla pale yellow, marked with dark red on upper lip............... ................................................................................................................ 5. M. delavayi 7b. Cymes with 1-sided branches; corolla uniformly white or pale yellow. 8a. Leaves large, average leaf length more than 9.5 cm; corolla tube base slightly dilated .................................................................................................. 6. M. wardii 8b. Leaves small, average leaf length around 6.5 cm; corolla tube base not dilated. 9a. Leaf margin coarsely serrate, apex long-acuminate; corolla pale yellow.............. .................................................................................................. 7. M. urticifolia 9ab. Leaf margin mucronate crenate, apex shortly acuminate; corolla white ................ ................................................................................................ 8. M. albescens 6b. Cymes compact in spikelike or ovoid panicles; bracts lanceolate. 10a. Leaf base truncate to subcordate; cymes axillary and terminal, in spikelike panicles.. ........................................................................................................... 9. M. stenocalyx 10b. Leaf base cuneate; cymes in terminal ovoid panicles. ....................... 10. M. miyiensis 5b. Stems sparsely puberulent to nearly glabrous; calyx teeth subequal. 11a. Calyx broadly campanulate; calyx teeth triangular-subulate, apex slightly curved or hooked. 12a. Cymes dichotomous, lax or compact in spikelike panicle; bract lanceolate to linear. 13a. Cymes lax; bracts lanceolate. .............................................. 11. M. megacalyx 13b. Cymes with 1-sided branches, sometimes compact in spikelike panicles; bract linear ......................................................................... 12. M. moupinensis 12b. Cymes compact in ovoid panicles; bracts ovate. ........................... 13. M. prainiana 11b. Calyx campanulate to tubular; calyx teeth triangular to lanceolate, apex straight. 14a. Leaf blade oblong-ovate; leaf base cuneate; cymes with 1-sided branches .............. ....................................................................................................... 14. M. muliensis 14b. Leaf blade ovate to triangular-ovate; leaf base truncate to subcordate; cymes not 1-sided. 15a. Leaf base truncate; calyx deeply split (up to halfway). 16a. Leaf blade triangular-ovate; bracts conspicuous. 17a. Inflorescence long pedunculate; calyx dusty glandular, teeth triangular ............................................................................... 15. M. omeiensis 17b. Inflorescence shortly pedunculate; calyx minutely puberulent, teeth lanceolate ..................................................... 16. M. brevipedunculata 16b. Leaf blade ovate; bracts tiny, inconspicuous. ................ 17. M. nepalensis 15b. Leaf base subcordate; calyx shallowly split (less than 1/3). 18a. Stems robust and erect; corolla white, marked with dark red on upper lip .......................................................................................... 18. M. robusta 18b. Stems base prostrate; corolla uniformly white.......................................... ........................................................................ 19. M. vanchingshanensis

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SECTION 1. Microtoena Herbs perennial. Cymes lax, sometimes with 1-sided branches, or compact in spikelike panicles. Bracts linear or oblong-ovate. Calyx 5-toothed, calyx teeth subequal. Corolla yellow or pale yellow, sometimes marked with red, dark red or scattered red spots on upper lip, corolla tube reduced to inconspicuous, apex of upper corolla lip emarginate, middle lobe of lower corolla lip oblong-ovate or ligulate, narrower than lateral lobes. Widely distributed in S and SW China, Laos, N India, Indonesia, Myanmar, Thailand, and Vietnam. Species 1–4. 1. Microtoena esquirolii H. Lév., Repert. Spec. Nov. Regni Veg. 9: 222. 1911. Type: China, Guizhou, Zhenfeng [Tchenfong], Oct. 1905, J. Esquirol 672 [holotype: E! (E00284286)]. Microtoena subspicata C.Y. Wu ex S.J. Hsuan, Acta Phytotax. Sin. 10: 45. 1965. Type: China, Guangxi, Longlin, Yancha, in valley, 900 m, 02 Nov 1957, C.C. Chang 10679 [holotype: IBSC! (IBSC 0005116)]. Microtoena subspicata var. intermedia C.Y. Wu ex S.J. Hsuan, Acta Phytotax. Sin. 10: 46. 1965. Type: China, Yunnan, Xinping, Mosha, Daxing, in valley, 700 m, 16 Oct. 1958, S.K. Wu 368 [holotype: KUN! (KUN 0484545); isotype: KUN! (KUN 0484546)]. Herbs perennial. Stems erect, base sometimes woody, 0.4–1.3m tall, sparsely puberulent. Leaf petiole 2–7 cm; leaf blade papery, adaxially sparsely white hirsutulous to subglabrous, abaxially sparsely hispid and minutely glandular, ovate, 3–9 cm long, 2–6 cm broad; base cuneate; margin serrate, mucrones absent; apex abruptly long- acuminate. Cymes dichotomous, glandular hairy, axillary and terminal, 3–10 flowered, compact in spikelike panicles; peduncle inconspicuous. Bracts oblong-ovate, 5–6 mm long, 1.5–2 mm broad. Calyx 2.8–3.3 mm long at anthesis, sparsely pubescent, dilated after anthesis, 5-toothed; calyx teeth subequal, 0.8–1 mm long, triangular-ovate, apex acute. Corolla 1–1.6 cm long, uniformly yellow to pale yellow, sparsely hirsutulous outside; upper lip laterally compressed, apex emarginate; lower lip 3-lobed, middle lobe oblong-ovate, narrower than lateral lobes. Nutlets black, glabrous, 1.2–1.5 mm long, 1.0–1.2 mm broad, triquetrous, subglobose. Figures. IV-1 & 2. Phenology: Flowering from September to November and fruiting from November to December. Distribution: This species is distributed in Guangxi, Guizhou and Yunnan of China (Map IV-2).

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Figure IV-1. Microtoena esquirolii H. Lév.: a. flowering stem; b. flower and dissected corolla; c. bract; d. calyx with dissection.

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Figure IV-2. Microtoena esquirolii H. Lév.: A. field individuals photographed by the author in Longlin County of Guangxi Province (China); B. cymes in spikelike panicle; C. sparsely puberulent stem; D. dissected corolla with calyx and bract; E. compact cyme with 10 flowers. Scale bars = 5 mm.

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Map IV-2. Microtoena esquirolii H. Lév.

Habitats: It grows in valleys, along stream sides, and on stony slopes, and hill sides at altitudes from 300 m in Daxin to 1200 m in Tianlin. Note: M. esquirolii was reduced to a synonym of M. insuavis by S.T. Dunn (1913) soon after being described by A.A.H. Léveillé (1911). This treatment is widely followed by later studies (Wu, 1959; Hsuan, 1965; Wu & Li, 1977; Li & Hedge, 1994). In fact, there are several distinct differences between M. esquirolii and M. insuavis: (1) stems sparsely puberulent for M. esquirolii but densely hirsute for M. insuavis (Figure IV-3A & B); (2) leaf surface sparsely hispid for M. esquirolii but densely villous and hispid for M. insuavis (Figure IV-3C & D); (3) hair surface granular for M. esquirolii but smooth for M. insuavis (Figure IV-3E & F); (4) cymes compact in spikelike panicles for M. esquirolii but lax and with 1-sided branches for M. insuavis (Figure IV-3G & H); (5) corolla uniformly yellow or pale yellow for M. esquirolii but marked with red on upper lip for M. insuavis (Figure IV-3I & J), and (6) middle lobe of lower corolla lip oblong-ovate for M. esquirolii but ligulate for M. insuavis (Figure IV-3K & L). The result of principal coordinate analysis (MVSP-Version 3.13b is used for the principal coordinate analysis of the present study) of multiple characters (Figure IV-4) also supports the separation of M. esquirolii and M. insuavis. In addition, the principal coordinate analysis reveals that M. esquirolii, M. subspicata and M. subspicata var. intermedia are conspecific to each other and circumscribed in the same entity (Figure IV-4). I have examined all the type specimens involved, and no distinct difference is observed for the the three taxa. S.J. Hsuan (1965) was apparently aware of the differences between M. esquirolii and M. insuavis, but failed to examine the type specimen of M. esquirolii. Consequently, she seems to just follow the treatment of S.T. Dunn (1913) but

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Figure IV-3. The comparison of M. esquirolii and M. insuavis: A. stem sparsely puberulent (M. esquirolii); B. stem densely hirsute (M. insuavis); C. leaf surface sparsely hispid (M. esquirolii); D. leaf surface densely villous and hispid (M. insuavis); E. hair surface granular (M. esquirolii); F. hair surface smooth (M. insuavis); G. cymes compact(M. esquirolii); H. cymes lax (M. insuavis); I. corolla uniformly yellow (M. esquirolii); J. upper corolla lip red (M. insuavis); K. middle lobe of lower corolla lip oblong-ovate (M. esquirolii); L. middle lobe of lower corolla lip ligulate (M. insuavis).

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Figure IV-4. Principal coordinate analysis shows the separation of M. esquirolii and M. insuavis: a. types of M. subspicata; b. types of M. subspicata var. intermedia; c. the holotype of M. esquirolii; d. the holotype with randomly sampled specimens of M. insuavis; k. population samples of M. esquirolii, D.Y. Hong & Q. Wang, H&W08006 (Longlin, Guangxi, China); l. population samples of M. esquirolii, D.Y. Hong & Q. Wang, H&W08007 (Longlin, Guangxi, China); n. population samples of M. insuavis, D. Y. Hong & Q. Wang, H&W09001 (Gengma, Yunnan, China); o. population samples of M. insuavis, D. Y. Hong & Q. Wang, H&W09002 (Gengma, Yunnan, China).

describe a new species M. subspicata, which is actually M. esquirolii. In addition, S.J. Hsuan (1965) described a new variety, M. subspicata var. intermedia, which is mainly based on its shorter spikelike panicles. However, based on my field observation, the length of spikelike panicle is highly variable and unstable. The shorter inflorescence of M. subspicata var. intermedia and the longer inflorescence of M. subspicata are absolutely in the same continuous variation range. Thus, I reduce M. subspicata var. intermedia as well as M. subspicata to synonyms of M. esquirolii. Additional specimens examined: CHINA, s. loc., s. d., s. coll. 396 (IBSC). GUANGXI, Daxin: Xialei, Xinfeng, stone slopes, 300-650 m, 16 Oct 2008, Chinese Botanic Union-Guangxi Exped. 4041 (IBK). Jingxi: Tongde, Guoli, in valley, 586-628 m, 24 Oct 2008, Chinese Botanic Union-Guangxi Exped. 4556 (IBK). Leye: Bailang station, road side, 360 m, 04 Dec 1986, Longtan Exped. 01859 (IBK). Longlin, Yancha, in thickets, 1250 m, 30 Nov 2008, Q. Wang HW08006 (K, PE, SZ); Yancha, Nongjin, River side, 910 m, 01 Dec 2008, Q. Wang HW08007 (K, PE, SZ). Longzhou: Banbitun, Mt. Paotai, in valley, 09 Dec 1957, P. X. Tan 57324 (IBSC). Napo: Delong, Pingda, stream side, 840 m, 26 Oct 1998, Z. Y. Li et al. 1569 (PE). Tianlin: Mt. Cengwanglao, Langping, in valley, 1200 m, 13 Oct 1989, South China sub-Exped. 2494 (IBSC); loc. eodem, in valley, 1200 m, 13 Oct 1989, South China Exped. 2530 (IBSC). GUIZHOU, Ceheng: Huarong, Balai, road side, 1000 m, 27 Oct. 1958, T. Y. Tsao 1147 (PE). Luodian: Yanglixiayi, road side, 410 m, 01 Nov 1986, Longtan Exped. 00590 (IBK).

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Panxian: Yangchang, Gesuo, ravine side, 600 m, 26 Sep 1988, C. Z. Dang 927 (HGAS). Zhenfeng: hill side, 16 Oct 1930, Y. Tsiang 7334 (IBSC, NAS). 2. Microtoena insuavis (Hance) Prain ex Briq., Nat. Pflanzenfam., div. 4, 3a: 269. 1895. = Gomphostemma insuave Hance, J. Bot. 22: 231. 1884. = Microtoena insuavis (Hance) Prain ex Dunn nom. illeg., Notes Roy. Bot. Gard. Edinburgh. 8(37): 169. 1913. Type: China, Guangdong, Shaoguan, N. River, 200 m, Nov 1883, B.C. Henry 22237 [holotype: BM! (BM 000950459)]. Microtoena siamica Stearn, J. Jap. Bot. 58(1): 11.1983. Type: Thailand, Payap, Doi Chiengdao, Doi Luang Chiang Dao, in evergreen forest, 1500 m, 03 Nov 1922, A.F.G. Kerr 6538 [holotype: BM! (BM 000950463); isotype: BM! (BM 000950464)]. Herbs perennial. Stems 0.6–1.5 m tall, base sometimes woody, densely hirsute. Leaf petiole 2-10 cm; leaf blade papery, adaxially white hirsutulous, abaxially densely villous, hispid and minutely glandular, ovate, 2.8–10.6 cm long, 1.6–7.8 cm broad; base cuneate; margin serrate, mucro absent; apex acute. Cymes axillary and terminal, dichotomous with 1-sided branches; peduncle 2–5 cm long. Bracts linear, 0.2–0.5 cm long, less than 0.5 mm broad. Calyxes 1.5–3 mm long at anthesis, puberulent, dilated after anthesis, 5-toothed; calyx teeth subequal, 0.7–1.5 mm long, triangular-lanceolate, apex acute. Corolla 0.7–1.4 cm long, yellow, marked with dark red on upper lip, sparsely hirsutulous outside; apex of upper lip emarginate; lower lip 3-lobed, middle lobe ligulate, lateral lobes triangular. Nutlets 0.8–1.3 mm long, 0.7–1.1 mm broad, smooth, sometimes slightly corrugate. Figures. IV-5 & 6. Phenology: Flowering from October to January and fruiting from December to February. Distribution: This species is widely distributed in South China, Indonesia, Thailand, and Vietnam (Map IV-3). Habitat: It grows at forest and thicket margins, and on grassy slopes, at altitudes from 90 m on Dinghu Mountain, Zhaoqing of Guangdong Province (China) to 1200 m in Huanglian Mountain, Luquan of Yunnan Province (China). Note: W.T. Stearn (1983) described M. siamica from Thailand as new species. This species is extremely similar to M. insuavis, but has shorter and somewhat compact inflorescence and shorter flowers. Based on my field observation, the short flowers of M. siamica are still within the variation range of M. insuavis. I have examined the types of M. siamica, the shorter and slightly compact inflorescence of which is apparently immature. Dissection of the short inflorescence reveals that the cymes are still dichotomous and with 1-sided branches. The result of principal coordinate analysis of multiple characters (Figure IV-7) confirms that M. insuavis and M. siamica are conspecific and circumscribed in the same entity. As a result, M. siamica should be reduced to a synonym of M. insuavis.

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Figure IV-5. Microtoena insuavis (Hance) Prain ex Briq.: a. flowering stem; b. flower marked with dark red on upper lip; c. bract with hairs; d. intact calyx; e. dissected calyx with five subequal teeth.

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Figure IV-6. Microtoena insuavis (Hance) Prain ex Briq.: A. field individuals photographed by the author in Gengma County of Yunnan Province (China); B. densely hirsute stem; C. lax dichotomous cymes with 1-sided branches; D. a densely villous leaf; E. population sampled flowers showing the stable dark red upper lip. Scale bars = 1 cm.

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Map IV-3. Microtoena insuavis (Hance) Prain ex Briq.

Additional specimens examined: s. loc., s. d., s. coll., s. n. (P 04333400). CHINA. s. loc., s. d., s. coll. L936 (P). GUANGDONG, s. loc. 1889, G. Forrest s. n. (K). Boluo, Mt. Luofu, Huashoutai, 245 m, 22 Dec 1928, Y. Tsiang 1693 (BM, NAS). Longzhou, Changtang, in forest, 28 Nov 1957, P. X. Tan 57102 (IBSC); Wulian, Banbitun, in valley, 350 m, 04 Oct 1956, Z. J. Li 3259 (IBSC); Wulian, Banbitun to Jiangshan, in thickets, 400 m, 04 Oct 1956, Z. J. Li 3003 (IBSC). Wengyuan, E Mt. Shangmiao, 09 Oct 1935, X. Q. Liu 24836 (IBK, PE, SZ, WUK); Heping, Mt. Bijia, in thickets, 01 Dec

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Figure IV-7. Principal coordinate analysis of M. siamica and M. insuavis: d. the holotype with randomly sampled specimens of M. insuavis; h. the holotype of M. siamica; n. population samples of M. insuavis, D. Y. Hong & Q. Wang, H&W09001 (Gengma, Yunnan, China); o. population samples of M. insuavis, D.Y. Hong & Q. Wang, H&W09002 (Gengma, Yunnan, China).

1935, X. Q. Liu 25348 (IBK, IBSC). HAINAN, Changjiang, in forest, 10 Sep 1933, X. R. Liang 66028 (IBK, IBSC); Mt. Rulong, in forest, 08 Jan 1934, S. K. Lau 3100 (P). Ledong, Mt. Jianfeng, Fangniu slope, in forest, 28 Jan 1935, S. K. Lau 5474 (IBSC). YUNNAN, s. loc., s. d., s. coll. 59-13833 (IBSC). Gengma, Huken, Back Hill, in thickets, 1330 m, 01 Jan 2009, Q. Wang HW09002 (K, PE, SZ); Huken, Mt. Dajian, Dapingzhang, in thickets, 1310 m, 01 Jan 2009, Q. Wang HW09001 (K, PE, SZ); Huken, Mt. Dajian, mountain slopes, 1100 m, 04 Dec 1958, T. P. Zhu 0467 (KUN). Luquan, Mt. Huanglian, Qimaba, in forest, 900-1200 m, 01 Nov 1995, S. K. Wu et al. 1124 (KUN). Mengla, s. loc. woods edge, 900 m, Nov 1936, C. W. Wang 80582 (KUN, PE); Menglun, Mt. Kongming, woods edge, 650 m, Dec 1959, X. Zhou 166 (KUN); Menglun, Tropic Botanic Garden, road side, 540 m, 24 Dec 1961, Y. H. Li 3857 (KUN); Mengyuan, Nanyang, woods edge, 05 Dec 2004, S. S. Zhou 2174 (PE); Rulong, in woods, 1000 m, Oct 1936, C. W. Wang 80153 (KUN, NAS, PE, WUK); Yiwu, Menghan, river side, 4 Nov 1959, S. J. Pei 59-11300 (KUN, WUK). Simao, Dongshan, 1524 m, s. d. A. Henry 12588 (A, K). INDONESIA, JAVA, s. loc. 11 Dec 1902, D. Bois 170 (P). Res Madioeu, Ngebel, 700 m, 25 May 1896, Kaarders 23161b (P). THAILAND, CHIANG MAI, Doi Inthanond, s. loc. in forest, 14 Dec 1964, K. Bunchuai 37088 (P). PHRAE, Mae Kray, s. loc. in forest, 700 m, 09 Jan 1972, C. F. van Beusekom et al. 4760 (P). VIETNAM, DE BAC GIANG, Lang Met, 1925, E. Colani s. n. (P). DE HOABUCH, s. loc. 550 m, 05 Dec 1926, E. Colani 5003 (P); s. loc. Feb 1935, s. coll. 5393 (P). DE LANG-SON, s. loc. 100 m, Nov 1925, E. Colani L.968 (P); s. loc. 19 Dec

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1913, A. Chevalier 29668 (P); Lang-Son, 20 Dec 1913, A. Chevalier 29750 (P). NINH BINH, Cuc Phuong, Cuc Phuong National Park, Biology station, in woods, 360-460 m, 16 Feb 1965, China-Vietnam Exped. 2316 (KUN); Cuc Phuong National Park, CP9: Big Tree Cinnamomam balansae site, 2 Km from Bong, in valley forest, 270 m, 29 Nov 1998, D. D. Soejarto et al. 10427 (P); Cuc Phuong National Park, from Cave of Prehistoric 2Km to the Main Park’s gate, forest edge, 180 m, 03 Jan 2001, P. K. Loc et al. P10531 (P). TONKIN, s. loc. 20 Dec 1887, B. Balansa 3582 (K, P); s. loc. 24 Nov 1889, B. Balansa 3584 (P); s. loc. 800 M, Apr 1914, M. Lemarie s. n. (P); s. loc. Apr 1914, M. Lemarie s. n. (P). Bau Phceug Nay, s. loc. 700 M, Apr 1914, M. Lemarie s. n. (P). Bieu Hieu, s. loc. 28 Jan 1892, s. coll. S034 (P). Cho Bo, 80 Km SW of Hanoi, 1887, B. Balansa 3583 (K, P). 3. Microtoena patchoulii (C.B. Clarke ex Hook. f.) C.Y. Wu & S.J. Hsuan, Acta Phytotax. Sin. 10: 44. 1965. = Plectranthus patchouli C.B. Clarke ex Hook. f., Fl. Brit. India 4: 624. 1885. = Microtoena cymosa Prain nom. illeg., Hooker’s Icon. Pl. 19: pl. 1872. 1889. Type: India, Assam, Khasia Mts, Johra, 1220 m, 09 Oct 1871, C.B. Clarke 15983 [Lectotype designated here: K! (K 000928190)]; India, Assam, s. d., Jenkins s. n. [syntype: K]. Microtoena pauciflora C.Y. Wu ex S.J. Hsuan, Acta Phytotax. Sin. 10: 44. 1965. Type: China, Yunnan, Tengchong, 1st District, in forest, 1600 m, 17 Dec 1958, R.L. Hsiung & S.K. Wen 580981 [holotype: KUN! (KUN 0484547)]. Herbs perennial. Stems 0.5–2 m tall, densely hispid, base sometimes woody. Leaf petiole 1–5 cm long; leaf blade papery, spreading hispid and minutely glandular, 2.8–11.3 cm long, 1.9–8.1 cm broad, triangular-ovate; base truncate to subcordate; margin serrate, mucrones absent; apex acute. Cymes axillary and terminal, dichotomous, lax, sometimes with 1-sided branches; peduncle 1.5–4 cm long. Bracts linear, 2.3–4.6 mm long, less than 0.5 mm broad. Calyx 2.3–3.8 mm long at anthesis, puberulent, dilated after anthesis, 5-toothed; calyx teeth subequal, 1–1.7 mm long, triangular-ovate, apex acute. Corolla 1.1–1.6 cm long, yellow, marked with red on upper lip, sparsely hirsutulous outside, upper lip apex emarginate, lower lip 3-lobed, middle lobe ligulate, narrower than lateral lobes. Nutlets dark brown to black, triquetrous, ovoid, 1.2–1.6 mm long, 1.0–1.3 mm broad, smooth. Figures. IV-8 & 9. Phenology: Flowering from October to December and fruiting in January to March. Distribution: This species is distributed in South China, North India and Myanmar (Map IV-4). Habitat: It grows at forest and thicket margins, and on open grass slope, at altitudes from 670 m in Yiwu of Mengla to 1700 m in Zhefang of Luxi of Yunnan province.

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Figure IV-8. Microtoena patchoulii (C. B. Clarke ex Hook. f.) C. Y. Wu & S. J. Hsuan: a. flowering stem; b. flower; c. linear bract; d. dissected calyx.

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Figure IV-9. Microtoena patchoulii (C. B. Clarke ex Hook. f.) C. Y. Wu & S. J. Hsuan: A. field individuals photographed by the author in Tengchong County of Yunnan Province (China); B. lax dichotomous cymes; C. densely hispid stem; D. population samples of flowers showing the uniformly red upper corolla lip; E. hispid leaf with truncate base. Scale bars = 1 cm.

Note: M. patchoulii was firstly placed in Plectranthus as P. patchouli by C.B. Clarke and J.D. Hooker (1885). In the protologue of P. patchouli, C.B. Clarke and J.D. Hooker cited another specimen (preserved in the herbarium K) collected by Jenkins from Assam (India) together with the specimen (preserved in the herbarium K) collected by C.B. Clarke from Khasia Mountain. Therefore, I make a lectotypification here and selected Clarke’s specimen as lectotype. D. Prain (1889) separated it from Plectranthus, and transferred it to his new genus Microtoena. However, D. Prain rejected the epithet ‘patchouli’ and renamed P. patchouli as M. cymosa, according to the now abandoned ‘Kew Rule’ of priority of epithets being limited to their use only under a given generic name. S.T. Dunn (1913) regarded M. cymosa (= M. patchoulii) as

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conspecific with M. insuavis, and reduced the former to a synonym. S.J. Hsuan (1965) separated M. patchoulii from M. insuavis as independent species, based on their differences in stem indumentum and leaf base, and reduced Prain's illegitimate name M. cymosa to a synonym of M. patchoulii. This treatment is followed by later studies (Wu & Li, 1977; Stearn, 1983; Li & Hedge, 1994) and supported by the principal coordinate analysis of multiple characters (Figure IV-10).

Map IV-4. Microtoena patchoulii (C. B. Clarke ex Hook. f.) C. Y. Wu & S. J. Hsuan.

Figure IV-10. Principal coordinate analysis of M. pauciflora, M. patchoulii and M. insuavis: d. holotype of M. insuavis; f. lectotype plus randomly sampled specimens of M. patchoulii; g. holotype of M. pauciflora; m. population samples of M. patchoulii, D.Y. Hong & Q. Wang, H&W08009 (Tengchong, Yunnan, China); n. population samples of M. insuavis, D. Y. Hong & Q. Wang, H&W09001 (Gengma, Yunnan, China); o. population samples of M. insuavis, D. Y. Hong & Q. Wang, H&W09002 (Gengma, Yunnan, China); s. population samples of M. patchoulii, D. Y. Hong & Q. Wang, H&W09010 (Jingdong, Yunnan, China); t. population samples of M. patchoulii, D. Y. Hong & Q. Wang, H&W09011 (Jingdong, Yunnan, China).

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In addition, S.J. Hsuan (1965) described a new species, M. pauciflora, which is extremely similar to M. patchoulii. This new species is just based on fewer flowers and smaller leaves than that of M. patchoulii. Although H.W. Li & I.C. Hedge (1994) accepted M. pauciflora in the Flora of China, they remarked that ‘This species may only be a variety of Microtoena patchoulii’. Based on my filed observation (Figure IV-11) in the type locality of M. pauciflora (Tengchong County, Yunnan Province, China), the flower number per cyme and the leaf size of M. pauciflora and M. patchoulii are within the same continuous variation range. The result of principal coordinate analysis of multiple characters (Figure IV-10) confirms that M. pauciflora and M. patchoulii are conspecific and circumscribed in the same entity. Therefore, M. pauciflora should be reduced to a synonym of M. patchoulii.

Figure IV-11. Two representative individuals from the same population of M. patchoulii (D.Y. Hong & Q. Wang, H&W08009, Tengchong, Yunnan, China): A. a plant with many flowered cymes and ‘larger’ leaves; B. a plant with 2–3 flowered comes and ‘smaller’ leaves, in accordance with the description of M. pauciflora.

Additional specimens examined: CHINA, s. loc., s. d., G. Forrest 26217 (P); s. loc., s. d., G. Forrest 25192 (P). YUNNAN, s. loc., 2750 m, Nov. 1924, G. Forrest 26017 (K, P); s. loc. 1952, R. C. Ching 50636 (KUN); 1952, R. C. Ching 50702 (KUN). Jingdong, Hsiao-Chang-Chien, in thickets, 1200 m, 07 Dec 1939, M. K. Li 1363 (IBSC, KUN, QTPMB); Huangcaoling, in thickets, 2450 m, 30 Oct 2009, Q. Wang HW09011 (K, PE, SZ); Huangcaoling, Xiaoheping, in woods, 2200 m, 30 Oct 2009, Q. Wang HW09010 (K, PE, SZ); Lansuo, road side, 1250 m, 02 Nov 1993, H. Peng 1904 (KUN). Lianghe,

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Dachangqu, in village, 1400 m, 11 Dec 1958, R. L. Hsiung & S. K. Wen 580890 (KUN). Longling, Bawan, in ravine, G. S. Yang 1426 (KUN). Luxi, Zhefang, Xishan, Nongbing to Yingpan, in village, 1700 m, 25 Nov 1960, W. Q. Yin 60-1521 (KUN); Zhefang, Nongqiu, in thickets, 01 Dec 1955, J. Chen 451 (KUN). Menghai, Mt. Nannuo, slopes, 1500-2000 m, 08 Nov 1933, Y. Tsiang 12597 (IBSC, KUN, NAS, PE). Mengla, Yiwu, in valley, 670 m, 22 Dec 1958, W. T. Wang 10429 (KUN); Hills NW of Yengyueh, 1924, G. Forrest 26082 (BM, IBSC, K, PE). Tengchong, Qushi, Shuanghe, Laojiazhai, in field, 1520 m, 25 Dec 2008, Q. Wang HW08009 (K, PE, SZ). INDIA, ASSAM, Khasia Hills, Feb 1888, S. Mann s. n. (K). LAOS, LOUANG PRABANG, Pres de Ban Sat, 30 Dec 1948, J. E. Vidal 777B (P). MYANMAR, SUMPRABUN, s. loc. open places along the forest edge, 610 m, 15 Dec 1953, F. Kingdon-Ward 21695 (BM, K). 4. Microtoena mollis H. Lév., Repert. Spec. Nov. Regni Veg. 9: 222. 1911. Type: China, Guizhou, Jiaozhuang [Kiaotsong], 13 Dec 1904, J. Esquirol 755 [lectotype designated here: E! (E 00284285)]; China, Guizhou, Kiaotsong [Jiaozhuang], 13 Dec 1904, J. Esquirol 330 [syntype: E! (E 00284283)]; China, Guizhou, Lofou, Mar 1909, J. Cavalerie 3548 [syntypes: E! (E 00284284), K! (K 000928183)]. Herbs perennial. Stems 0.5–2 m tall, densely hispid. Leaf petiole 2–6 cm long; leaf blade papery, adaxially white hirsutulous, abaxially densely hispid and minutely glandular, ovate, 3.3–11.8 cm long, 2.4–8.6 cm broad; base cordate; margin mucronate crenate; apex shortly acuminate. Cymes, axillary and terminal, dichotomous, lax, usually with 1-sided branches; peduncle 1–5 cm long. Bracts linear, 3.2–6.4 mm long, ca. 0.5 mm broad. Calyx 3.1–4.8 mm long at anthesis, puberulent, dilated after anthesis, 5-toothed; calyx teeth subequal, lanceolate, 1.2–2 mm long. Corolla 1.2–1.8 cm long, yellow, marked with red spots on upper lip, sparsely hirsutulous outside; apex of upper lip emarginate, lower lip 3-lobed, middle lobe ligulate, narrower than lateral lobes, reflexed before opening of flower. Nutlets 1.3–1.6 mm long, 1.1–1.3 mm broad, black to brown, smooth. Figures. IV-12 & 13. Phenology: Flowering from November to March and fruiting from April to May. Distribution: This species is distributed in Guizhou, Yunnan and Guangxi province of China (Map V-5). Habitat: It grows at the margins of woods and thickets, at altitudes from 400 m in Tian’e of Guangxi and Luodian of Guizhou to 1200 m in Shizong of Yunnan province. Note: Together with M. cymosa and M. esquirolii, M. mollis was also reduced to a synonym of M. insuavis by S.T. Dunn (1913), soon after being described by A.A.H. Léveillé (1911). Based on its distinct indumentum, leaf base and leaf margin, S.J. Hsuan (1965) reinstated it as independent species. This treatment is followed by later studies (Wu & Li, 1977; Li & Hedge, 1994) and supported by the principal coordinate analysis of multiple characters (Figure IV-14).

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Figure IV-12. Microtoena mollis H. Lév.: a. flowering stem; b. a flower with bract; c. a calyx and a dissected calyx; d. linear bracts.

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Figure IV-13. Microtoena mollis H. Lév.: A. field individuals photographed by the author in Mile County of Yunnan Province (China); B. lax dichotomous cymes with 1-sided branches; C. a densely hispid stem; D. a hispid leaf with cordate base; E. population samples of flowers showing the highly variable and unstable red spots scattered on upper corolla lip. Scale bars = 1 cm.

In the protologue of M. mollis, A.A.H. Léveillé (1911) cited three collections, and recorded one collection ‘J. Esquirol 755’ as ‘J. Esquirol 155’ apparently by mistake. Here the specimen J. Esquirol 755 preserved in the herbarium E (E00284285) is designated as lectotype of M. mollis.

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Map IV-5. Microtoena mollis H. Lév..

Figure IV-14. Principal coordinate analysis of M. mollis and M. insuavis: d. the holotype of M. insuavis; e. the lectotype with randomly sampled specimens of M. mollis; i. population samples of M. mollis, D. Y. Hong & Q. Wang, H&W08004 (Mile, Yunnan, China); j. population samples of M. mollis, D. Y. Hong & Q. Wang, H&W08005 (Mile, Yunnan, China); n. population samples of M. insuavis, D. Y. Hong & Q. Wang, H&W09001 (Gengma, Yunnan, China); o. population samples of M. insuavis, D. Y. Hong & Q. Wang, H&W09002 (Gengma, Yunnan, China); p. population samples of M. mollis, D. Y. Hong & Q. Wang, H&W09003 (Mengla, Yunnan, China); q. population samples of M. mollis, D. Y. Hong & Q. Wang, H&W09004 (Mengla, Yunnan, China); r. population samples of M. mollis, D. Y. Hong & Q. Wang, H&W09006 (Mile, Yunnan, China).

Additional specimens examined: CHINA, GUANGXI, Longlin, Wuchong, in valley, 01 May 1957, Z. F. Liang & D. L. Wu 32050 (IBK, IBSC). Tian’e, Nazhi, Baidongtun, ditch side, 400 m, 05 May 1989, Beijing Exped. 890847 (PE). GUIZHOU, Luodian, Bamao, Xiaotianba, in woods, 400 m, 22 Mar 1960, Guizhou Exped. 198 (PE). YUNNAN, Jinghong, Mengyang, Manzhanghuiha, hill side, 950 m, 27 Mar

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1957, China-Soviet Union Exped. 5710 (IBSC, KUN). Malipo, Ba-Buh, woods edge, 1000 m, 01 Feb 1940, C. W. Wang 86480 (IBSC, KUN, PE). Mengla, Manda, Mt. Jiulong, in woods, s. d., Y. H. Li 02841 (KUN); Menglun, Baqiaxiaozhao, stream side, 620 m, 05 Jan 2009, Q. Wang HW09004 (K, PE, SZ); loc. eodem, stream side, 620 m, 29 Apr 2013, Q. Wang W13003 (PE); Menglun, Manrun, stream side, 680 m, 11 Dec 1961, Y. H. Li 3821 (HITBC, IBK, IBSC, KUN); loc. eodem, stream side, 24 Feb 1964, Y. H. Li 4768 (HITBC, IBSC, KUN); Menglun, Mengkuan, stream side, 24 Dec 1963, Y. H. Li 5084 (HITBC, IBSC, KUN); Menglun, Tropic Botanic Garden, 19 Nov 2010, G. Yao 378 (IBSC); loc. eodem, in woods, 18 Jan 1964, Y. H. Li 5175 (HITBC, IBK, IBSC, KUN); Menglun to Menghai, road side, 19 Nov 2010, G. Yao 377 (IBSC); Menglun, Xiaola Road, road side, 700 m, 05 Jan 2009, Q. Wang HW09003 (K, PE, SZ); Menglun, Xiaola Road 55 Km, road side, 27 Dec 1959, Y. H. Li 1732 (HITBC, KUN); Menglun, Xiaola Road 59 Km, in woods, 560 m, 11 Jan 1959, Y. H. Li 309 (HITBC, IBK, IBSC, KUN). Mile, Dongshan, Luona, in valley, 1180 m, 09 Dec 2005, E. D. Liu 1464 (KUN); loc. eodem, in thickets, 1140 m, 26 Nov 2008, Q. Wang HW08004 (K, PE, SZ); Laozhai, in thickets, 1180 m, 26 Nov 2008, Q. Wang HW08005 (K, PE, SZ); Dayubang, in thickets, 1070 m, 08 Jan 2009, Q. Wang HW09006 (K, PE, SZ). Shizong, dark ravine, 1200 m, s. d., A. Henry 11583 (K) SECTION. 2. Delavayana Q. Wang sect. nov. —Type: Microtoena delavayi Prain. Herbs perennial. Cymes dichotomous, lax, sometimes with 1-sided branches, or compact in spikelike or ovoid panicles. Bracts linear to lanceolate. Calyxes 5-toothed, sometimes one tooth conspicuously elongated. Corolla white, pale yellow, yellow, sometimes marked with dark red on upper lip, corolla tube conspicuous, apex of upper corolla lip entire, middle lobe of lower corolla lip subcircular or ovate, wider than lateral lobes. Distributed in Central and Southwest China (Hunan, Hubei, Guizhou, Sichuan, Yunnan, Tibet), Bhutan, North India (Assam), Nepal. Species 5–19. 5. Microtoena delavayi Prain, Bull. Soc. Bot. France 42: 424. 1895. = Microtoena delavayi var. vera Prain, nom. inval., Bull. Soc. Bot. France 42: 425. 1895. Type: China, Yunnan, Heqing, 19 Oct 1886, M.J. Delavay 2463 [holotype: P! (P00743569); isotypes: P! (P00743570, P00743571, P00743572, P04444595)]. Microtoena affinis C.Y. Wu & S.J. Hsuan, Acta Phytotax. Sin. 10: 48–49. 1965. Type: China, Yunnan, Xundian, Mt. Dabei, in woods, 2800 m, 01 Nov 1940, Y.P. Chang 817 [holotype: KUN! (KUN0484539)]. Microtoena bhutanica Stearn, J. Jap. Bot. 58(1): 11. 1983. syn. nov. Type: Bhutan, Hinglai La, 2438 m [8000 ft], 24 Aug 1949, F. Ludlow et al. 19640 [holotype: BM! (BM000950461)]. Microtoena delavayi var. grandiflora Prain, Bull. Soc. Bot. France 42: 425. 1895. syn. nov. Type: China, Yunnan, Ta-long-tan to Ta-pin-tze, 1800 m, 21 Jul 1886, M.J.

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Delavay 2190 [lectotype designated here: P! (P00743575); isolectotypes: P! (P00743573, P00743574)]; China, Yunnan, Ta-long-tan, 10 Oct 1889, M.J. Delavay 4206 [syntypes: P! (P00743576, P00743577, P00743578)]. Microtoena tenuiflora C.Y. Wu, Acta Phytotax. Sin. 8: 47–48. 1959. Type: China, Yunnan, Weishan, Mt. Weibao, in valley, 2300 m, 08 Sep 1933, Y. Tsiang 11884 [holotype: PE! (PE00031251); isotypes: IBSC! (IBSC0583597, IBSC0584279), KUN! (KUN0217754), NAS! (NAS00072411, NAS00223564, NAS00223566, NAS00223568, NAS00223574), PE! (PE00824243), WUK! (WUK0120785)]. Herbs perennial. Stems erect, 1–2 m tall, densely hispid, base sometimes woody. Leaf petiole 2–16 cm; leaf blade papery, adaxially hirsutulous, abaxially hispid and minutely glandular, ovate, 4.6–17.2 cm long, 3–11.4 cm broad; base truncate to subcordate; margin mucronate crenate; apex shortly caudate-acuminate. Cymes axillary and terminal, dichotomous, lax, branches sometimes 1-sided; peduncle 1–6 cm long. Bracts linear, 4.3–5.6 mm long, ca. 0.5 mm broad. Calyx 3.8–6.1 mm long at anthesis, hispid, dilated after anthesis, 5-toothed; calyx teeth lanceolate, unequal, one calyx tooth conspicuously elongated; tooth apex acuminate. Corolla 1.4–2.6 cm long, yellow to pale yellow, marked with dark red on upper lip, sparsely hirsutulous outside; corolla tube 0.8–1.5 cm long; upper corolla lip laterally compressed, apex entire; lower corolla lip 3-lobed, middle lobe subcircular, wider than lateral lobes. Nutlets 1.5–2.1 mm long, 1.1–1.6 mm broad, dark brown, smooth. Figures. IV-15 & 16. Phenology: Flowering from July to November and fruiting from September to January of the next year. Distribution: This species is widely distributed in Southwest China (Sichuan and Yunnan provinces) and Bhutan (Map IV-6). Habitat: It grows in forests, at forest and thicket margins, on open grass slopes, at altitudes from 1700 m in Yunnan province to 3500 m in Sichuan province. Note: S.J. Hsuan (1965) described M. affinis as new species, based on its 1-flowered axillary cyme, small leaf blade and remotely crenate leaf margin. H.W. Li & I.C. Hedge (1994) accepted M. affinis in the Flora of China, but indicated that ‘this species may be a form of M. delavayi’ I have carefully examined the holotype of M. affinis, and found that its ‘1-flowered axillary cyme’ is actually a cyme with five flowers (Figure IV-17). The holotype of M. affinis is collected from a new shoot. Based on my field observation, it is common in M. delavayi that new shoots of this perennial species generally have smaller leaves and fewer flowers compared with the main stem. In addition, the leaf margins on new shoots are generally more sparsely toothed than those of the main stem. No distinct difference has been observed between the leaves of M. affinis and those of the new shoots of M. delavayi (Figure IV-18). It is interesting that the holotype collector, Y.P. Zhang, collected four specimens (China, Yunnan, Xundian, Mt. Dabei, 01 Nov 1940, Y.P. Zhang 0811, IBSC, KUN, PE), together with the holotype of M. affinis at the same day in the type locality, all of which should be

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Figure IV-15. Microtoena delavayi Prain: a. flowering stem; b. a part of cyme; c. linear bract; d. a dissected calyx with one elongated tooth.

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Figure IV-16. Microtoena delavayi Prain: A. field individuals photographed by Dr. Y.S. Chen in Lijiang County of Yunnan Province (China); B. dichotomous cymes; C. a fruiting stem; D. a calyx with linear bract and one extremely elongated tooth, scale bar = 2 mm; E. a leaf with truncate base and mucronate crenate margin, scale bar = 2 cm.

identified as M. delavayi. The result of principal coordinate analysis of multiple characters shows that the holotype of M. affinis, the four duplicates (Y.P. Zhang 0811) and randomly sampled specimens of M. delavayi are conspecific with each other, and form a distinct entity (Figure IV-19). Therefore, M. affinis should be reduce to a synonym of M. delavayi. Microtoena delavayi var. vera is actually an invalid name (Art. 24. 3), and the autonym M. delavayi var. delavayi must be used instead. D. Prain (1895) described the new variety M. delavayi var. grandiflora, mainly based on its large flower. This variety is accepted in later studies (Hsuan, 1965, Wu & Li, 1977; Li & Hedge, 1994). C.Y. Wu (1959) described M. tenuiflora as a new species, based on the large leaves and slim corolla tube. S.J. Hsuan (1965) recognized

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Map IV-6. Microtoena delavayi Prain.

Figure IV-17. the ‘1-flowered’ cyme from the holotype of M. affnis, which actually consists of one flower with four flower buds. Scale bar = 1 mm.

Figure IV-18. The leaves of M. affinis and M. delavayi Prain: a. small and sparsely crenate leaf from the holotype of M. affinis; b. small and sparsely crenate leaf of new shoot of M. delavayi, sampled from the population D.Y. Hong & Q. Wang, H&W08005 (Tengchong, Yunnan, China); c. large and densely mucronate crenate leaf from of main stem of M. delavayi, sampled from the population D.Y. Hong & Q. Wang, H&W08005 (Tengchong, Yunnan, China). Scale bar = 1 cm.

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Figure IV-19. Principal coordinate analysis of multiple characters of M. affinis and M. delavayi: C. the holotype of M. affinis; D. the types with randomly sampled specimens of M. delavayi; E. the four duplicates of Y.P. Chang 0811.

M. tenuiflora as conspecific with M. delavayi, and reduced the former to a synonym. This treatment is also followed by later studies (Wu & Li, 1977; Li & Hedge, 1994). W.T. Stearn (1983) described M. bhutanica from Bhutan as a new species, but this is also similar to M. delavayi. I have examined all the types involved, and found that all the ‘diagnostic’ characters of these taxa are within the continuous variation range of M. delavayi. Therefore, I here reduce M. delavayi var. grandiflora and M. bhutanica to synonyms of M. delavayi, and also agree with the treatment of S. J. Hsuan (1965) on M. tenuiflora. Additional specimens examined: CHINA, s. loc., s. d., s. coll. 74-19 (SM). SICHUAN, Butuo, Gengbu, grassy slopes, 06 Sep 1979, s. coll. 0896 (SM). Miyi, Pingshan, Longxingzi, grassy slopes, 2300 m, 18 Sep 1978, s. coll. 388 (SM). Mianning, Xinxing, 2000 m, 06 Oct 1978, Mianning Exped. 627 (SM). Muli, 913# field, Zamin, in valley, 2500 m, 24 Aug 1983, Qinghai-Tibet Exped. 13317 (KUN); Baidiao, in forest, 3500 m, 16 Sep 1959, S. K. Wu 3703 (CDBI, KUN, PE); Baidiao to Linjia, ditch side, 2500-2700 m, 12 Sep 1959, S. K. Wu 2591 (KUN); Lilanggangding, forest edge, 2800 m, 31 Jul 1938, T. T. Yü 7517 (A, KUN, PE). YUNNAN, s. loc. s. d., s. coll. s. n. (KUN); s. loc. s. d. M. J. Delavay s. n. (P). Chuxiong, Tou Tsu, stream side, 2400 m, 20 Sep 1939. M. K. Li 246 (KUN). Heqing, Huangping, Junhua, Shangdaping, ditch side, 2450 m, 17 Aug 1963, NW Yunnan-Jinsha River Exped. 6596 (KUN). Jingdong, Nanma, in woods, 2200 m, 17 Jun 1940, M. K. Li 2990 (KUN, WUK); Raw Mau Lun, ditch side, 2400 m, 29 Dec 1939, M. K. Li 2690 (IBSC, KUN, WUK). Songming, Shaodian, Aziying, Guodong, hill side, 25 Aug 1957, B. Y. Qiu 55017 (IBSC, KUN, PE); loc. eodem. 2400 m, 21 Sep 1957, B. Y. Qiu 55154 (PE). Tengchong, Qushi, in woods, 1930-2120 m, 12 Sep 1960, W. Q. Yin 60-1010 (KUN); Qushi, Shuanghe, Maluchong, in thickets, 1470 m, 25 Dec 2008, Q. Wang HW08008 (K, PE, SZ). Weixi, s. loc. in ravine, 2300m, 03 Nov 1933, H. T. Tsai 59995 (A, IBSC, KUN, NAS, PE, SZ); s. loc. in ravine, 2300 m, 04 Nov 1934, H. T. Tsai 63007 (A, KUN,

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NAS, PE, SZ); Cangjiang side, Yanwaguan, 08 Nov 1940, K. M. Feng 8778 (KUN, PE). Shangrila, Jinshajiang side, Shangsuoyi, 2300 m, 18 Nov 1939, K. M. Feng 3396 (KUN, PE). Xundian, Mt. Dabei, grassy slopes, 2600 m, 01 Nov 1940, Y. P. Chang 0811 (IBSC, KUN, PE). Yingjiang, s. loc. in woods, 1700 m, 14 Nov 1958, S. K. Wen 580696 (KUN). Yongde, Xiaoxueshan, in thickets, 3200 m, 20 Aug 2002, s. coll. L5862 (KUN); Wumulong, Yinchang Street, road side, 2400 m, 08 Sep 2003, E. D. Liu & D. C. Yang 260 (KUN). Yulong, s. loc. stone slopes, 3650 m, Sep 1921, G. Forrest 20586 (K, P); Judian, Wuhou, grassy land, 2300 m, 13 Sep 1958, W. T. Wang 247 (KUN); Mt. Yulong, Baishui River, road side, 2900 m, 18 Aug 1956, Yunnan University-Biology department 00812 (IBSC, KUN); Mt. Yulong, Heishui River, 15 Aug 1962, s. coll. s. n. (HITBC); loc. eodem, river side, 10 Jul 1962, A. L. Zhang & S. W. Yu 100913 (KUN); Mt. Yulong, Muzhu ditch, grassy slopes, 2850 m, 10 Jul 1959, s. coll. 22363 (KUN); loc. eodem, stream side, 2800 m, Oct 1970, S. W. Yu s. n. (KUN); Mt. Yulong, Dahuoshan, Gezhifang, in forest, 2700 m, 23 Aug 1961, Y. F. Qi 612615 (KUN). 6. Microtoena wardii Stearn, J. Jap. Bot. 58(1): 11. 1983. Type: Driang Dzong, stream side, 1828 m, 12 Sep 1938, F. Kingdon-Ward 14235 [holotype: BM! (BM 000950462)]. Microtoena delavayi var. amblyodon C.Y. Wu & S.J. Hsuan, Acta Phytotax. Sin. 10: 48. 1965. syn. nov. Type: China, Yunnan, Lanping, in woods, 2900 m, 25 Aug 1933, H.T. Tsai 54088 [holotype: PE! (PE 00031250); isotypes: A! (A00001326), IBSC! (IBSC0583620), KUN! (KUN0217778), NAS! (NAS00223563), PE! (PE00824263), SZ! (SZ00241927)]. Microtoena delavayi var. lutea C.Y. Wu & S.J. Hsuan, Acta Phytotax. Sin. 10: 48. 1965. syn. nov. Type: China, Yunnan, Gingtung [Jingdong], Huangcaoling, Xiaoheping, in woods, 2000 m, 11 Sep 1959, S.K. Shui 4931 [holotype: KUN! (KUN0217786); isotype: KUN! (KUN0217787)]. Microtoena urticifolia var. subedentata Hand.-Mazz., Symb. Sin. 7: 927. 1936. syn. nov. Type: China, Yunnan, Bahan [Pehalo], Nujiang, 2400-2600 m, 26 Sep 1915, HandelMazzetti 8412 [holotype: WU! (WU0060512)]. Herbs perennial. Stems 1–2 m tall, hispid. Leaf petiole 2–10 cm long; leaf blade broadly ovate, 7–12.5 cm long, 5–9 cm broad; base truncate to subcordate; margin mucronate crenate; apex shortly acuminate. Cymes axillary and terminal, dichotomous, lax, sometimes with 1-sided branches; peduncle 1–3 cm long. Bracts linear, 3.6–5.6 mm long; ca. 0.5 mm broad. Calyx 3.3–5.0 mm long at anthesis, puberulent, dilated after anthesis, 5-toothed; calyx teeth 1–1.5 mm long, lanceolate, unequal, one calyx tooth conspicuously elongated. Corolla uniformly yellow to pale yellow, 1.5–2.3 cm long, sparsely hirsutulous outside; corolla tube conspicuous, 0.9–1.5 cm long, tube base slightly dilated; upper corolla lip laterally compressed, apex entire; lower corolla lip 3-lobed, middle lobe subcircular, wider than lateral lobes. Nutlets 1.5–2 mm long, 1–1.5 mm broad, black to brown. Figures. IV-20 & 21.

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Figure IV-20. Microtoena wardii Stearn: a. flowering stem; b. a part of cyme with a dilated fruiting calyx and a flower slightly dilated at base of corolla tube.

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Figure IV-21. Microtoena wardii Stearn: A. field individuals photographed by Dr. B. Liu in Chayu County (Tibet, China); B. a leaf with truncate base and mucronate crenate margin, scale bar = 2 cm; C. flowers in axillary and terminal panicles, with 1-sided branches; D. a pale-yellow corolla with slightly dilated base of corolla tube.

Phenology: Flowering from August to September and fruiting in October. Distribution: This species is distributed Yunnan (Jingdong and Lushui) and Tibet (Chayu) of Southwest China, and North India (Assam) (Map IV-7). Habitat: It grows in forests and at forest margins, at altitudes from 2440 m in Senge Dzong to 2600 m in Beiqingtou, Lianping, Heqing, Yunnan province of China. Note: For the three varieties here treated as new synonyms, M. urticifolia var. subedentata was the first one to be described, mainly based on the entire apex of the upper corolla lip and the ‘short’ cymes. C.Y. Wu (1959) and S.J. Hsuan (1965) did not see the holotype preserved in herbarium WU, and described two other varieties, M. delavayi var. amblyodon and M. delavayi var. lutea. The former was based on the somewhat obtuse apex of the calyx teeth, and the latter was based on the uniformly yellow corolla. However, based on my field observations and examination of specimens, the three varieties all have uniformly yellow to pale yellow corolla and entire apex of

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Map IV-7. Microtoena wardii Stearn.

upper lip just like M. wardii. The ‘small’ cyme of M. urticifolia var. subedentata was just not fully developed, and the slightly obtuse apex of the calyx teeth of M. delavayi var. amblyodon are still within the variation range of M. wardii. Although M. wardii was described later, the three variety names have no priority over the species name. Therefore, I here reduce these variety names to synonyms of M. wardii. Additional specimens examined: CHINA, TIBET, Senge Dzong [Shan Chi], in forest, 2438 m [8000 ft], 16 Aug 1938, F. Kingdon-Ward 14098 (BM). YUNNAN, s. loc. s. d. H.T. Tsai 57468 (KUN); Heqing, Lianping, N. Qingtou, in forest, 2700 m, 18 Aug 1940, R.C. Ching 23689 (PE, KUN); Weixi, Pantiange, Luoshuidong, 27 Oct 1961, G.S. Yang 610123 (KUN) 7. Microtoena urticifolia Hemsl., J. Linn. Soc., Bot. 26: 308. 1890. Type: China, Hubei, Badong, May 1888, A. Henry 7339 [holotype: K! (K000928187); isotype: K! (K000928188)]. Herbs perennial. Stems 1–1.5 m tall, densely hispid, base sometimes woody. Leaf petiole 1.5–10 cm long; leaf blade papery, ovate, spreading hispid, 6.9–7.8 cm long, 4.7–5.4 cm broad; base truncate to subcordate; margin coarsely serrate, teeth sometimes mucronate; apex long-acuminate. Cymes axillary and terminal, dichotomous, lax, usually with 1-sided branches; peduncle 1–4 cm long. Bracts linear, 3.3–3.8 mm long, less than 0.5 mm broad. Calyx 2.8–3.3 mm long at anthesis, puberulent, dilated after anthesis, 5-toothed; calyx teeth lanceolate, apex sometimes hooked, one tooth conspicuously elongated. Corolla pale yellow to white, 1.6–2.7 cm long, sparsely hirsutulous outside; corolla tube 0.8–1.4 cm long; upper corolla lips laterally compressed, apex entire; lower corolla lips 3-lobed, middle lobe obovate, wider than lateral lobes. Nutlets brown, ovoid, smooth. Figure IV-22. Phenology: Flowering from March to April and fruiting in May. Distribution: This species is endemic to Sichuan and Hubei (Badong and Xingshan) Provinces (Map IV-8).

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Figure IV-22. Microtoena urticifolia Hemsl.: a. flowering stem; b. a flower with conspicuously elongated tube; c. a linear bract.

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Map IV-8. Microtoena urticifolia Hemsl.

Habitat: It grows in forests and thickets and at their margins. Additional specimens examined: CHINA, CHONGQING, Qianjiang, Huiqian TV station, ditch side, 1830 m, 18 Aug 1988, Z.C. Zhao 88-2009 (PE). Youyang, Qingkou, Cangling, mountain slopes, 1100 m, 07 Sep 1978, s. coll. 1117 (SM). GUIZHOU, Yinjiang, Xiaojiahe, Shaishiban, in thickets, 1050 m, 06 Sep 1963, Z.B. Jian et al. 31157 (PE). HUBEI, Badong, Feb 1887, A. Henry 2536 (K); Hefeng, s. loc. forest edge, 1230 m, 09 Aug 1988, Y.M. Wang 5952 (PE); s. loc. in woods, 1470 m, 06 Aug 1988, Y.M. Wang 5638 (PE). HUNAN, Sangzhi, s. loc. 28 Aug 1988, Beijing Exped. 4241 (PE); Mt. Badagong, Doupeng Hill, 01 Sep 1988, Beijing Exped. 4384 (PE); Houping, stream sides, 1400 m, 07 Aug 1984, s. coll. 1197 (KUN); Mt. Tianping, path sides, 1450m, 21 Aug 1988, Beijing Exped. 3832 (PE); loc. eodem, in forest, 1200 m, 18 Jul 1988, L.Q. Li 133 (PE). SICHUAN, s. loc. Apr 1937, E.H. Wilson 4372 (A, BM, IBSC). 8. Microtoena albescens C.Y. Wu & S.J. Hsuan, Acta Phytotax. Sin. 10: 49. 1965. Type: China, Guizou, Qingzhen, Moupo, 07 Oct 1935, S.W. Teng 1330 [holotype: IBSC! (IBSC 0005113)]. Herbs perennial. Stems erect, ca. 0.6 m tall, hispid. Leaf petiole 1–5 cm long; leaf blade broadly ovate, 5.2–5.7 cm long, 4.1–4.4 cm broad, sparsely hirsute; base truncate; margin mucronate crenate; apex shortly acuminate. Cymes axillary, dichotomous, lax; peduncle inconspicuous, 3–5 mm long. Bracts linear, ca. 5.3 mm long, ca. 0.5 mm broad. Calyx glandular puberulent, ca. 4.8 mm long at anthesis, dilated after anthesis, 5-toothed; calyx teeth lanceolate, one calyx tooth conspicuously elongated. Corolla uniformly white, ca. 2.1 cm long; corolla tube conspicuous, ca. 1.2 cm long; apex of upper corolla lip entire; lower lips 3-lobed, middle lobe subcircular, wider than lateral lobes. Nutlets unknown. Figure IV-23.

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Figure IV-23. Microtoena albescens C. Y. Wu & S.J. Hsuan: a. flowering stem; b. a flower with conspicuous tube; c. a linear bract; d. a calyx with one tooth conspicuously elongated.

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Phenology: Flowering in October. Distribution: This species is endemic to Qingzhen of Guizhou province (Map IV-9).

Map IV-9. Microtoena albescens C. Y. Wu & S. J. Hsuan.

Habitat: It is only recorded for the holotype, from forest and thicket margins. Additional specimens examined: CHINA, GUIZHOU, s. loc. 1931, S.S. Sin 50809 (IBSC). 9. Microtoena stenocalyx C.Y. Wu & S.J. Hsuan, Acta Phytotax. Sin. 10: 49. 1965. Type: China, Yunnan, Zhengkang, snow range, in thickets, 2400 m, 25 Jul 1938, T.T. Yü 16966 [holotype: KUN! (KUN0217858); isotypes: A! (A00001328), KUN! (KUN0217859), PE !(PE00824295, PE00824327)]. Herbs perennial. Stems erect, 1–2 m tall, densely hirsute, base sometimes woody. Leaf petiole 1–4 cm long; leaf blade oblong-ovate, 4.7–7.8 cm long, 3–4.9 cm broad, densely hirsute; base truncate to subcordate; margin mucronate crenate; apex long-acuminate. Cymes axillary and terminal, dichotomous, in compact, narrow panicles; lower panicles with peduncle 1–3 cm long, peduncle inconspicuous in upper inflorescences. Bracts lanceolate, 5–6.9 mm long, 1.7–2.4 mm broad. Calyx 4.8–5.8 mm long at anthesis, puberulent, dilated after anthesis, 5-toothed; calyx teeth linear-lanceolate, unequal, one tooth conspicuously elongated. Corolla uniformly pale yellow, 1.9–2.5cm long, sparsely hirsutulous outside; corolla tube conspicuous, 1–1.4cm long; upper corolla lips laterally compressed, apex entire; lower corolla lips 3-lobed, middle lobe subcircular, wider than lateral lobes. Nutlets dark brown, 1.5–2 mm long, 1–1.2 mm broad, smooth. Figures. IV-24(a–d) & 25.

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Figure IV-24. Microtoena stenocalyx C.Y. Wu & S.J. Hsuan (a–d): a. flowering stem; b. a flower with conspicuous tube; c. a lanceolate bract; d. a dissected calyx with linear-lanceolate teeth. Microtoena miyiensis C.Y. Wu & H.W. Li. (e–f): e. a linear-lanceolate bract; f. a dissected calyx with one elongated tooth.

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Figure IV-25. Microtoena stenocalyx C.Y. Wu & S.J. Hsuan: A. field individuals photographed by the author in Fugong County (Yunnan Province, China); B. compact cymes in a sub-spikelike panicle; C. densely hirsute stem; D. profile of a flower with conspicuous tube; E. front of a flower with laterally compressed upper lip and subcircular middle lobe of lower lip.

Phenology: Flowering from July to September and fruiting from September to October. Distribution: This species is restricted to Zhenkang, Fugong, Gongshan, Yongde of Yunnan Province (Map IV-10). Habitat: It grows at forest and thicket margins, at altitudes from 2000 m in Fugong to 2520 m in Yongde of Yunnan province.

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Map IV-10. Microtoena stenocalyx C. Y. Wu & S. J. Hsuan.

Additional specimens examined: CHINA, YUNNAN, s. loc. s. d. Gaoligong Shan Biodiversity Survey 2003 18871 (GH). Fugong, Guquan, in forest, 1700-2100 m, 16 Jul 2013, X.H. Jin et al. ST0278 (PE); in forest, 2000 m, 18 Sep 1933, H.T. Tsai 56519 (A, IBSC, KUN, PE, SZ). Gongshan, to Dulongjiang, in woods, 2200-2400 m, 21 Jul 1982, Qinghai-Tibet Exped. 8280 (HITBC, KUN). Lushui, Luobenzhuo, Ega, 2300 m, 18 Jul 2013, X.H. Jin et al. ST0464 (PE). Yongde, Wumulong, Ganhe, Xiaoheihuo, ditch side, 2520 m, 15 Jul 2003, S.S. Zi 155 (KUN). 10. Microtoena miyiensis C.Y. Wu & H.W. Li, Fl. Reipubl. Popularis. Sin. 66: 579. 1977. Type: China, Sichuan, Miyi, Puwei, Ertaizi, Longdong, in woods, 2500 m, 17 Aug 1958, Sichuan Medic. Plant Resources Surv. 11287 [holotype: SM! (SM64)]. Herbs perennial. Stems erect, ca. 45 cm tall, densely hispid. Leaf petiole slender, 1.5–8 cm long; leaf blade papery, rhombic-ovate, 3.5–7 cm long, 2.4–4.7 cm broad, adaxially sparsely appressed bristly, strigose on veins, abaxially densely white bristly, with scattered glands; base cuneate; margin coarsely serrate, mucrones absent; apex acuminate. Cymes dichotomous, compact in terminal ovoid panicles, densely white pilose and strigose; peduncle inconspicuous. Bracts linear-lanceolate, ca. 4.2 mm long, ca. 1 mm broad. Calyx ca. 3.5 mm long at anthesis, densely pilose, 5-toothed; calyx teeth lanceolate, unequal, one calyx tooth extremely elongated, ca. 3× as long as other teeth. Corolla white to pale yellow, ca. 2.2 cm long, puberulent and glandular outside; corolla tube slender, conspicuous, ca. 1.4 cm long; upper corolla lip laterally

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compressed, apex entire; lower corolla lip 3-lobed, middle lobe obovate. Nutlets unknown. Figure IV-24(e–f). Phenology: Flowering in August, fruiting phase unknown. Distribution: This species is endemic to Miyi of Sichuan province (Map IV-11).

Map IV-11. Microtoena miyiensis C. Y. Wu & H. W. Li.

Habitat: It is only recorded for the holotype growing in forests, at altitude of 800 m in Miyi of Sichuan province. 11. Microtoena megacalyx C.Y. Wu, Acta Phytotax. Sin. 8: 48. 1959. Type: China, Yunnan, Wenshan, in forest, 2000-2200 m, 19 Aug 1947, K.M. Feng 11344 [holotype: PE! (PE 00031247); isotypes: A! (A 00001327), KUN! (KUN 0484541, KUN 0484542)]. Herbs perennial. Stems erect, 0.3–1.5 m tall, puberulent. Leaf petiole 2–10 cm long; leaf blade ovate, 8.6–10.6 cm long, 5.2–6.4 cm broad, sparsely minutely hispid; base cuneate; margin mucronate crenate; apex acuminate. Cymes axillary and terminal, dichotomous, lax; peduncle 1–5 cm long. Bracts lanceolate, 3.5–5 mm long, 0.9– 1.3 mm broad. Calyx campanulate, glandular puberulent, 4.6–5.9 mm long at anthesis, strongly dilated and netted after anthesis, 5-toothed; calyx teeth subequal, 1.4–1.8 mm long, triangular-subulate, teeth usually with a hooked apex. Corolla uniformly white to pale yellow, 1.7–2.7 cm long, puberulent outside; corolla tube conspicuous, 0.9–1.4 cm long; apex of upper lip entire; lower lip 3-lobed, middle lobe subcircular, wider than lateral lobes. Nutlets dark brown, 1.4–1.8 mm long, 1–1.3 mm broad, smooth. Figure IV-26. Phenology: Flowering from August to October and fruiting from October to November.

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Figure IV-26. Microtoena megacalyx C.Y. Wu: a. flowering stem; b. a flower with conspicuous tube; c. a lanceolate bract; d. a dissected calyx with subequal teeth; e. a dilated and netted calyx; f. a dissected dilated calyx.

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Distribution: This species is distributed in Yunnan (Wenshan county) and Guizhou (Hezhang county) provinces of China, and Vietnam (Map IV-12).

Map IV-12. Microtoena megacalyx C. Y. Wu.

Habitat: It grows at woods and thickets margins, on ravine grassy slopes, at altitudes from 1500 m in Caishentang, Hezhang of Guizhou province to 2100 m in Wenshan of Yunnan province. Additional specimens examined: CHINA, GUIZHOU, Hezhang, Caishentang, grassy slopes, 1500 m, 11 Oct 1957, P. H. Yu 1261 (KUN). HUBEI, Yichang, Dalaoling, Laochangbu, Zhucaogou, in forest, 1650 m, 07 Sep 2007, C. M. Zhao EX0551 (PE). YUNNAN, Wenshan, Zuodi Volcano, ditch side, 2100 m, 27 Sep 1958, H. T. Tsai 58-8185 (KUN). 12. Microtoena moupinensis (Franch.) Prain, Bull. Soc. Bot. France 42: 426. 1895. = Clerodendrum moupinense Franch., Nouv. Arch. Mus. Hist. Nat., ser. 2, 10: 68. 1887. Type: China, Sichuan, Baoxing, 1870, M.I. David s. n. [lectotype designated here: P! (P 00743579)]. Microtoena griffithii Prain, Journ. Asiatic Soc. Bengal 59(2): 310. 1890. Type: India, Assam, probably Mishmi Hills, Apr 1863, Griffith 4059 [lectotype designated here: K! (K 000928194)]. Microtoena longisepala C.Y. Wu ex S.J. Hsuan, Acta Phytotax. Sin. 10: 55. 1965. Type: China, Sichuan, Butuo, Jiaoji River, in woods, 2300 m, 24 Aug 1959, Sichuan Econ. Plant Exped. 5842 [holotype: KUN! (KUN 0484540); isotypes: PE! (PE 00031248, PE 00824276), SM! (SM 0161009)]. Microtoena maireana Hand.-Mazz., Symb. Sin. 7: 927. 1936. Type: China, Yunnan,

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Huize, Zhehai, 2550 m, s. d. E.E. Maire s. n. [holotype: W! (W 5977)]. Herbs perennial. Stems erect, 0.6–1.6 m tall, base sometimes woody, sparsely puberulent. Leaf petiole 1–11 cm long; leaf blade oblong-ovate, triangular-ovate, ovate to broadly ovate, 2.9–13.6 cm long, 1.9–9.2 cm broad, sparsely puberulent, hispid, or nearly glabrous; base cuneate, truncate or cordate, margin coarsely serrate to mucronate crenate; apex acuminate to shortly caudate-acuminate. Cymes axillary and terminal, dichotomous, lax, slightly to very compact in spikelike panicles, sometimes with 1-sided branches; peduncle inconspicuous to 4 cm long. Bracts usually linear, 3.2–7.3 mm long, 0.3–0.8 mm broad. Calyx 3.2–6.9 mm long at anthesis, puberulent, dilated after anthesis, 5-toothed; calyx teeth triangular-lanceolate, linear-lanceolate to subulate, subequal, with apex usually hooked. Corolla yellow to pale yellow, 1.6–3.7 cm long, sparsely hirsutulous outside; corolla tube conspicuous, 1.1–2.5 cm long, sometimes slender; upper corolla lip laterally compressed, apex entire; lower corolla lip 3-lobed, middle lobe subcircular, wider than lateral lobes. Nutlets dark brown to black, 1.6–2 mm long, 1–1.5 mm broad, smooth. Figures. IV-27 & 28. Phenology: Flowering from August to September and fruiting in October. Distribution: This species is widely distributed in Hubei, Sichuan, Yunnan provinces of China (Map IV-13). Note: D. Prain (1890) used ‘probably Mishmi hill’ of Assam in its protologue. However, the location of M. griffithii is not indicated clearly on its holotype, and might be somewhere near to S Tibet of China. Since the location is ambiguous, the type locality of M. griffithii is not marked on Map IV-13. Habitat: It grows at forest and thicket margins, or on open grassy slopes or stony slope, at altitudes from 800 m in Shennongjia of Hubei province to 2850 m in Shimian of Sichuan province. Note: The results of standard deviation analysis in the Chapter II has already shown that many characters of M. moupinensis are highly variable (with wide variation range) and unstable (with high standard deviation), both at population and species level. In fact, M. moupinensis can be treated as a complicated species complex. The type of inflorescence is usually uniform in Microtoena, such as the lax cymes in M. patchoulii and the finely compact cymes in M. esquirolii. However, the cymes of M. moupinensis is highly variable from strongly compact in spikelike panicles to extremely lax or with 1-sided branches (Figures. 28E & 29A–C, G–I). The cultivation experiments of sampled species (M. delavayi & M. esquirolii) of Microtoena (Figure II-10) has shown that the leaf base of Microtoena is usually uniform from young individuals to flowering individuals (Figure II-10). Based on my field observation and specimen examination, most species of Microtoena have stable shape of leaf base. Again, M. moupinensis presents a special case as many types of leaf base, such as cordate, truncate, cuneate and intermediate types, can be observed in M. moupinensis, even within the same population (Figures. 28D & 29D–F).

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Figure IV-27. Microtoena moupinensis (Franchet) Prain: a. flowering stem; b. a leaf; c. a flower with conspicuous and slightly slender tube; d. a linear bract; e. a dissected calyx; f. a dilated calyx with hooked teeth.

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Figure IV-28. Microtoena moupinensis (Franchet) Prain: A. field individuals photographed by the author in type locality (Baoxing County, Sichuan Province, China); B. lax cyme with flowers; C. sparsely hispid and puberulent stem; D. two leaves sampled from H&W09007 (Q. Wang & D.Y. Hong, Baoxing County, Sichuan Province, China) with cuneate and cordate base respectively, scale bar = 2 cm; E. variable inflorescences from the same individual of Microtoena moupinensis, scale bar = 1 cm.

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Map IV-13. Microtoena moupinensis (Franchet) Prain.

Figure IV-29. The variation of characters within a sampled population (Q. Wang & D.Y. Hong, H&W09007, Baoxing County, Sichuan Province, China) of Microtoena moupinensis (Franchet) Prain: A–C. cymes from compact, slightly compact to lax, scale bar = 1 cm; D–F. leaf bases from subcordate, subtruncate to cuneate, scale bar = 2 cm; G. ‘M. maireana’ kind of M. moupinensis, scale bar = 2cm; H. ‘M. griffithii’ kind of M. moupinensis, scale bar = 2cm; I. ‘M. longisepala’ kind of M. moupinensis, scale bar = 2cm.

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In addition, M. moupinensis is also highly variable and unstable in indumentum, leaf size and shape, leaf margin, bract length, calyx tooth size and shape, corolla length as well as corolla tube length. Therefore, it does not surprise me that individuals from the same population (D.Y. Hong & Q. Wang, H&W09007, collected from the type locality of M. moupinensis) can be identified as M. moupinensis, M. griffithii, M. maireana and M. longisepala respectively. As a result, I here reduce M. griffithii, M. maireana and M. longisepala to synonyms of M. moupinensis. Additional specimens examined: CHINA, HUBEI, Shennongjia, Mt. Guanshi, Shiban Ditch, in forest, 1750 m, 11 Aug 1976, Hubei Shennongjia Exped. 11130 (PE); Songluo, Biandan River, river side, 800 m, 31 Aug 1976, Hubei Shennongjia Exped. 2494 (PE). SICHUAN, s. loc. 1933, T. H. Tu 4607 (IBSC); s. loc. s. d. K. L. Chu 4601 (NAS, PE). Baoxing, Suburb, in thickets, 1560 m, 22 Sep 2009, Q. Wang HW09008 (K, PE, SZ); Dengchi Ravine, in thickets, 1470 m, 22 Sep 2009, Q. Wang HW09007 (K, PE, SZ); loc. eodem, in thickets, 1500 m, 30 Aug 2010, Q. Wang XZ015 (K, PE, SZ); Dengchi Ravine, Shijiancao, grass land, 2000 m, 26 Sep 1936, K. L. Chu 3951 (IBSC, PE, SZ); Donghe, Dadi Ditch, in forest, 1500 m, 15 Oct 1958, S. C. Chang & Y. X. Ren 07624 (PE); Donghe, Dadi Ditch, Timberyard, in forest, 2200 m, 12 Aug 1961, J. S. Ying 10091 (PE); Ganyang, baiqianchang, 1220 m, 12 Aug 1936, K. L. Chu 3558 (IBSC, PE); Ganyang, Shijiaotou, grass land, 2000 m, 22 Aug 1936, K. L. Chu 3633 (BM, IBSC, PE, SZ); Leimu Ravine, in thickets, 1450 m, 24 Aug 1963, K. J. Guan & W. T. Wang 2525 (PE); Yemaoping to Hetaoping, 1500 m, 30 Aug 1938, Y. Pei 8121 (NAS, PE). Beichuan, Qingpian, Xiaozhaizi Ravine, Big Geoge, river side, 1550 m, 13 Aug 2007, D. H. Zhu et al. 4763 (HX). Butuo, Zeluo, grassy slopes, 2400 m, 09 Aug 1979, s. coll. 0713 (SM). Dechang, Cida, Heping, Mazhailing, stream side, 2000 m, 31 Jul 1978, Hu 0275 (SM). Dujiangyan, Mt. Zhaogong, in thicket, 2200 m, 09 Aug 2006, Z. B. Feng et al. 20060455 (HX); Longxi River, Mashanping to Qinglongzui, on moist slope, 1400 m, 11 Sep 1988, D. E. Boufford & B. Bartholomew 24821 (A, KUN). Ganluo, Aer, Luomu, grassy slopes, 1500 m, 30 Jul 1979, s. coll. 742 (SM). Lushan, Kuaile, Xiaoxing 7th Group, ditch side, 27 Aug 1978, Lushan Exped. 78-0834 (SM). Luding, Moxi, E Gongga, Hailuo Ravine, 2130 m, 15 Aug 1997, D. E. Boufford et al. 27189 (A, CDBI); Moxi, Gonghe 4th Group, in forest, 1800 m, 30 Jul 1980, G. R. Xu 23295 (CDBI, IBSC). Meigu, Majing, Shuanghekou, stream side, 2230 m, 02 Aug 1959, Z. T. Guan 9118 (PE). Mianning, Daqiao, Xiaolin Ditch, in valley, 1950 m, 26 Feb 1959, S. K. Wu 1993 (PE); Mt. Caogu, ditch side, 2100 m, 26 Oct 1993, Z. C. Zhao 4777 (PE). Ningnan, Xingfu, in forest, 2800 m, 27 Sep 1978, Ningnan Exped. 757 (SM). Pingwu, Sier, Yangliuba, Yaozi Ditch, in forest, 2380 m, 28 Aug 1958, H. L. Tsiang 11053 (IBSC, SZ). Puge, Xiatiandi, road side, 2700 m, 14 Sep 1979, Puge Exped. 854 (SM); loc. eodem, road side, 2700 m, 14 Sep 1979, Puge Exped. 879 (SM). Shifang, s. d., s. coll. 0941 (SM). Shimian, s. loc. 1955, C. C. Hsieh 42606 (IBSC, PE); s. loc. 1955, C. C. Hsieh 42137 (PE, SZ); Wajiao, Gonggangping, 2850 m, 22 Aug

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1978, Shimian Exped. 78-1016 (SM); Caoke, Tanyao Ditch, ditch side, 2300 m, 30 Aug 1978, Shimian Exped. 78-1151 (SM); Zhongkang Lumbering Team, ditch side, 2500 m, 20 Sep 1978, Shimian Exped. 78-1252 (SM). Tianquan, Laba River, grassy slopes, 1500 m, 22 Aug 2004, X. Y. Xiong & H. M. Shao T159 (HX); New Ditch, road side, 1500 m, 02 Sep 2012, D. Y. Hong et al. H12056 (PE); New Ditch, Mapiantang, in valley, 1800 m, 05 Sep 1963, K. J. Guan & W. T. Wang 3219 (PE); Tianquanzheng River, Yinchang District, 07 Aug 1954, W. P. Fang & W. K. Hu 51-1117 (SZ, WUK). Yanyuan, Shengou, ditch side, 24200 m, 17 Sep 1979, s. coll. 0911 (SM). YUNNAN, Daguan, Mugan Forest Centre, Wife Cliff, in woods, 1900 m, 17 Aug 1972, NE Yunnan Exped. 39 (KUN). Qiaojia, Dongping, Babao Cave, 2000-2200 m, 18 Aug 2003, H. Wang et al. 03-0951 (KUN) 13. Microtoena prainiana Diels, Bot. Jahrb. Syst. 29: 556. 1900. Type: China, Chongqing, Nanchuan, Qianniuping, 06 Sep 1891, Erdhügel 788 [holotype: K! (K 11146)]. Herbs perennial. Stems erect, 0.6–1.2 m tall, subglabrous. Leaf petiole 3–9 cm long; leaf blade ovate, 5–11.6 cm long, 3.1–7.2 cm broad, sparsely puberulent; base cuneate; margin coarsely serrate; apex acuminate. Cymes axillary and terminal, strongly compact in ovoid panicle; peduncle 1–3 cm long. Bracts ovate, 4.1–5.4 mm long, 1.8–2.4 mm broad. Calyx 3.7–4.7 mm long at anthesis, glabrous, dilated after anthesis, 5-toothed; calyx teeth subulate, subequal, 1.4–1.8 mm long, apex conspicuously hooked. Corolla uniformly pale yellow, 1.6–2.5 cm long, hirsutulous outside; corolla tube conspicuous, 0.9–1.5 cm long; upper corolla lips laterally compressed, apex entire; lower corolla lips 3-lobed, middle lobe subcircular, wider than lateral lobes. Nutlets brown to black, obovoid, 1.5–1.9 cm long, 1–1.3 cm broad, smooth. Figures. IV-30 & 31. Phenology: Flowering from July to August and fruiting in September. Distribution: This species is distributed in Nanchuan of Chongqing City, and the adjacent Sichuan, Yunnan, Guizhou Province (Map IV-14). Habitat: It grows in forests, or at forest and thicket margins, at altitudes from 1050 m in Jinfo mountain, Nanchuan of Chongqing to 2300 m in Tiantai, Jinyang of Sichuan province. Additional specimens examined: CHINA, CHONGQING, Fuling, Chepan, Yangziping, in woods, 1300 m, 11 Aug 1979, X. Z. Zhu 1028 (SM). Nanchuan, s. loc., 18 Jul 1957, s. coll. s. n. (SM); s. loc., Sep 1978, s. coll. s. n. (SM); Mt. Jinfo, s. d., M. Tseng 48 (PE); loc. eodem, s. d., M. Tseng 856 (PE); loc. eodem, in forest, 1600 m, 26 Aug 2009, C. L. Xiang 340 (KUN); Mt. Jinfo, Huangcaoping, in woods, 1700 m, 10 Sep 2012, Q. Wang H12079 (PE); Mt. Jinfo, Lvcaoping, Nian Ditch, in forest, 1300-1500 m, 21 Jul 1979, s. coll. 0829 (SM); Mt. Jinfo, Shuiyuan, Santang Ditch, forest edge, 1002 m, 20 Aug 1957, G. F. Li 63622 (IBSC, KUN, SZ); loc. eodem, ditch side, 1050 m, 12 Sep 1957, G. H. Hsiung & Z. L. Chow 93340 (IBSC, KUN, PE, SZ);

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Figure IV-30. Microtoena prainiana Diels: a. flowering stem; b. a flower with conspicuous tube; c. dissected corolla; d. dissected calyx; e. style; f. immature and mature anthers; g. nutlet; h. bract.

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A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

Figure IV-31. Microtoena prainiana Diels: A. field individuals photographed by the author (Mt. Jinfo, Nanchuan County, Chongqing, China); B. compact cymes in an ovoid panicle; C. flowers; D. a fruiting panicle.

Mt. Jinfo, Mazui, Xujiawan, ditch side, 1400 m, 24 Aug 1957, G. F. Li 63726 (PE, SZ); Mt. Jinfo, Tuangudeng, 1530 m, 28 Aug 1957, G. F. Li 63841 (KUN, PE, SZ); Mt. Jinfo, Phenix Temple, mountain slopes, 2000 m, 10 Aug 1957, G. H. Hsiung & Z. L. Chow 92513 (PE, SZ); Mt. Jinfo, Phenix Temple to Linjiaba, Liuqing Hill, grass slopes, 1800 m, 14 Aug 1957, G. H. Hsiung & Z. L. Chow 92690 (IBSC, PE, SZ); Xiaohe, Lanba, Zengjia Cliff, forest edge, 1560 m, 13 Jul 1957, G. H. Hsiung & Z. L. Chow 91984 (PE, SZ); Xiaohe, Baojiaping, forest edge, 1700 m, 19 Jul 1957, G. H. Hsiung & Z. L. Chow 95257 (SM). GUIZHOU, Mt. Duopeng, mountain slope, 1800-2500 m, 02 Aug 1931,

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T. S. Kang 50809 (NAS). Dafang, Baina, Mt. Jiulong, mountain slopes, 2000 m, 16 Aug 1959, Bijie Exped. 903 (KUN, PE); loc. eodem, mountain slopes, 2000 m, 18 Aug 1959, Bijie Exped. 993 (PE). Leishan, meadow of Mt. Leigongin woods, 26 Aug 1959, S Guizhou Exped. 3636 (KUN, PE). Nayong, Juren, mountain slopes, 2000 m, 02 Aug 1959, Bijie Exped. 670 (PE). SICHUAN, Huidong, 4th District reservoir, stream side, 2400 m, 31 Aug 1978, Huidong Exped. 0685 (SM); 4th District, Yuejin, in woods, 2300 m, 11 Aug 1978, Huidong Exped. 0504 (SM). Jinyang, Tiantai, 2300 m, 07 Aug 1978, s. coll. 504 (SM); loc. eodem, 2100 m, 03 Aug 1978, s. coll. 248 (SM). Shimian, s. loc., 1955, C. C. Hsieh 42079 (IBSC, PE, SZ). YUNNAN, Yongshan, s. loc., in ravine, 1400 m, 13 Jul 1932, H. T. Tsai 51139 (A, KUN, NAS, PE, SZ, WUK).

Map IV-14. Microtoena prainiana Diels.

14. Microtoena muliensis C.Y. Wu & S.J. Hsuan, Acta Phytotax. Sin. 10: 50. 1965. Type: China, Sichuan, Muli, Baidiao to Linjia, in forest, 2700m, 12 Sep 1959, S.K. Wu 2589 [holotype: PE; isotypes: CDBI!(CDBI 0172354), KUN! (KUN 0484543, KUN 0484544)]. Herbs perennial. Stems erect, 0.8–1.5 m tall, glabrous, base sometimes woody. Leaf petiole 3–6 cm long; leaf blade oblong-ovate, 7–12.4 cm long, 3.3–5.9 cm broad, subglabrous; base cuneate; margin mucronate crenate; apex shortly caudate-acuminate. Cymes axillary and terminal, dichotomous, lax, usually with 1-sided branches; peduncle 1.5–6 cm long. Bracts linear, 2.8–3.9 mm long, less than 0.5 mm broad. Calyx 3–5 mm long at anthesis, puberulent, dilated after anthesis, 5-toothed; calyx teeth triangular, subequal, 1.1–1.9 mm long, not hooked. Corolla yellow, 1.6–2.6 cm long, hirsutulous outside; corolla tube conspicuous, 0.8–1.4 cm long; upper corolla lips laterally compressed, apex entire; lower corolla lips 3-lobed, middle lobe circular, wider than lateral lobes. Nutlets brown to black, ovoid, 1.2–1.7 cm long, 0.9–1.1 cm broad, smooth. Figure IV-32.

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A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

Figure IV-32. Microtoena muliensis C. Y. Wu & S. J. Hsuan: a. flowering stem; b. flower with conspicuous tube; c. calyx.

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107

Phenology: Flowering from August to October and fruiting in November. Distribution: This species is distributed in Muli, Huili, Junlian of Sichuan Province, and Lushui of Yunnan Province (Map IV-15).

Map IV-15. Microtoena muliensis C. Y. Wu & S. J. Hsuan.

Habitat: It grows at forest and thicket margins, at altitudes from 1300 m in Huge snow mountain of Junlian to 2380 m in Mazong of Huili of Sichuan province. Additional specimens examined: CHINA, s. loc., s. d., s. coll. s. n. (SM); Luchanglang, 12 Oct 1978, s. coll. s. n. (SM). SICHUAN, Huili, Taiping, Mazong, in forest, 2380 m, 18 Aug 1978, Taiping Exped. 0486 (SM). Muli, Sept. 1922, G. Forrest 22408 (K, P). YUNNAN, Lushui, Yaojiaping to Jiangqiao, Gutan River, in valley, 2200 m, 21 Aug 1964, S. K. Wu 8496 (HITBC, KUN). 15. Microtoena omeiensis C.Y. Wu & S.J. Hsuan, Acta Phytotax. Sin. 10: 51. 1965. Type: China, Sichuan, Mt. Emei, Zhanglaoping, 20 Aug 1952, C.H. Hsiung et al. 32020 [holotype: SZ! (SZ 00241919); isotypes: IBSC! (IBSC 0005114), PE! (PE 00824299)]. Herbs perennial. Stems erect, 1–2 m tall, sparsely puberulent. Leaf petiole 3–15 cm long; leaf blade papery, triangular-ovate, 5.5–11.1 cm long, 3.4–6.9 cm broad, sparsely hispid, usually flushed with purple on the abaxial leaf surface; base truncate to cuneate; margin mucronate crenate; apex caudate-acuminate. Cymes axillary, dichotomous, lax; peduncle 1–2 cm long. Bracts linear, 2.8–3.9 mm long, less than 0.5 mm broad. Calyx 3.7–4.8 mm long at anthesis, glandular, dilated after anthesis, 5-toothed; calyx teeth triangular, 1.2–1.5 mm long, one tooth slightly elongated. Corolla uniformly yellow to pale yellow, 1.6–2.8 cm long, hirsutulous outside; corolla tube conspicuous, 1–1.7 cm long; upper corolla lips laterally compressed, apex entire; lower corolla lips 3-lobed, middle lobe subcircular, wider than lateral lobes. Nutlets brown to black, obovoid, 1.5–2 mm long, 1–1.4 cm broad, smooth. Figure IV-33.

108

A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

Figure IV-33. Microtoena omeiensis C. Y. Wu & S. J. Hsuan: a. flowering stem; b. stem base; c. a flower with conspicuous tube; d. a linear bract; e. fruiting calyx; f. dissected fruiting calyx; g. dissected calyx at anthesis.

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Phenology: Flowering from July to September and fruiting in October. Distribution: This species is endemic to Emei Mountain and adjacent region of Sichuan province (Map IV-16).

Map IV-16. Microtoena omeiensis C. Y. Wu & S. J. Hsuan.

Habitat: It grows on grassy slopes and at thicket margins, at altitudes from 1300 m in Luobodihan, Emei Mountain to 2100 m in Meigu, Sichuan province. Additional specimens examined: CHINA, SICHUAN, s. loc., 1930, F.T. Wang 22040 (KUN, NAS). Emeishan, Mt. Emei, road side, 1830 m, 06 Aug 1928, W.P. Fang 2640 (GH, NAS, K, P); loc. eodem, 1936, W.C. Cheng 6509 (NAS, PE); Mt. Emei, Xianfeng Temple, 1600 m, 11 Aug 2009, H. Peng et al. 113 (KUN); Mt. Emei, Longchi, Jinwu, Minzhu 5th Group, ditch side, 13 Jul 1978, s. coll. 1301 (SM); Mt. Emei, Jingshui, 13 Aug 1978, Mt. Emei Exped. 1827 (SM); Mt. Emei, Oct 1939, L.Y. Lin 1148 (IBSC); Mt. Emei, Jiulaodong, 1700 m, 18 Aug 1957, K.H. Yang 56702 (IBSC, KUN, PE, SZ); Mt. Emei, Yuxian Temple, 29 Aug 1957, S.Y. Chen et al. 4226 (NAS, SM, SZ); Mt. Emei, Jiulaodong, grassy slope, 12 Aug 1935, Y.Y. Ho 6014 (NAS); Mt. Emei, Doushiti, grassy slope, 25 Aug 1938, T.Y. Chow & K.J. Hsu 343 (NAS); Mt. Emei, Oct 1936, W.C. Cheng 6777(PE); Mt. Emei, Zhanglaoping, 28 Aug 1939, C.W. Yao 4842 (PE); Mt. Emei, 18 Aug 1928, W.P. Fang 3132 (PE); Mt. Emei, Luobodidang, 1300 m, 11 Aug 1956, Y.H. Tao 51446 (SZ). Gulin, Deyue, Hongqi, Shisunping, 1400 m, 03 Aug 1976, s. coll. 878 (SM). Junlian, Luomurou, Jiefang, Big Snow Mountain, 1300 m, 05 Sep 1964, M.F. Zhong & S.G. Tang 1033 (SM). Meigu, s. loc., 2100 m, 27 Jun 1979, Meigu-Puge Exped. 0310 (SM). Yingjing, Paocaowan, stream side, 2000 m, 14 Sep 1978, Yingjing Exped. 78-0837 (SM).

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A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

16. Microtoena brevipedunculata (C.Y. Wu & S.J. Hsuan) Q. Wang. comb. & stat. nov. Basionym: Microtoena urticifolia var. brevipedunculata C.Y. Wu & S.J. Hsuan, Acta Phytotax. Sin. 10: 54. 1965. Type: China, Hunan, Qianyang, Mt. Taipo, 08 Aug 1954, T.T. Lee 2985 [holotype: PE! (PE 00824264); isotype: IBSC! (IBSC 0005117)]. Herbs perennial. Stems erect, 0.6–1 m tall, puberulent. Leaf petiole 2–6 cm long, sometimes slender; leaf blade papery, triangular-ovate, 7.3–10 cm long, 5.5–7.6 cm broad, hirsutulous; base truncate to subcordate; margin mucronate crenate; apex shortly acuminate. Cymes axillary, dichotomous, lax; peduncle 0.5–1 cm long. Bracts linear, ca. 4.1 mm long, less than 0.5 mm broad. Calyx 3.5–3.8 mm long at anthesis, puberulent, dilated after anthesis, 5-toothed; calyx teeth lanceolate, subequal, ca. 1.4 mm long, not hooked. Corolla pale yellow to white, ca.1.8 cm long, sparsely hirsutulous outside; corolla tube ca. 1 cm long; upper corolla lips laterally compressed, apex entire; lower corolla lips 3-lobed, middle lobe obovate, wider than lateral lobes. Nutlets brown, ovoid, smooth. Figure IV-34. Phenology: Flowering from August to September and fruiting from September to October. Distribution: This species is endemic to Hunan province (Map IV-17).

Map IV-17. Microtoena brevipedunculata (C. Y. Wu & S. J. Hsuan) Q. Wang.

Habitat: It grows in forest, and at forest and thicket margins, at an altitude of 800m in Zhijiang, Hunan Province of China. Note: This species was described as M. urticifolia var. brevipedunculata, a new variety of M. urticifolia (Hsuan, 1965). However, it has long and slender leaf petiole, mucronate crenate leaf margin, shortly acuminate leaf apex, axillary cymes, short peduncle, subequal, lanceolate and straight calyx teeth, which are all distinct from those of M. urticifolia. Therefore, I here raise it as an independent species, M. brevipedunculata. Additional specimens examined: CHINA, HUNAN, Zhijiang, Muyexi, Jiaoyitang, in valley, 800 m, 30 Sep 1988, Wuling Exped. 1610 (IBSC).

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Figure IV-34. Microtoena brevipedunculata (C. Y. Wu & S. J. Hsuan) Q. Wang: a. flowering stem; b. flower with conspicuous tube; c. dissected calyx; d. calyx; e. root system.

112

A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

17. Microtoena nepalensis Stearn, Enum. Flowering Pl. Nepal 3: 158. 1982. Type: Nepal, Tinjure Danda, in wet forest, 2280 m, 06 Sep 1967, Williams & Stainton 8384 [holotype: BM! (BM 000521969); isotype: K!]. Herbs perennial. Stems erect, ca. 0.6 m tall, puberulent. Leaf petiole 1–5 cm long; leaf blade ovate to oblong-ovate, 6.4–7.7 cm long, 4.4–5.3 cm broad, hirsutulous; base truncate to subcordate; margin mucronate crenate; apex acuminate. Cymes axillary and terminal, dichotomous, lax; peduncle 2–4.5 cm long. Bracts linear, minute, 1.6–2.1 mm long. Calyx 4.5–5 mm long at anthesis, puberulent, dilated after anthesis, 5-toothed; calyx teeth triangular-lanceolate, subequal, 1.4–1.7 mm long, one tooth slightly elongated. Corolla yellow, 1.8–2.4 cm long, sparsely pubescent outside; corolla tube conspicuous, 0.9–1.2 cm long; upper corolla lips laterally compressed, apex entire; lower corolla lips 3-lobed, middle lobe obovate, wider than lateral lobes. Nutlets brown, obovoid, ca. 1.8 mm long. Figure IV-35. Phenology: Flowering from August to October and fruiting in November. Distribution: This species is endemic to Nepal (Map IV-18).

Map IV-18. Microtoena nepalensis Stearn.

Habitat: It grows in forests, forest and thicket margins, open grassy slopes, at altitudes from 2110 m in Dhankuta district to 2200 m in Chittre of Sinduwa of Nepal. Additional specimens examined: NEPAL, SINDUWA, Dhankuta district, 2110m, 1963. 10. Hara, H., Kanai, H., Kurosawa, S. & Murata, G. 6306496 (BM, K); Chittre, 27.06N; 87.25E, 2200m, 1972. 8. 16. Dobremez, J. F. 1482 (BM). 18. Microtoena robusta Hemsl., Journ. Linn. Soc. Bot. 26: 307.1890. Type: China, Hubei, Hsingshan [Xingshan], Mar 1889, A. Henry 6482A [lectotype designated here: K! (K 000928185); isolectotype: BM! (BM 000950460), K! (K 000928184), P! (P 00743580)].

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Figure IV-35. Microtoena nepalensis Stearn: a. flowering stem; b. flower with conspicuous tube and minute bract; c. dissected calyx.

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A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

Herbs perennial. Stems erect, 1–2 m tall, densely puberulent, sometimes appressed, base sometimes woody. Leaf petiole 2–10 cm long; leaf blade ovate, 5.4–12.2 cm long, 3.9–8.9 cm broad, densely hispid; base truncate to subcordate; margin mucronate crenate; apex shortly acuminate. Cymes axillary and terminal, dichotomous, lax; peduncle 1–3 cm long. Bracts linear, tiny, 2.9–3.5 mm long, less than 0.5 mm broad. Calyx 2.6–3.5 mm long at anthesis, puberulent, dilated after anthesis, 5-toothed; calyx teeth triangular, short, subequal, 0.6–0.9 mm long, not hooked, connivent after anthesis. Corolla white, marked with red on upper lip, 1.5–2.2 cm long, hirsutulous outside; corolla tube conspicuous, 0.8–1.2 cm long; upper corolla lips laterally compressed, apex entire; lower corolla lips 3-lobed, middle lobe subcircular, wider than lateral lobes. Nutlets brown to black, obovoid, smooth. Figure IV-36 & 37. Phenology: Flowering from May to August and fruiting in September. Distribution: This species is distributed in Fangxian, Xingshan, and Shennongjia of Hubei and Wenxian of Gansu province (Map IV-19).

Map IV-19. Microtoena robusta Hemsl.

Habitat: It grows at forest and thicket margins, at altitude from 820m in Wenxian, Gansu Province to 1700m in Wuxi, Chongqing City of China. Additional specimens examined: CHINA, CHONGQING, Chengkou, s. loc. 1500 m, 24 Jul 1893, R. P. Farges 1192 (P); loc. eodem, s. d. R. P. Farges s. n. (P). Wushan, Dengjia Tea Farm, in woods, 1330-1570 m, 09 Sep 2009, The Yangtse three Gorges Exped. 3595 (PE). Wuxi, Shuangyang, Baiguo, in woods, 1408 m, 12 Jul 2004, Y. S. Chen 1406 (WUK); loc. eodem, in woods, 1700 m, 13 Jul 2004, Y. S. Chen 1593 (WUK). GANSU, Wenxian, Fanba, Wenjia Ditch, grassy slopes, 820 m, 01 Sep 1976, J. Q. Fu 2756 (PE). HUBEI, Fangxian, s. d., A. Henry 7631 (K). SICHUAN, Pingwu, Laohe Ditch, Hualian Slope, forest edge, 1270 m, 08 Aug 2005, D. H. Zhu et al. 95 (HX).

CHAPTER IV. TAXONOMIC REVISION OF MICROTOENA

Figure IV-36. Microtoena robusta Hemsl.: a. flowering stem; b. flower; c. calyx

115

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A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

Figure IV-37. Microtoena robusta Hemsl.: A. a field individual photographed by Dr. C. L. Xiang; B. white flower marked with red on upper lip; C. leaf, scale bar = 2 cm; D. a fruiting cyme and a fruiting calyx with connivent teeth, scale bar = 5 mm.

19. Microtoena vanchingshanensis C.Y. Wu & S.J. Hsuan, Acta Phytotax. Sin. 10: 52. 1965. Type: China, Guizhou, Yinjiang, Mt. Fanjing, stream side, 1650 m, 12 Aug 1932, T.H. Tu 3486 [holotype: PE! (PE 00031128); isotype: IBSC! (IBSC 0583626)]. Herbs perennial. Stems erect, 0.5–0.8 m tall, puberulent, base sometimes prostrate. Leaf petiole 2–5.2 cm long; leaf blade ovate, 5.4–6.1 cm long, 3.5–3.9 cm broad, densely hispid; base cordate; margin mucronate crenate; apex shortly acuminate. Cymes axillary and terminal, dichotomous, slightly compact; peduncle 0.5–2 cm long. Bracts linear, tiny, 4.9–5.3 mm long, 2.91–2.4 mm broad. Calyx 4.1–4.5 mm long at anthesis, pilose outside, dilated after anthesis, 5-toothed; calyx teeth triangular, short, subequal, 0.9–1 mm long, not hooked. Corolla uniformly white, 1.6–1.8 cm long, hirsutulous outside; corolla tube conspicuous, 0.9–1 cm long; upper corolla lips laterally compressed, apex entire; lower corolla lips 3-lobed, middle lobe obovate, wider than lateral lobes. Nutlets unknown. Figure IV-38.

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Figure IV-38. Microtoena vanchingshanensis C. Y. Wu & S. J. Hsuan: a. flowering stem with a prostrate base; b. flower with conspicuous tube; c. fruiting calyx; d. dissected calyx.

118

A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

Phenology: Flowering from August to September and fruiting in October. Distribution: This species is endemic to Fanjing mountain of Guizhou province (Map IV-20).

Map IV-20. Microtoena vanchingshanensis C. Y. Wu & S. J. Hsuan.

Habitat: It grows in forest, and at forest and thicket margins, at an altitude of 1650m in Mt. Fanjing of Yinjiang, Guizhou Province of China. Additional specimens examined: CHINA, CHONGQING, Nanchuan, s. loc., s. d., M. Tseng 795 (NAS).

EXCLUDED NAME Microtoena coreana H. Lév., Repert. Spec. Nov. Regni Veg. 9: 223. 1911. Type: Korea, Seoul, Mt. Nam-San, 23. Sept. 1901, U. Faurie 752 (holotype: E! (E00284127)). Note: Wu (1959) & Hsuan (1965) considered that M. coreana did not belong to Microtoena based on its distribution pattern. I have critically examined the holotype of M. coreana collected by Faurie from Seoul of South Korea. It is clearly conspecific with Tripora divaricata (Maxim) Cantino (=Caryopteris divaricata Maxim). Therefore, I exclude this name from Microtoena (Wang, 2015).

BIBLIOGRAPHY Baranova M. 1987. Historical development of the present classification of morphological types of stomata. Botanical Review, 53: 53-79. Baranova M. 1992. Principles of comparative stomatographic studies of flowering plants. Botanical Review, 58: 1-99. Bentham G. 1876. Labiatae. Genera Plantarum, vol. 2. London: Reeve. 1160-1223. Briquet J. 1895. Labiatae. Die Naturlichen Pflanzenfamilien. div 4.3a. Leipzig: Wilhelm Engelmann. 183-380. Clarke C B. 1889. On the plants of Kohima and Muneypore. Journal of the Linnaean Society, 25: 58. Coyne J A, Orr H A. 2004. Speciation. Sunderland: Sinauer Associates. Diels L. 1900. Flora von Central-China. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie, 29(3-4): 556-557. Dunn M S. 1913. A Key to the Labiatae of China. Notes from the Royal Botanic Garden, Edinburgh, 8: 169. Erdtman G. 1945. Pollen morphology and plant taxonomy. IV. Labiatae, Verbenaceae, and Avicenniaceae. Svensk Botanisk Tidskrift, 39: 279-285. Hance H F. 1884. A new Chinese Gomphostemma. Journal of Botany, British and Foreign, 22(8), 231-232. Handel-Mazzetti H F V. 1936. Microtoena Prain. Symbolae Sinicae, 7(4): 927. Harley R M, Paton A J, Ryding O. 2003. New synonymy and taxonomic changes in the Labiatae. Kew Bulletin, 58: 485-489. Harris J G, Harris W H. 1994. Plant Identification Terminology. Payson: Spring Lake Publishing, 121-188. Hemsley W B. 1890. Labiatae. The Journal of the Linnean Society, Botany, 26: 307. Hong D Y. 1983. On pollen shape in some groups of dicotyledons. Grana, 22: 73-78. Hong D Y. 2010. Peonies of the World. Richmond: Royal Botanic Gardens, Kew. 8-10. Hooker J D. 1885. The Flora of British India. vol. 4. London: L. Reeve & Co. 624. Hsuan S J. 1965. Revisio Generis Microtoena Labiatarum Sinensium. Acta Phytotaxonomica Sinica, 10(1): 41-56. Kadereit J W. 2004. Flowering Plants, Dicotyledons, Lamiales (except Acanthaceae including Avicenniaceae). In: Kubitzki K (Ed.) The Families and Genera of Vascular Plants Vol. VII. Berlin: Springer-Verlag. Léveillé H. 1911. Microtoena. Repertorium Specierum Novarum Regni Vegetabilis, 9(208-210): 222-223. Li C Q, Li B G, Qi H Y, et al. 2004. Three cyclooctapeptides and one glycoside from Microtoena prainiana. Journal of natural products, 67: 978-982. Li C Q, Li B G, Wang F P, et al. 2006. A Diterpene Endoperoxide from Microtoena insuavis (Hance) Prain ex Dunn. Journal of Integrative Plant Biology, 48 (5): 613-616. Li X W, Hedge I C. 1994. Microtoena Prain. In: Wu Z Y, Raven P H & Hong D Y (Eds.) Flora of China, vol. 17. Bejing/St. Louis: Science Press & Missouri Botanical Garden Press, 189-194.

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Lohith K, Vijayakumar G R, Somashekar B R, et al. 2006. Glycosides and amino acyl esters of carbohydrates as potent inhibitors of angiotensin converting enzyme. European Journal of Medicinal Chemistry, 41(9): 1059-1072. Lowe R T. 1868. A Manual Flora of Madeira and the Adjacent Islands Porto Santo and the Desertas. vol.1. London: van Voorst. 490. Manami S, Kazuya N, Michiho I, et al. 2007. Chemical composition of the essential oil of Microtoena patchoulii [(C.B. Clarke Ex J.D. Hooker) C.Y. Wu et Hsuan]. Journal of essential oil research, 19(4): 336-337. McNeill J. et al. 2012. International code of botanical nomenclature (Melbourne Code). Adopted by the 18th Int. Bot. Congr. Melbourne, Australia, July 2011. – Reg. Veg. 154, ARG Gantner Verlag KG. Pollen group of morphology department, Institute of Botany, Chinese academy of Sciences. 1960. Pollen morphology of Chinese plant. Beijing: Science Press. Prain D. 1889. Microtoena cymosa. Hooker's Icones Plantarum, 19: 1872. Prain D. 1890. Some additional species of Labiatae. Journal of the Asiatic Society of Bengal, 59(2): 310. Prain D. 1895. Microtoena Prain. Bulletin de la Société Botanique de France, 42: 421-427. Prain D. 1908. Patchouli. (Pogostemon Patchouli, var. suavis, Hk. f. = P. Cablin, Benth.). Bulletin of Miscellaneous Information (Royal Gardens, Kew), 1908: 78-82. Scheen A C, Bendiksby M, Ryding O, et al. 2010. Molecular phylogenetics, character evolution, and suprageneric classification of Lamioideae (Lamiaceae). Annals of the Missouri Botanical Garden, 97(2): 191-217. Stearn W T. 1982. Microtoena Prain. An Enumeration of the Flowering Plants of Nepal, 3: 158. Stearn W T. 1983. New species of Microtoena (Labiatae) from Nepal, Bhutan, Assam, and Thailand. The Journal of Japanese Botany, 58(1): 1-13. Stebbins G L. 1950. Variation and Evolution in Plants. New York: Columbia University Press. Wang F X, Qian N F, Zhang Y L, et al. 1995. Pollen morphology of China. Beijing: Science Press. Wang Q, Hong D Y. 2011. Character analysis and taxonomic revision of the Microtoena insuavis Complex (Lamiaceae). Botanical Journal of the Linnean Society, 165: 315-327. Wang Q, Hong D Y. 2012. Identity of Microtoena affinis (Lamiaceae). Systematic Botany, 37(4): 1031-1034. Wang Q, Hong D Y. 2015. Identity of Microtoena coreana (Lamiaceae), a doubtful species from Korea. Phytotaxa, 195(1): 079-085. Wu Z Y. 1959. Revisio labiatarum Sinensium. Acta Phytotaxonomica Sinica, 8(1): 44-49.

INDEX TO SPECIMENS CITED* B Balansa, B. 3582 (K, P), 3583(K, P), 3584 (K, P) [M. insuavis] Beijing Exped. 3832 (PE), 4241 (PE), 4384 (PE) [M. urticifolia]; 890847 (PE) [M. mollis] Beusekom, C. F. van et al. 4760 (P) [M. insuavis] Bijie Exped. 670 (PE), 903 (KUN, PE), 993 (PE) [M. prainiana] Bois, D. 170 (P) [M. insuavis] Boufford, D. E. & Bartholomew, B. 24821 (A, KUN) [M. moupinensis] Boufford, D. E. et al. 27189 (A, CDBI) [M. moupinensis] Bunchuai, K. 37088 (P) [M. insuavis]

C Cavalerie, J. 3548 [syntypes of M. mollis: E (E 00284284), K (K 000928183)] [M. mollis] Chang, C. C. 10679 [holotype of M. subspicata: IBSC (IBSC 0005116)] [M. esquirolii] Chang, S. C. & Ren, Y. X. 07624 (PE) [M. moupinensis] Chang, Y. P. 0811 (IBSC, KUN, PE) [M. delavayi], 817 [holotype of M. affinis: KUN (KUN 0484539)] [M. delavayi] Chen, J. 451 (KUN) [M. patchoulii] Chen, Y. S. 1406 (WUK), 1593 (WUK) [M. robusta] Chen, S. Y. et al. 4226 [paratypes of M. omeiensis: NAS (NAS 00072410), SM (SM 0161053, SM 0161054, SM 0161055, SM 0161056), SZ (SZ 00241922)] [M. omeiensis] Cheng, W. C. 6509 (NAS, PE) [M. omeiensis], 6777 [paratype of M. omeiensis: PE (PE 00824297)] [M. omeiensis] Chevalier, A. 29668 (P), 29750 (P) [M. insuavis]

China-Soviet Union Exped. 5710 (IBSC, KUN) [M. mollis] China-Vietnam Exped. 2316 (KUN) [M. insuavis] Chinese Botanic Union-Guangxi Exped. 4041 (IBK), 4556 (IBK) [M. esquirolii] Ching, R. C. 23689 [paratypes of M. delavayi var. amblyodon: KUN (KUN 0217782, KUN 0217783), PE (PE 00824241, PE 00824242)] [M. wardii]; 50636 (KUN, PE), 50702 (KUN) [M. patchoulii] Chow, T. Y. & Hsu, K. J. 343 [paratype of M. omeiensis: NAS (NAS 00223586)] [M. omeiensis] Chu, K. L. 3558 (IBSC, PE), 3633 (BM, IBSC, PE, SZ), 3951 (IBSC, PE, SZ), 4601 (NAS, PE) [M. moupinensis] Clarke, C. B. 15983 [lectotype of Plectranthus patchouli: K (K 000928190)] [M. patchoulii] Colani, E. 5003 (P), L.968 (P), s. n. (P) [M. insuavis]

D Dang, C. Z. 927 (HGAS) [M. esquirolii] Delavay, M. J. s. n. (P) [M. delavayi]; 2190 [lectotype of M. delavayi var. grandiflora: P (P 00743575); isolectotypes: P (P 00743573, P 00743574)] [M. delavayi]; 2463 [holotype of M. delavayi: P (P 00743569); isotypes: P (P 00743570, P 00743571, P 00743572, P 04444595)] [M. delavayi]; 4206 [syntypes of M. delavayi var. grandiflora: P (P 00743576, P 00743577, P 00743578)] [M. delavayi] David, M. I. s. n. [lectotype of Clerodendrum moupinense: P (P 00743579)] [M. moupinensis] Dobremez, J. F. 1482 (BM) [M. nepalensis]

* The accepted names are in bold and presented in square parentheses at the end of each gathering.

122

A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

E Erdhügel 788 [holotype of M. prainiana: K (K 11146)] [M. prainiana] Esquirol, J. 330 [syntype of M. mollis: E (E 00284283)] [M. mollis]; 672 [holotype of M. esquirolii: E (E 00284286)] [M. esquirolii]; 755 [lectotype of M. mollis: E (E 00284285)] [M. mollis]

F Fang, W. P. & Hu, W. K. 51-1117 (SZ, WUK) [M. moupinensis] Fang, W. P. 2640 (GH, NAS, K, P); 3132 [paratype of M. omeiensis: PE (PE 00824302)] [M. omeiensis] Farges, R. P. 1192 (P), s. n. (P) [M. robusta] Faurie, U. 752 [holotype of M. coreana (excluded name): E (E 00284127)] [Tripora divaricata] Feng, K. M. 3396 (KUN, PE), 8778 (KUN, PE) [M. delavayi]; 11344 [holotype of M. megacalyx: PE (PE 00031247); isotypes: A (A 00001327), KUN (KUN 0484541, KUN 0484542)] [M. megacalyx] Feng, Z. B. et al. 20060455 (HX) [M. Moupinensis] Forrest, G. 20586 (K, P) [M. delavayi]; 22408 (K, P) [M. muliensis]; 25192 (P), 26017 (K, P), 26082 (BM, IBSC, K, PE), 26217 (P) [M. patchoulii]; s. n. (K) [M. insuavis] Fu, J. Q. 2756 (PE) [M. robusta]

G Gaoligong Shan Biodiversity Survey 2003 18871 (GH) [M. stenocalyx] Griffith 4059 [lectotype of M. griffithii: K (K 000928194)] [M. moupinensis] Guan, K. J. & Wang, W. T. 2525 (PE), 3219 (PE) [M. moupinensis] Guan, Z. T. 9118 (PE) [M. moupinensis] Guizhou Exped. 198 (PE) [M. mollis]

H Handel-Mazzetti, H. F. V. 8412 [holotype of

M. urticifolia var. subedentata: WU (WU 0060512)] [M. wardii] Hara, H., Kanai, H., Kurosawa, S. & Murata, G. 6306496 (BM, K) [M. nepalensis] Henry, A. 2536 [paratype of M. urticifolia: K (K 000928189)] [M. urticifolia]; 6482A [lectotype of M. robusta: K (K 000928185); isolectotypes: BM (BM 000950460), K (K 000928184), P (P 00743580)] [M. robusta]; 7339 [holotype of M. urticifolia: K (K 000928187); isotype: K (K 000928188)] [M. urticifolia]; 7631 [syntype of M. robusta: K (K 000928186)] [M. robusta]; 11583 (K) [M. mollis]; 12588 (A, K) [M. insuavis]. Henry, B. C. 22237 [holotype of Gomphostemma insuave: BM (BM 000950459)] [M. insuavis] Ho, Y. Y. 6014 [paratypes of M. omeiensis: NAS (NAS 00223584, NAS 00223585)] [M. omeiensis] Hsieh, C. C. 42079 (IBSC, PE, SZ) [M. prainiana]; 42137 (PE, SZ), 42606 (IBSC, PE) [M. moupinensis] Hsiung, C. H. et al. 32020 [holotype of M. omeiensis: SZ (SZ 00241919); isotypes: IBSC (IBSC 0005114), PE (PE 00824299)] [M. omeiensis] Hsiung, G. H. & Chow, Z. L. 91984 (PE, SZ), 92513 (PE, SZ), 92690 (IBSC, PE, SZ), 93340 (IBSC, KUN, PE, SZ), 95257 (SM) [M. prainiana] Hsiung, R. L. & Wen, S. K. 580890 (KUN), 580981 [holotype of M. pauciflora: KUN (KUN 0484547)] [M. patchoulii] Hu 0275 (SM) [M. moupinensis] Hubei Shennongjia Exped. 2494 (PE), 11130 (PE) [M. moupinensis] Huidong Exped. 0504 (SM), 0685 (SM) [M. prainiana]

J Jenkins s. n. [syntype of P. patchouli: K] Jian, Z. B. et al. 31123 (PE), 31157 (PE) [M. urticifolia] Jin, X. H. et al. ST0278 (PE), ST0464 (PE) [M. stenocalyx]

INDEX TO SPECIMENS CITED

K Kaarders 23161b (P) [M. insuavis] Kang, T. S. 50809 (NAS) [M. prainiana] Kerr, A. F. G. 6538 [holotype of M. siamica: BM (BM 000950464); isotype: BM (BM 000950463)] [M. insuavis] Kingdon-Ward, F. 14098 [paratype of M. wardii: BM], 14235 [holotype of M. wardii: BM (BM 000950462)] [M. wardii]; 21695 (BM, K) [M. patchoulii]

L Lau, S. K. 3100 (P), 5474 (IBSC) [M. insuavis] Lee, T. T. 2985 [holotype of M. urticifolia var. brevipedunculata: PE (PE 00824264); isotype: IBSC (IBSC 0005117)] [M. brevipedunculata] Lemarie, M. s. n. (P) [M. insuavis] Li, G. F. 63622 (IBSC, KUN, SZ), 63726 (PE, SZ), 63841 (KUN, PE, SZ) [M. prainiana] Li, L. Q. 133 (PE) [M. urticifolia] Li, M. K. 1363 (IBSC, KUN, QTPMB) [M. patchoulii]; 246 (KUN), 2690 (IBSC, KUN, WUK), 2990 (KUN, WUK) [M. delavayi] Li, Y. H. 02841 (KUN), 1732 (HITBC, KUN), 309 (HITBC, IBK, IBSC, KUN), 3821 (HITBC, IBK, IBSC, KUN), 4768 (HITBC, IBSC, KUN), 5084 (HITBC, IBSC, KUN), 5175 (HITBC, IBK, IBSC, KUN) [M. mollis]; 3857 (KUN) [M. insuavis] Li, Z. J. 3003 (IBSC), 3259 (IBSC) [M. insuavis] Li, Z. Y. et al. 1569 (PE) [M. esquirolii] Liang, X. R. 66028 (IBK, IBSC) [M. insuavis] Liang, Z. F. & Wu, D. L. 32050 (IBK, IBSC) [M. mollis] Lin, L. Y. 1148 [paratype of M. omeiensis: IBSC (IBSC 0583632)] [M. omeiensis] Liu, E. D. & Yang, D. C. 260 (KUN) [M. delavayi] Liu, E. D. 1464 (KUN) [M. mollis] Liu, X. Q. 24836 (IBK, KUN, PE, SZ, WUK), 25348 (IBK, IBSC) [M. insuavis] Loc, P. K. et al. P10531 (P) [M. insuavis] Longtan Exped. 00590 (IBK), 01859 (IBK) [M. esquirolii]

123

Ludlow F. et al. 19640 [holotype of M. bhutanica: BM (BM 000950461)] [M. delavayi] Lushan Exped. 78-0834 (SM) [M. moupinensis]

M Maire, E. E. s. n. [holotype of M. maireana: W (W 5977)] [M. moupinensis] Mann, S. s. n. (K) [M. patchoulii] Meigu-Puge Exped. 0310 (SM) [M. omeiensis] Mianning Exped. 627 (SM) [M. delavayi] Mt. Emei Exped. 1827(SM) [M. omeiensis]

N NE Yunnan Exped. 39 (KUN) [M. moupinensis] Ningnan Exped. 757 (SM) [M. moupinensis] NW Yunnan-Jinsha River Exped. 6596 (KUN) [M. delavayi]

P Pei, S. J. 59-11300 (KUN, WUK) [M. insuavis] Pei, Y. 8121 (NAS, PE) [M. moupinensis] Peng, H. 1904 (KUN) [M. patchoulii] Peng, H. et al. 113 (KUN) [M. omeiensis] Puge Exped. 854 (SM), 879 (SM) [M. moupinensis]

Q Qi, Y. F. 612615 (KUN) [M. delavayi] Qinghai-Tibet Exped. 13317 (KUN) [M. delavayi]; 8280 (HITBC, KUN) [M. Stenocalyx] Qiu, B. Y. 55017 (IBSC, KUN, PE), 55154 (PE) [M. delavayi]

S S Guizhou Exped. 3636 (KUN, PE) [M. prainiana] s. coll. 0713, 0911, 0941, 742 (SM) [M. moupinensis]; 0829, 248, 504, s. n. (SM) [M. prainiana]; 0896, 388, 74-19 (SM), 22363, L5862, s. n. (KUN), s. n. (HITBC) [M. delavayi]; 1117 (SM), 1197 (KUN) [M. urticifolia]; 1301, 878 (SM) [M. omeie-

124

A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

nsis]; 396 (IBSC) [M. esquirolii]; 59-13833 (IBSC), 5393, L936, S034, s. n. (P) [M. insuavis]; s. n. (SM) [M. muliensis] Shimian Exped. 78-1016, 78-1151, 78-1252 (SM) [M. moupinensis] Shui, S. K. 4931 [holotype of M. delavayi var. lutea: KUN (KUN 0217786); isotype: KUN (KUN 0217787)] [M. wardii] Sichuan Econ. Plant Exped. 5842 [holotype of M. longisepala: KUN (KUN 0484540); isotypes: PE (PE 00031248, PE 00824276), SM (SM 0161009)] [M. moupinensis] Sichuan Medic. Plant Resources Surv. 11287 [holotype of M. miyiensis: SM (SM 64)] [M. miyiensis] Sin, S. S. 50809 (IBSC) [M. albescens] Soejarto, D. D. et al. 10427 (P) [M. insuavis] South China sub-Exped. 2494, 2530 (IBSC) [M. esquirolii]

T Taiping Exped. 0486 (SM) [M. muliensis] Tan, P. X. 57102 (IBSC) [M. insuavis]; 57324 (IBSC) [M. esquirolii] Tao, Y. H. 51446 [paratypes of M. omeiensis: SZ (SZ 00241915, SZ 00241916, SZ 00241923)] [M. omeiensis] Teng, S. W. 1330 [holotype of M. albescens: IBSC (IBSC 0005113)] [M. albescens] The Yangtse three Gorges Exped. 3595 (PE) [M. robusta] Tsai, H. T. 51139 (A, KUN, NAS, PE, SZ, WUK) [M. prainiana]; 54088 [holotype of M. delavayi var. amblyodon: PE (PE 00031250), isotypes: A (A 00001326), IBSC (IBSC 0583620), KUN (KUN 0217778), NAS (NAS 00223563), PE (PE 00824263), SZ (SZ 00241927)] [M. wardii]; 56519 [paratypes of M. stenocalyx: A (A 00145875), IBSC (IBSC 0583659), KUN (KUN 0217857), PE (PE 00824281, PE 00824296), SZ (SZ 00241925)] [M. stenocalyx]; 57468 [paratypes of M. delavayi var. amblyodon: KUN ( KUN 0217779, KUN 0217780, KUN 0217781)] [M. wardii]; 58-8185 (KUN) [M. mega-

calyx]; 59995 (A, IBSC, KUN, NAS, PE, SZ), 63007 (KUN, NAS, PE, SZ) [M. delavayi] Tsao, T. Y. 1147 [paratypes of M. subspicata: PE (PE 00824328, PE 00824329)] [M. esquirolii] Tseng, M. 48, 856 (PE) [M. prainiana]; 795 (NAS) [M. vanchingshanensis] Tsiang, H. L. 11053 (IBSC, SZ) [M. moupinensis] Tsiang, Y. 11884 [holotype of M. tenuiflora: PE (PE 00031251); isotypes: IBSC (IBSC 0583597, IBSC 0584279), KUN (KUN 0217754), NAS (NAS 00072411, NAS 00223564, NAS 00223566, NAS 00223568, NAS 00223574), PE (PE 00824243), WUK (WUK 0120785)] [M. delavayi]; 12597 (IBSC, KUN, NAS, PE) [M. patchoulii]; 1693 (BM, NAS) [M. insuavis]; 7334 (IBSC, NAS) [M. esquirolii] Tu, T. H. 3486 [holotype of M. vanchingshanensis: PE (PE 00031128); isotype: IBSC (IBSC 0583626)] [M. vanchingshanensis]; 4607 (IBSC) [M. moupinensis]

V Vidal, J. E. 777B (P) [M. patchoulii]

W Wang, C. W. 80153 (IBSC, KUN, NAS, PE, WUK), 80582 (IBSC, KUN, PE) [M. insuavis]; 86480 (IBSC, KUN, PE) [M. mollis] Wang, F. T. 22040 (KUN, NAS) [M. omeiensis] Wang, H. et al. 03-0951 (KUN) [M. Moupinensis] Wang, W. T. 10429 (KUN) [M. patchoulii]; 247 (KUN) [M. delavayi] Wang, Y. M. 5638, 5952 (PE) [M. urticifolia] Wen, S. K. 580696 (KUN) [M. delavayi] Williams & Stainton 8384 [holotype of M. nepalensis: BM (BM 000521969)] [M. nepalensis] Wilson, E. H. 4372 (A, BM, IBSC) [M. urticifolia] Wu, S. K. 368 [holotype of M. subspicata var.

INDEX TO SPECIMENS CITED intermedia: KUN (KUN 0484545); isotype: KUN (KUN 0484546)] [M. esquirolii]; 1993 (PE) [M. moupinensis]; 2589 [holotype of M. muliensis: PE; isotypes: CDBI (CDBI 0172354), KUN (KUN 0484543, KUN 0484544)] [M. muliensis]; 2591 (KUN), 3703 (CDBI, KUN, PE) [M. delavayi]; 8496 (HITBC, KUN) [M. muliensis] Wu, S. K. et al. 1124 (KUN) [M. insuavis] Wuling Exped. 1610 (IBSC) [M. brevipedunculata]

X Xiang, C. L. 340 (KUN) [M. prainiana] Xiong, X. Y. & Shao, H. M. T159 (HX) [M. moupinensis] Xu, G. R. 23295 (CDBI, IBSC) [M. moupinensis]

Y Yang, G. S. 1426 (KUN) [M. patchoulii]; 610123 [paratype of M. delavayi var. amblyodon: KUN (KUN 0217788)] [M. wardii] Yang, K. H. 56702 [paratypes of M. omeiensis: IBSC (IBSC 0583648), KUN (KUN 0217820), PE (PE 00824298), SZ (SZ 00241920, SZ 00241921)] [M. omeiensis] Yao, C. W. 4842 [paratype of M. omeiensis: PE (PE 00824300)] [M. omeiensis]

125

Yao, G. 377, 378 (IBSC) [M. mollis] Yin, W. Q. 60-1010 (KUN) [M. delavayi]; 60-1521 (KUN) [M. patchoulii] Ying, J. S. 10091 (PE) [M. moupinensis] Yingjing Exped. 78-0837 (SM) [M. omeiensis] Yu, P. H. 1261 (KUN) [M. megacalyx] Yu, S. W. s. n. (KUN) [M. delavayi] Yü, T. T. 7517 (A, KUN, PE) [M. delavayi]; 16966 [holotype of M. stenocalyx: KUN (KUN 0217858); isotypes: A (A 00001328), KUN (KUN 0217859), PE (PE 00824295, PE 00824327)] [M. stenocalyx] Yunnan University-Biology department 00812 (IBSC, KUN) [M. delavayi]

Z Zhang, A. L. & Yu, S. W. 100913 (KUN) [M. delavayi] Zhao, C. M. EX0551 (PE) [M. megacalyx] Zhao, Z. C. 4736, 4777 (PE) [M. moupinensis]; 88-2009 (PE) [M. urticifolia] Zhong, M. F. & Tang, S. G. 1033 (SM) [M. omeiensis] Zhou, S. S. 2174 (PE) [M. insuavis] Zhou, X. 166 (KUN) [M. insuavis] Zhu, D. H. et al. 4763 (HX) [M. moupinensis]; 95 (HX) [M. robusta] Zhu, T. P. 0467 (KUN) [M. insuavis] Zi, S. S. 155 (KUN) [M. stenocalyx]

INDEX TO BOTANICAL NAMES AND SYNONYMS Clerodendrum moupinense Franch. (1887) =Microtoena moupinensis (Franch.) Prain Gomphostemma insuave Hance (1884) =Microtoena insuavis (Hance) Prain ex Briq. Microtoena Prain (1889) affinis C.Y. Wu & S.J. Hsuan (1965) =Microtoena delavayi Prain albescens C.Y. Wu & S.J. Hsuan (1965) bhutanica Stearn (1983) =Microtoena delavayi Prain brevipedunculata (C.Y. Wu & S.J. Hsuan) Q. Wang (2017) = var. brevipedunculata C.Y. Wu & S.J. Hsuan (1965) coreana H. Lév. (1911) =Tripora divaricata (Maxim.) P.D. Cantino cymosa Prain (1889) =Microtoena patchoulii (C.B. Clarke ex Hook. f.) C.Y. Wu & S.J. Hsuan delavayi Prain (1895) var. amblyodon C.Y. Wu & S.J. Hsuan (1965) =Microtoena wardii Stearn var. grandiflora Prain (1895) =Microtoena delavayi Prain var. lutea C.Y. Wu & S.J. Hsuan =Microtoena wardii Stearn var. vera Prain (1895) =Microtoena delavayi Prain esquirolii H. Lév. (1911) griffithii Prain (1890) =Microtoena moupinensis (Franch.) Prain insuavis (Hance) Prain ex Briq. (1895) = Gomphostemma insuave Hance (1884) insuavis (Hance) Prain ex Dunn. (1913) = Gomphostemma insuave Hance =Microtoena insuavis (Hance) Prain ex Briq. longisepala C.Y. Wu ex S.J. Hsuan (1965)

INDEX TO BOTANICAL NAMES AND SYNONYMS =Microtoena moupinensis (Franch.) Prain maireana Hand.-Mazz. (1936) =Microtoena moupinensis (Franch.) Prain megacalyx C.Y. Wu (1959) miyiensis C.Y. Wu & H.W. Li (1977) mollis H. Lév. (1911) moupinensis (Franch.) Prain (1895) =Clerodendrum moupinense Franch. (1887) muliensis C.Y. Wu ex S.J. Hsuan (1965) nepalensis Stearn (1982) omeiensis C.Y. Wu & S.J. Hsuan (1965) patchoulii (C.B. Clarke ex Hook. f.) C.Y. Wu & S.J. Hsuan (1965) =Plectranthus patchouli C.B. Clarke ex Hook. f. (1885) pauciflora C.Y. Wu ex S.J. Hsuan (1965) =Microtoena patchoulii (C.B. Clarke ex Hook. f.) C.Y. Wu & S.J. Hsuan prainiana Diels (1900) robusta Hemsl. (1890) siamica Stearn (1983) =Microtoena insuavis (Hance) Prain ex Briq. stenocalyx C.Y. Wu & S.J. Hsuan (1965) subspicata C.Y. Wu ex S.J. Hsuan (1965) =M. esquirolii H. Lév. var. intermedia C.Y. Wu ex S.J. Hsuan (1965) =M. esquirolii H. Lév. tenuiflora C.Y. Wu (1959) =M. delavayi Prain urticifolia Hemsl. (1890) var. subedentata Hand.-Mazz. (1936) =Microtoena wardii Stearn var. brevipedunculata C.Y. Wu & S.J. Hsuan (1965) =M. brevipedunculata (C.Y. Wu & S.J. Hsuan) Q. Wang vanchingshanensis C.Y. Wu & S.J. Hsuan (1965) wardii Stearn (1983) Plectranthus patchouli C.B. Clarke ex Hook. f. (1885) =Microtoena patchoulii (C.B. Clarke ex Hook. f.) C.Y. Wu & S.J. Hsuan

127

DATA OF STANDARD DEVIATION ANALYSIS Data-1. The standard deviation analysis of leaf length of sampled populations. Population

Species

H&W08004

M. mollis

Variation range (cm) 4.26-8.26

Mean (cm) 6.255

Standard deviation (cm) 0.968

H&W08005

M. mollis

4.33-8.23

5.822

0.869

H&W08006

M. esquirolii

4.72-8.57

6.204

1.358

H&W08007

M. esquirolii

3.24-6.78

4.303

1.269

H&W08009

M. patchoulii

2.84-5.95

4.146

1.149

H&W09001

M. insuavis

3.43-4.89

4.051

0.535

H&W09002

M. insuavis

2.87-4.30

3.541

0.581

H&W09003

M. mollis

4.23-6.47

5.206

0.891

H&W09004

M. mollis

5.37-11.84

8.171

2.557

H&W09006

M. mollis

3.31-9.23

5.910

1.314

H&W09007

M. moupinensis

3.22-9.28

5.569

1.740

H&W09008

M. moupinensis

4.75-8.63

6.753

1.013

H&W09010

M. patchoulii

3.62-5.03

4.227

0.532

H&W09011

M. patchoulii

3.77-4.88

4.462

0.432

Data-2. The standard deviation analysis of leaf length of species. Species

Variation range (cm)

Mean (cm)

Standard deviation (cm)

1. M. esquirolii

3.24-8.85

5.695

1.965

2. M. insuavis

2.87-10.51

5.412

1.795

3. M. patchoulii

2.84-11.31

5.456

1.803

4. M. mollis

3.31-11.84

6.361

1.555

5. M. delavayi

4.63-17.16

8.708

2.781

6. M. wardii

7.06-12.49

9.588

1.822

7. M. urticifolia

6.98-7.74

7.398

0.304

8. M. albescens (holotype)

5.59

—

—

9. M. stenocalyx

4.71-7.77

6.510

0.802

10. M. miyiensis (holotype)

6.21

—

—

11. M. megacalyx

8.60-10.59

9.649

0.723

12. M. moupinensis

2.91-13.58

6.824

2.332

13. M. prainiana

5.00-11.58

8.280

1.955

14. M. muliensis

7.07-12.37

10.170

1.641

15. M. omeiensis

5.50-11.03

7.793

1.772

16. M. brevipedunculata

7.35-10.02

8.686

1.338

17. M. nepalensis

6.46-7.69

7.103

0.419

18. M. robusta

5.41-12.24

9.740

2.619

19. M. vanchingshanensis

5.49-6.09

5.774

0.248

DATA OF STANDARD DEVIATION ANALYSIS

129

Data-3. The standard deviation analysis of leaf shape of sampled populations. Population

Species

Variation range

Mean

Standard deviation

H&W08004

M. mollis

0.67-0.79

0.736

0.041

H&W08005

M. mollis

0.62-0.76

0.708

0.037

H&W08006

M. esquirolii

0.61-0.67

0.634

0.019

H&W08007

M. esquirolii

0.65-0.69

0.663

0.018

H&W08009

M. patchoulii

0.67-0.77

0.721

0.030

H&W09001

M. insuavis

0.55-0.70

0.643

0.062

H&W09002

M. insuavis

0.61-0.74

0.689

0.044

H&W09003

M. mollis

0.71-0.76

0.732

0.021

H&W09004

M. mollis

0.72-0.77

0.739

0.014

H&W09006

M. mollis

0.64-0.76

0.716

0.038

H&W09007

M. moupinensis

0.52-0.84

0.693

0.092

H&W09008

M. moupinensis

0.56-0.83

0.629

0.065

H&W09010

M. patchoulii

0.64-0.71

0.668

0.028

H&W09011

M. patchoulii

0.69-0.79

0.721

0.035

Data-4. The standard deviation analysis of leaf shape of species. Species

Variation range

Mean

Standard deviation

1. M. esquirolii

0.59-0.72

0.644

0.033

2. M. insuavis

0.45-0.74

0.640

0.061

3. M. patchoulii

0.58-0.79

0.704

0.046

4. M. mollis

0.62-0.82

0.729

0.041

5. M. delavayi

0.52-0.87

0.662

0.088

6. M. wardii

0.61-0.90

0.723

0.079

7. M. urticifolia

0.58-0.77

0.689

0.079

8. M. albescens (holotype)

0.78

—

—

9. M. stenocalyx

0.58-0.66

0.630

0.023

10. M. miyiensis (holotype)

0.68

—

—

11. M. megacalyx

0.54-0.67

0.606

0.046

12. M. moupinensis

0.52-0.89

0.677

0.084

13. M. prainiana

0.47-0.73

0.618

0.061

14. M. muliensis

0.35-0.57

0.476

0.067

15. M. omeiensis

0.55-0.71

0.627

0.040

16. M. brevipedunculata

0.71-0.82

0.763

0.058

17. M. nepalensis

0.65-0.73

0.683

0.027

18. M. robusta

0.66-0.77

0.729

0.042

19. M. vanchingshanensis

0.63-0.67

0.649

0.016

130

A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

Data-5. The standard deviation analysis of bract length of sampled populations. Population

Species

H&W08004

M. mollis

Variation range (cm) 0.46-0.60

Mean (cm) 0.504

Standard deviation (cm) 0.033

H&W08005

M. mollis

0.40-0.63

0.505

0.041

H&W08006

M. esquirolii

0.49-0.58

0.543

0.030

H&W08007

M. esquirolii

0.47-0.59

0.538

0.047

H&W08009

M. patchoulii

0.34-0.38

0.370

0.013

H&W09001

M. insuavis

0.27-0.28

0.275

0.003

H&W09002

M. insuavis

0.26-0.28

0.268

0.005

H&W09003

M. mollis

0.57-0.61

0.593

0.014

H&W09004

M. mollis

0.58-0.63

0.594

0.015

H&W09006

M. mollis

0.57-0.64

0.604

0.019

H&W09007

M. moupinensis

0.37-0.73

0.479

0.095

H&W09008

M. moupinensis

0.38-0.57

0.466

0.056

H&W09010

M. patchoulii

0.37-0.39

0.384

0.007

H&W09011

M. patchoulii

0.36-0.39

0.379

0.009

Data-6. The standard deviation analysis of bract length of species. Species

Variation range (cm)

Mean (cm)

Standard deviation (cm)

1. M. esquirolii

0.47-0.59

0.525

0.044

2. M. insuavis

0.21-0.51

0.273

0.051

3. M. patchoulii

0.23-0.46

0.367

0.032

4. M. mollis

0.33-0.64

0.508

0.082

5. M. delavayi

0.34-0.56

0.450

0.053

6. M. wardii

0.36-0.56

0.405

0.058

7. M. urticifolia

0.33-0.38

0.358

0.020

8. M. albescens (holotype)

0.53

—

—

9. M. stenocalyx

0.50-0.69

0.581

0.053

10. M. miyiensis (holotype)

0.42

—

—

11. M. megacalyx

0.35-0.50

0.432

0.064

12. M. moupinensis

0.32-0.73

0.446

0.071

13. M. prainiana

0.41-0.54

0.481

0.036

14. M. muliensis

0.28-0.39

0.315

0.029

15. M. omeiensis

0.29-0.41

0.347

0.034

16. M. brevipedunculata

0.41-0.42

0.418

0.007

17. M. nepalensis

0.16-0.21

0.183

0.017

18. M. robusta

0.29-0.35

0.325

0.023

19. M. vanchingshanensis

0.49-0.53

0.510

0.014

DATA OF STANDARD DEVIATION ANALYSIS

131

Data-7. The standard deviation analysis of bract shape of sampled populations. Population

Species

Variation range

Mean

Standard deviation

H&W08004

M. mollis

0.10-0.13

0.115

0.009

H&W08005

M. mollis

0.10-0.13

0.114

0.009

H&W08006

M. esquirolii

0.27-0.33

0.296

0.018

H&W08007

M. esquirolii

0.27-0.31

0.290

0.012

H&W08009

M. patchoulii

0.08-0.10

0.089

0.008

H&W09001

M. insuavis

0.12-0.13

0.123

0.005

H&W09002

M. insuavis

0.11-0.13

0.121

0.006

H&W09003

M. mollis

0.09-0.12

0.104

0.008

H&W09004

M. mollis

0.10-0.12

0.110

0.005

H&W09006

M. mollis

0.10-0.12

0.111

0.007

H&W09007

M. moupinensis

0.11-0.16

0.124

0.011

H&W09008

M. moupinensis

0.11-0.13

0.118

0.005

H&W09010

M. patchoulii

0.07-0.10

0.086

0.009

H&W09011

M. patchoulii

0.08-0.09

0.089

0.004

Data-8. The standard deviation analysis of bract shape of species. Species

Variation range

Mean

Standard deviation

1. M. esquirolii

0.27-0.44

0.323

0.049

2. M. insuavis

0.11-0.14

0.124

0.005

3. M. patchoulii

0.07-0.10

0.090

0.007

4. M. mollis

0.09-0.13

0.116

0.009

5. M. delavayi

0.09–0.14

0.114

0.010

6. M. wardii

0.09-0.12

0.108

0.006

7. M. urticifolia

0.10-0.13

0.114

0.009

8. M. albescens (holotype)

0.11

—

—

9. M. stenocalyx

0.29-0.39

0.348

0.026

10. M. miyiensis (holotype)

0.26

—

—

11. M. megacalyx

0.23-0.27

0.256

0.018

12. M. moupinensis

0.10-0.16

0.116

0.008

13. M. prainiana

0.37-0.59

0.447

0.052

14. M. muliensis

0.11-0.13

0.114

0.006

15. M. omeiensis

0.10-0.12

0.113

0.006

16. M. brevipedunculata

0.10-0.12

0.110

0.007

17. M. nepalensis

0.10-0.13

0.113

0.009

18. M. robusta

0.11-0.12

0.115

0.005

19. M. vanchingshanensis

0.44-0.46

0.448

0.011

132

A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

Data-9. The standard deviation analysis of calyx length of sampled populations. Population

Species

Variation range

Mean

Standard deviation

H&W08004

M. mollis

0.36-0.41

0.395

0.013

H&W08005

M. mollis

0.38-0.43

0.401

0.012

H&W08006

M. esquirolii

0.28-0.33

0.306

0.018

H&W08007

M. esquirolii

0.28-0.32

0.306

0.013

H&W08009

M. patchoulii

0.34-0.37

0.360

0.010

H&W09001

M. insuavis

0.28-0.30

0.290

0.005

H&W09002

M. insuavis

0.29-0.30

0.296

0.005

H&W09003

M. mollis

0.40-0.44

0.418

0.015

H&W09004

M. mollis

0.40-0.42

0.410

0.007

H&W09006

M. mollis

0.39-0.45

0.415

0.014

H&W09007

M. moupinensis

0.34-0.69

0.500

0.087

H&W09008

M. moupinensis

0.39-0.56

0.443

0.045

H&W09010

M. patchoulii

0.35-0.37

0.360

0.009

H&W09011

M. patchoulii

0.35-0.36

0.353

0.004

Data-10. The standard deviation analysis of calyx length of species. Species

Variation range

Mean

Standard deviation

1. M. esquirolii

0.28-0.33

0.306

0.014

2. M. insuavis

0.15-0.30

0.247

0.047

3. M. patchoulii

0.23-0.38

0.327

0.043

4. M. mollis

0.31-0.48

0.403

0.026

5. M. delavayi

0.38-0.61

0.457

0.058

6. M. wardii

0.33-0.50

0.434

0.046

7. M. urticifolia

0.28-0.33

0.303

0.017

8. M. albescens (holotype)

0.48

—

—

9. M. stenocalyx

0.48-0.58

0.507

0.032

10. M. miyiensis (holotype)

0.35

—

—

11. M. megacalyx

0.46-0.59

0.500

0.050

12. M. moupinensis

0.32-0.69

0.464

0.070

13. M. prainiana

0.37-0.47

0.420

0.025

14. M. muliensis

0.30-0.50

0.369

0.053

15. M. omeiensis

0.37-0.48

0.438

0.029

16. M. brevipedunculata

0.35-0.38

0.362

0.014

17. M. nepalensis

0.45-0.50

0.473

0.014

18. M. robusta

0.26-0.35

0.320

0.037

19. M. vanchingshanensis

0.41-0.45

0.429

0.013

DATA OF STANDARD DEVIATION ANALYSIS

133

Data-11. The standard deviation analysis of calyx split ratio of sampled populations. Population

Species

Variation range

Mean

H&W08004

M. mollis

0.39-0.48

0.423

Standard deviation

H&W08005

M. mollis

0.38-0.46

0.417

0.023

H&W08006

M. esquirolii

0.27-0.31

0.284

0.012

H&W08007

M. esquirolii

0.28-0.32

0.293

0.014

H&W08009

M. patchoulii

0.46-0.49

0.473

0.009

H&W09001

M. insuavis

0.47-0.52

0.486

0.013

H&W09002

M. insuavis

0.47-0.50

0.487

0.009

H&W09003

M. mollis

0.43-0.47

0.450

0.018

H&W09004

M. mollis

0.42-0.47

0.435

0.015

H&W09006

M. mollis

0.40-0.44

0.428

0.010

H&W09007

M. moupinensis

0.34-0.46

0.400

0.041

H&W09008

M. moupinensis

0.36-0.45

0.405

0.020

H&W09010

M. patchoulii

0.44-0.47

0.455

0.008

H&W09011

M. patchoulii

0.45-0.46

0.454

0.005

0.028

Data-12. The standard deviation analysis of calyx split ratio of species. Species

Variation range

Mean

Standard deviation

1. M. esquirolii

0.27-0.32

0.287

0.012

2. M. insuavis

0.39-0.52

0.474

0.026

3. M. patchoulii

0.40-0.49

0.448

0.017

4. M. mollis

0.38-0.48

0.424

0.020

5. M. delavayi

0.32-0.50

0.421

0.050

6. M. wardii

0.26-0.35

0.313

0.027

7. M. urticifolia

0.27-0.32

0.297

0.016

8. M. albescens (holotype)

0.44

—

—

9. M. stenocalyx

0.40-0.45

0.418

0.016

10. M. miyiensis (holotype)

0.37

—

—

11. M. megacalyx

0.27-0.33

0.300

0.025

12. M. moupinensis

0.34-0.49

0.417

0.036

13. M. prainiana

0.36-0.42

0.391

0.017

14. M. muliensis

0.31-0.46

0.392

0.041

15. M. omeiensis

0.26-0.36

0.319

0.027

16. M. brevipedunculata

0.36-0.38

0.370

0.008

17. M. nepalensis

0.31-0.35

0.327

0.013

18. M. robusta

0.24-0.30

0.264

0.021

19. M. vanchingshanensis

0.22-0.25

0.233

0.009

134

A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

Data-13. The standard deviation analysis of tooth ratio of sampled populations. Population

Species

Variation range

Mean

Standard deviation

H&W08004

M. mollis

1.02-1.11

1.057

0.027

H&W08005

M. mollis

1.01-1.13

1.066

0.035

H&W08006

M. esquirolii

1.07-1.20

1.109

0.039

H&W08007

M. esquirolii

1.04-1.27

1.125

0.083

H&W08009

M. patchoulii

1.04-1.17

1.101

0.038

H&W09001

M. insuavis

1.04-1.20

1.081

0.052

H&W09002

M. insuavis

1.02-1.30

1.131

0.099

H&W09003

M. mollis

1.03-1.15

1.087

0.043

H&W09004

M. mollis

1.02-1.20

1.105

0.060

H&W09006

M. mollis

1.02-1.20

1.096

0.050

H&W09007

M. moupinensis

1.11-1.41

1.264

0.087

H&W09008

M. moupinensis

1.10-1.37

1.237

0.065

H&W09010

M. patchoulii

1.04-1.19

1.103

0.054

H&W09011

M. patchoulii

1.02-1.14

1.087

0.042

Data-14. The standard deviation analysis of tooth ratio of species. Species

Variation range

Mean

Standard deviation

1. M. esquirolii

1.02-1.13

1.091

0.032

2. M. insuavis

1.03-1.21

1.106

0.042

3. M. patchoulii

1.01-1.19

1.102

0.040

4. M. mollis

1.01-1.21

1.112

0.043

5. M. delavayi

1.68-2.31

1.903

0.160

6. M. wardii

1.47-2.02

1.668

0.183

7. M. urticifolia

1.83-2.24

2.033

0.152

8. M. albescens (holotype)

1.89

—

—

9. M. stenocalyx

1.47-1.60

1.520

0.041

10. M. miyiensis (holotype)

2.87

—

—

11. M. megacalyx

1.01-1.10

1.055

0.036

12. M. moupinensis

1.03-1.49

1.206

0.092

13. M. prainiana

1.01-1.13

1.050

0.037

14. M. muliensis

1.08-1.21

1.143

0.043

15. M. omeiensis

1.42-1.62

1.500

0.046

16. M. brevipedunculata

1.40-1.43

1.412

0.016

17. M. nepalensis

1.38-1.67

1.463

0.106

18. M. robusta

1.10-1.20

1.140

0.041

19. M. vanchingshanensis

1.08-1.17

1.127

0.036

DATA OF STANDARD DEVIATION ANALYSIS

135

Data-15. The standard deviation analysis of corolla length of sampled populations. Population

Species

Variation range

Mean

H&W08004

M. mollis

1.40-1.60

1.513

Standard deviation 0.063

H&W08005

M. mollis

1.24-1.64

1.500

0.091

H&W08006

M. esquirolii

1.02-1.53

1.303

0.162

H&W08007

M. esquirolii

1.33-1.63

1.489

0.103

H&W08009

M. patchoulii

1.45-1.49

1.477

0.017

H&W09001

M. insuavis

1.26-1.40

1.324

0.050

H&W09002

M. insuavis

1.30-1.37

1.328

0.025

H&W09003

M. mollis

1.44-1.51

1.485

0.029

H&W09004

M. mollis

1.33-1.71

1.538

0.119

H&W09006

M. mollis

1.42-1.77

1.539

0.091

H&W09007

M. moupinensis

2.75-3.67

3.016

0.280

H&W09008

M. moupinensis

2.70-3.44

3.004

0.253

H&W09010

M. patchoulii

1.43-1.56

1.507

0.036

H&W09011

M. patchoulii

1.48-1.52

1.490

0.013

Data-16. The standard deviation analysis of corolla length of species. Species

Variation range

Mean

Standard deviation

1. M. esquirolii

1.02-1.63

1.361

0.143

2. M. insuavis

0.78-1.40

1.191

0.166

3. M. patchoulii

1.10-1.64

1.438

0.123

4. M. mollis

1.24-1.77

1.530

0.096

5. M. delavayi

1.47–2.53

2.026

0.305

6. M. wardii

1.47-2.28

1.855

0.266

7. M. urticifolia

1.62-2.64

2.015

0.422

8. M. albescens (holotype)

2.08

—

—

9. M. stenocalyx

1.91-2.48

2.201

0.165

10. M. miyiensis (holotype)

2.18

—

—

11. M. megacalyx

1.73-2.70

2.297

0.370

12. M. moupinensis

1.64-3.67

2.829

0.420

13. M. prainiana

1.62-2.54

2.163

0.230

14. M. muliensis

1.66-2.60

1.994

0.272

15. M. omeiensis

1.67-2.75

2.326

0.356

16. M. brevipedunculata

1.79-1.85

1.822

0.029

17. M. nepalensis

1.87-2.36

2.103

0.168

18. M. robusta

1.48-2.16

1.906

0.265

19. M. vanchingshanensis

1.68-1.76

1.714

0.036

136

A MONOGRAPH OF THE GENUS MICROTOENA (LAMIACEAE)

Data-17. The standard deviation analysis of corolla tube ratio of sampled populations. Population

Species

Variation range

Mean

Standard deviation

H&W08004

M. mollis

0.12-0.17

0.138

0.016

H&W08005

M. mollis

0.11-0.17

0.130

0.014

H&W08006

M. esquirolii

0.16-0.19

0.177

0.009

H&W08007

M. esquirolii

0.18-0.20

0.188

0.007

H&W08009

M. patchoulii

0.19-0.24

0.214

0.014

H&W09001

M. insuavis

0.12-0.15

0.135

0.010

H&W09002

M. insuavis

0.11-0.13

0.124

0.006

H&W09003

M. mollis

0.11-0.15

0.127

0.012

H&W09004

M. mollis

0.11-0.15

0.126

0.014

H&W09006

M. mollis

0.11-0.16

0.127

0.012

H&W09007

M. moupinensis

0.66-0.71

0.683

0.013

H&W09008

M. moupinensis

0.63-0.72

0.674

0.022

H&W09010

M. patchoulii

0.19-0.24

0.218

0.012

H&W09011

M. patchoulii

0.18-0.24

0.214

0.023

Data-18. The standard deviation analysis of corolla tube ratio of species. Species

Variation range

Mean

Standard deviation

1. M. esquirolii

0.16-0.20

0.183

0.009

2. M. insuavis

0.11-0.15

0.128

0.008

3. M. patchoulii

0.18-0.25

0.220

0.014

4. M. mollis

0.11-0.17

0.128

0.013

5. M. delavayi

0.51-0.64

0.577

0.031

6. M. wardii

0.59-0.66

0.621

0.021

7. M. urticifolia

0.52-0.55

0.531

0.010

8. M. albescens (holotype)

0.59

—

—

9. M. stenocalyx

0.52-0.57

0.542

0.017

10. M. miyiensis (holotype)

0.62

—

—

11. M. megacalyx

0.52-0.55

0.528

0.012

12. M. moupinensis

0.63-0.72

0.677

0.016

13. M. prainiana

0.54-0.63

0.581

0.023

14. M. muliensis

0.51-0.54

0.525

0.011

15. M. omeiensis

0.60-0.65

0.621

0.016

16. M. brevipedunculata

0.52-0.55

0.535

0.013

17. M. nepalensis

0.48-0.53

0.509

0.020

18. M. robusta

0.53-0.58

0.552

0.020

19. M. vanchingshanensis

0.55-0.58

0.566

0.013