UNIVERSITY OF LONDON
IMPERIAL COLLEGE OF SCIENCE AND TECHNOLOGY
Department of Botany
A numerical - taxonomic study of the Labiatae tribe Pogostemoneae ( Benth. ex Endl. ) Briq.
by
John Robert Press, B.Sc.
Thesis submitted for the degree of Master of Philosophy in the Faculty of Science.
LONDON, January 1981 ii.
ABSTRACT.
A numerical study incorporating cluster analyses and ordination methods was used to investigate the taxonomy of 138 species from ten genera of Labiatae tribe Pogostemoneae on the basis of morphological variation in sixty four characters. On the basis of the results the tribe Pogostemoneae is divided into five subtribes, the divisions being supported by other independent data sets of pollen and stomata morphology. Seven genera, Keiskea, Tetradenia, Eurysolen,
Rostrinucula, Colebrookea, Comanthosphace and Leucosceptrum are upheld unchanged. The union of Comanthosphace and Leucosceptrum
( Kitamura & Murata 1962 ) is rejected. A reassessment of the importance of the so-called " tribal characters " within the Labiatae is required to solve this problem. Pogostemon and Dysophylla are considered to be congeneric as Pogostemon sensu lato. The sections suggested by Keng ( 1978 ), section Pogostemon and section Eusteralis, are supported although the diagnosis for each one is modified.
Seventeen new combinations are made and two new names mentioned in
Pogostemon. The genus Elsholtzia was formerly divided into three sections. Two of these, sections Elsholtzia and Cyclostegia are now considered to be consectional and Cyclostegia is reduced to synonomy.
The third section, section Aphanochilus, formerly contained two series. Now, series Platyelasmeae is raised to the level of section while the species of series Stenelasmeae remain in section
Aphanochilus sensu stricto. A number of species within the various genera are indicated as being conspecific. iii.
TABLE OF CONTENTS.
1. INTRODUCTION 1
2. HISTORICAL REVIEW 7
2.1. Taxonomic position of the Pogostemoneae ?
2.2. Previous taxonomic treatments of the
Pogostemoneae 10
3. MATERIALS AND METHODS 26
3.1. Materials 26
3.2. Methods 2?
3.2.1. Reconstitution of material 27
3.2.2. Compilation of the data set 2?
3.2.3. Measurements of characters 3^
3*3. Numerical methods
3.3«/|. Selection of characters 35
3.3.2. Coding of character states ¥4
3.3.3. Similarities : match and mis-match 45
3.3.4. Assessment of results 4?
3.3.5. Missing values 4?
3.3.0. Sampling ••••• 48
3.3.7. Dependent characters 48
3.4. The analyses 49
3.4.1. Similarity matrices 49
3.4.2. Nearest neighbours 49
3-4.3. Single-linkage analysis 50 iv.
Principal co-ordinates analysis 51
3.4.5. Clustering to maximise within-
group mean similarity ( WGMS ) ••••• 53
3.4.6. Second runs of analyses 5^
3.5. Summary 55
4. CHARACTER VARIATION %
4.1. Leaves 5^
4.2. Heterophylly 5$
4.3. Leaf size 5$
4.4. Phyllotaxis 58
4.5. Petiole 65
4.6. Bracts 65
4.7. Bracteoles 66
4.8. Indumentum 67
4.9. Calyx 68
4.10. Corolla %
4.11. Stamens 77
4.12. Style 78
4.13. Disc 79
4.14. Nutlets 79
4.15* Inflorescence 80
5. RESULTS OF ANALYSES 87
5.1. The similarity matrices ...••••• 87
5.2. Clustering to maximise within- group
mean similarity ( WGMS ) 92 V.
5.3- Principal co-ordinates analysis 10?
5,4. Single-linkage analysis 126
6. DISCUSSION OF RESULTS 133
6.1. Ranks 133
6.2. Subtribes 134
6.3* Subtribe Comanthosphacineae 144
6.3-1 • Comanthosphace and Rostrinucula 144
6.3.2. Rostrinucula 148
6.4. Subtribe Elsholtzineae 152
6.4.1. Elsholtzia 152
6.4.2. Elsholtzia sections Elsholtzia
and Cyclostegia 153
6.4.3- Elsholtzia section Aphanochilus
series Stenelasmeae 159
6.4.4. Elsholtzia section Aphanochilus
series Platyelasmeae 163
6.4.5. Relocated 0TU»s 170
6.4.6. Elsholtzia beddomei and E.
kachinensis 170
6.4.7. Elsholtzia penduliflora 170
6.4.8. Elsholtzia hunanensis, E. flava
and E. fruticosa 171
6.4.9- Elsholtzia concinna 172
6.4.10. Leucosceptrum plectranthoideum 173
6.4.11. Dysophylla mairei 173
6.4.12. Elsholtzia integrifolia 174 vi.
6.4.13. Elsholtzia chinense 175
6.4.14. Distribution of Elsholtzia 176
6.4.15. Keiskea 177
6.4.16. Tetradenia 179
6.5. Subtribe Eurysolenineae 182
6.5.1. Eurysolen 182
6.6. Subtribe Pogostemoneae 187
6.6.1. Pogostemon and ffysophylla ...... 187
6.6.2. Subgeneric divisions 20k
6.6.3. Relocated OTU's 208
6.6.4. Elsholtzia aquatica 208
6.6.5. Distribution of Pogostemon 210
6.7. Subtribe Colebrookineae 210
6.7.1. Colebrookea 210
7. GENERAL DISCUSSION 213
8. TAXONOMIC CONSPECTUS 216
8.1. Key to genera and sections 216
8.2. Conspectus 219
ACKNOWLEDGEMENTS 232
REFERENCES 233
APPENDICES.
1. Specimens used for scoring and vii.
producing averaged data 241
2. Coded data scored for 138 OTU's 24-7
3. Full, similarity matrix 253
4. Simplified similarity matrix 274
5. Simplified similarity matrix run 2A 275
6. Simplified similarity matrix run 2B 276
7. The five nearest neighbours and their
similarities for each OTU 277
8. Distribution of OTU's in the six and
nine group schemes for the clustering to
maximise WGMS analysis 282
9. Distribution of OTU's in the four and
five group schemes for the clustering to
maximise WGMS analysis run 2A 284
10. Distribution of OTU's in the four and
five group schemes for the clustering to
maximise WGMS analysis run 2B 285
LIST OF FIGURES.
1. Distribution of the Pogostemoneae 2
2. Diagrammatic scheme showing the major
changes in the taxonomy of the Pogostemoneae 21
3. Leaf shapes, margins and petioles in
the Pogostemoneae •••• 59
4. Phyllotaxis in the Pogostemoneae 61
5. Bract types in the Pogostemoneae 63
6. Hair types in the Pogostemoneae 70 viii.
7. Calyx shapes and vejnation in
the Pogostemoneae 72
8. Corolla structure in the Pogostemoneae 74
9. Stamen types in the Pogostemoneae 81
10. Style and disc types in the Pogostemoneae 85
11. Nutlet types in the Pogostemoneae 85
12. Simplified diagram of the nine-group scheme
in the simplified similarity-matrix 88
13. Simplified diagram of the six-group scheme
in the simplified similarity-matrix 89
,15i16. Two-dimensional plots of the principal co-
ordinates analysis using the four largest
eigenvalues as axes :
14. vector 1 v. vector 2 110
15* vector 1 v. vector 3 111
16. vector 1 v. vector 4 112
17,18,19- Two-dimensional plots of the principal co-
ordinates analysis run 2A using the four
largest eigenvalues as axes.
17. vector 1 v. vector 2 114
18. vector 1 v. vector 3 115
19- vector 1 v. vector 4 116
20,21,22* Two-dimensional plots of the principal co-
ordinates analysis run 2A showing the
distribution of species of Elsholtzia.
20. vector 1 v. vector 2 118
21. vector 1 v. vector 3 119 ix.
/ 22. vector 1 v. vector 4 120
23,24,25. Two-dimensional plots of the principal co-
ordinates analysis run 2B using the four
largest eigenvalues as axes.
23. vector 1 v. vector 2 122
24. vector 1 v. vector 3 123
25. vector 1 v. vector 4 124
26. A single-linkage dendrogram and the box-
in-box classification derived from it 126
27. Single-linkage dendrogram for 138 OTU's 130
28. Diagrammatic representation of the BGMS
values for subtribes derived from a
combination of all analyses 143
29. Rostrinucula dependens 149
30. Elsholtzia luteola 157
31. Elsholtzia densa 168
32. Eurysolen gracilis 185
33« Dysophylla glabrata 190
3^,35i36. Two-dimensional plots of the principal co-
ordinates analysis run 2B.
34. vector 1 v. vector 2 •••••••••••••••• 194
35• vector 1 v. vector 3 195
36. vector 1 v. vector 4 196
37- Pogostemon micangensis 202
38. Distribution of Pogostemon
sensu lato 209 X.
LIST OF TABLES.
1. Summary of the classifications of the Labiatae by
Endlicher ( 1838 ), Bentham ( 1848 ) and Briquet ( 1897 ) 8
2. Numbers of nuclei and colpi in pollen grains
of the Pogostemoneae 25
3. The OTU's used in the analyses 28
4. The characters scored, their states and character types
according to Gower's coefficient of similarity 36
5. Designation of character types according to
Gower's coefficient of similarity 46
6. WGMS and BGMS values for six groups in the clustering
to maximise WGMS analysis 93
7. WGMS and BGMS values for nine groups in the clustering
to maximise WGMS analysis 94
8. WGMS and BGMS values for ten groups in the clustering
to maximise WGMS analysis 95
9. WGMS and BGMS values for species of Comanthosphace,
Rostrinucula and Leucosceptrum canum 99
10. WGMS and BGMS values for species of Comanthosphace,
Rostrinucula and Comanthosphace nanchuanensis 99
11. WGMS and BGMS values for four groups in the clustering
to maximise WGMS analysis run 2A 100
12. WGMS and BGMS values for five groups in the clustering
to maximise WGMS analysis run 2A 101
13. WGMS and BGMS values for six groups in the clustering
to maximise WGMS analysis run 2A 102
14. WGMS and BGMS values for five groups in the clustering xi.
to maximise WGMS analysis run 2B 105
15. WGMS and BGMS values for four groups in the clustering
to maximise WGMS analysis run 2B 106
16. Name hierarchy for eight taxa represented in the box-
in-box classification shown in fig. 26 128
17. Classification of the Pogostemoneae produced by use
of three phenon-lines in the single-linkage dendrogram 131
18. Six arrangements of subtribes within the Pogostemoneae
derived from different analyses 135
19. WGMS and BMGS values for subtribes, derived from the
principal co-ordinates and clustering to maximise
WGMS analyses ( arrangement one ) 138
20. WGMS and BGMS values for subtribes, derived from
the single-linkage analysis ( arrangement two ) •••• 139
21. WGMS and BGMS values for subtribes, derived from
a combination of all analyses ( arrangement three ) 140
22. Bract characters in Elsholtzia sections Elsholtzia,
and Cyclostegia Bent ham ••••• 155
23. Bract characters in Elsholtzia 166
24. Species which share high similarities with Dysophy11a
and Pogostemon in two analyses ••••• ••••• 189
25. Names and diagnoses of Briquet's subdivisions
of Pogostemon 205
26. Names and diagnoses of Briquet's subdivisions
of Dysophylla 206 1
1. INTRODUCTION,
The Pogostemoneae is a tribe containing some 202 species divided among ten genera. The group has a wide distribution through the warm temperate and tropical regions of the Old World ( fig. 1 ).
Most species are herbs or suffruticose perennials, some e.g. species of Dysophylla are aquatic or amphibious and a few e.g. Elsholtzia fruticosa and Colebrookea oppositifolia are shrubs.
Several species, particularly of Pogostemon and Elsholtzia, have commercial uses and are cultivated on a limited scale. The most important are Pogostemon heynianus which provides the patchouli of perfumery and Elsholtzia ciliata which is used as a condiment, carminative and popular drug in folk-medicine. It is grown in France for use in perfumery and the oil has an antibiotic effect. A number of other species are used as condiments and in folk-medicine, especially in eastern and south-eastern Asia.
Pogostemon mutanaba ( fig. 37 ) from southern tropical Africa produces starchy tubers which are eaten by the local people. A number of species are valuable food sources for bees, e.g.
Pogostemon parviflorus, from which pangol honey is produced.
This relatively small but quite well-known tribe has been rather neglected by recent workers in the Labiatae. Studies on anatomy, essential oils, susceptibility to rust-fungi and especially pollen morphology indicate the need for a reappraisal of the taxonomy of the Pogostemoneae. The integrity, status and phenetic relationships of this group as a tribe are also in doubt. While it H' oq
t> H' CctO H H' a1 £ c+ H- O P O Hj
&C+ ro 3 is not within the scope of this study to offer solutions to problems at the tribal level, those problems within the tribe are enumerated below. 1) Generic delimitations in Pogostemon and Dysophylla. The distinction between these genera is unclear. Many earlier authors regarded the two as distinct genera while others ( Hasskarl 1842, Miquel 1859i Kuntze 1891 ) expressed doubts about the validity of such a separation. Such differences of opinion are still widespread today. Keng ( 1978 ) states that a single character, phyllotaxis, is insufficient for a separation of genera and reduces the status of Dysophylla to Pogostemon section Eusteralis. On the other hand El-Gazzar & Watson ( 1967 ) maintain Dysophylla at generic rank based on a number of micro-characters such as phyllotaxis, leaf shape, indumentum and crystal inclusions. 2) Status and relationships of Comanthosphace and Leucosceptrum. On the basis of the tribal characters of basal nutlet-attachment, gynobasic style and confluent anther-locules, Comanthosphace is usually placed in the Pogostemoneae ( e.g. Briquet 1897 )• Leucosceptrum, with lateral nutlet-attachment, terminal style and unilocular anthers is placed in the Ajugoideae ( e.g. Bentham 1832^36, Briquet 1897 )• Kitamura & Murata ( 1962 ) believed these characters to be misconstrued in Leucosceptrum and considered it- congeneric with Comanthosphace, removing the redefined genus to the Satureieae. This view has not been widely accepted, leaving two problems; are Leucosceptrum and Comanthosphace congeneric and if so, in which of the three tribes, Ajugeae, Satureieae or jpogostemoneae should Leucosceptrum sensu Kitamura & Murata be placed? 3) Phenetic relationships of Rostrinucula. The formerly monotypic genus Rostrinucula is based on Elsholtzia dependens, a species which Rehder ( 1917 ) placed in Elsholtzia on the grounds of its broadly-bracted spikes, although it differed in the corolla shape, the presence of an annulus and the rostrate nutlets. Wunderlich's work ( 1963 ) suggests that Rostrinucula is both phenetically and phylogenetically closer to Comanthosphace, Pogostemon and Dysophylla than to Elsholtzia and therefore the systematic value of the bract, flower and nutlet characters needs to be re-examined before any firm conclusions can be made. k) Phenetic relationships of Keiskea. Little is known about the genus Keiskea, which in most classifications ( e.g. Bentham & Hooker 1876, Briquet 1897 ) is placed close to Elsholtzia. The genus is recognised on the basis of the deeply-divided calyx, the four-lobed corolla and the bilocular anthers. Several new species have been published in recent years and the availability of additional material will allow these characters to be investigated more fully. This may help to give a clearer indication of its phenetic relationships. 5) Phenetic relationships of Tetradenia. This unusual genus is a Madagascan endemic, a place where no other Pogostemoneae occur. Like Keiskea little is known of its phenetic relationships, some authors placing it next to Elsholtzia ( Bentham 1832 •..36, Endlicher 1838 ) while others place it with Pogostemon, Dysophylla and Colebrookea ( Bentham 8c Hooker 1876 ). All of these authors knew Tetradenia as a monotypic genus. It has now been expanded to include three species but no recent assessment of 5 its affinities has been carried out. 6) Phenetic relationships of Eurysolen. The genus Eurysolen is not usually considered to be a member of the Pogostemoneae and is placed more often in the tribe Ajugeae on the basis of the oblique attachment of the nutlets ( Briquet in Prain, 1901, Mukerjee ^^bO ). Chermisirivathana ("1963 ) and, Keng C 1969,1978 ) have expressed doubts as to the validity of this since the fruit characters agree well with other genera in the Pogostemoneae. 7) Subgeneric groupings. Problems of subgeneric groupings mainly concern species of Elsholtzia which is usually divided into three sections and two series. Some authors, e.g. Kudo ( 1929 ), have recognised some sections but not others, often without commenting on their decisions. Three species, E. densa, E. eriostachya and E. manshurica.have alternatively been recognised as a series ( series Platyelasmae ), a distinct genus ( Platyelasma ) and a series again by the same author ( Kitagawa 1935* 1939 ) while E. densa was recognised as another separate genus by Briquet ( 1908 ). In recent years many new species have been described without being assigned to any particular section. 8) Disjunct distributions. Elsholtzia and Pogostemon have very disjunct distributions. They are primarily Indo-Chinese and Malaysian plants, with one species of Elsholtzia and two of Pogostemon occuring in southern tropical Africa. These last three species are very isolated, the closest relative occuring in Madagascar. There are no species of either genus occuring in surrounding African areas or the near Middle-East. Previous treatments of relationships within the Pogostemoneae have all been based on relatively few characters. The problems require more radical solutions and a detailed method of assessment has been used here. To ensure a thorough analysis of all the available data numerical phenetic methods giving direct comparisons between groups have been used to investigate these problems. 7 / 2. HISTORICAL REVIEW. 2.1. TAXONOMIC POSITION OF THE POGOSTEMONEAE. The only complete monographic treatments of the Labiatae are those of Bentham ( 1832 - 36 , 1848 ). In his earlier work Bentham ( 1832 - 36 ) divided the family into eleven tribes, and described but did not give names to a number of divisions within these tribes. Endlicher ( 1838 ) accepted Bentham1 s arrangement and also assigned the rank of subtribe to Bentham1s unnamed divisions. Subtribes Pogostemeae ( containing Pogostemon and Dysophylla ) and Elsholtzieae ( containing Elsholtzia and Tetradenia ) were placed in the tribe Menthoideae ( see table 1 ). In his second account of the family Bentham ( 1848 ) rearranged his tribes, reduced the number to eight and recognised a number of subtribes. His subtribe Elsholtzieae ( containing Pogostemon, Dysophylla, Colebrookea, Tetradenia and Elsholtzia ) was placed in the tribe Satureieae ( see table 1 ). Later classifications were minor variations of that of Bentham until Briquet ( 1897 ) produced an apparently very different classification, with more subdivisions than in the earlier ones. He recognised eight subfamilies, fourteen tribes and eleven subtribes ( see table 1 ). Although Briquet altered much of the order of Benthamfs and Endlicher's systems, the essential difference between the classifications is one of rank, Briquet's divisions generally being made at one rank higher than those of the other authors. For example Endlicher's tribes Stachydeae, Nepeteae and Monardeae were retained by Briquet in the subfamily Stachyoideae. The subtribes Salvieae, Hormineae, Marrubieae, Bagostemoneae and Meriandreae were raised to tribes and only V Endlicher, 1838 ( based Bentham, 1848 Briquet, 1897 on Bentham, 1832 - 36 ). Subfamilies, Tribes & subtribes Tribes & subtribes Tribes 8c subtribes I Ocimoideae I Ocimoideae Ajugoideae a H 1. Moschosmeae 3 unnamed divisions 1. Ajugeae g. (D 2. Plectrantheae II Satureieae 2. Rosmarineae II Prostantheroideae 00 3. Hyptideae 1. Elsholtzeieae v>l 00 4. Lavanduleae 2. Menthoideae III Prasioideae IP Menthoideae 3. Thymeae IV Scutellarioideae a c+ {3- 1. Pogostemeae 4. Melisseae V Lavanduloideae 2. Elsholtzieae 5. Genera anomala VI Stachyoideae OO 3. Mentheae III Monardeae 1 • Marrubieae £ Meriandreae IV Nepeteae 2. Perilomieae IH Monardeae V Stachydeae 3. Nepeteae H- 1• Salvieae 1. Scutellarieae 4. Stachydeae g e+ 2. Rosmarineae 2. Melitteae a. Prune11inae ^ 00 3. Hormineae 3. Marrubieae b. Melittinae jY Satureineae 4. Lamieae c• Laminae ^ 1• Origaneae VI Prasieae 5. Glechoneae Endlicher, 1838 Bentham, 1848 Briquet, 1897 2. Hyssopeae VII Prostanthereae 6. Salvieae 3. Cunileae VII Ajugeae 7. Meriandreae V Mellisinae 8. Monardeae VI Scutellarineae 9. Hormineae VII Prostanthereae 10. Lepechinieae VIII Nepeteae 11. Satureieae IX Stachydeae a. Melissinae 1• Melitteae b. Hyssopinae 2. Lamieae c• Thyminae 3. Marrubieae d. Menthinae 4. Balloteae e. Perillinae X Prasieae 12. Pogostemoneae XI Ajugoideae VII Ocimoideae a. Hyptidinae b. Plectranthinae c. Moschosminae VIII Catopherioideae 10 Melittinae and Lamiinae retained as subtribes. Although some authors ( e.g. Erdtman 19^51 El-Gazzar & Watson 1970 ) have expressed dissatisfaction with Briquet's classification it is accepted by most modern authors ( e.g. Keng 1969 ) as being superior to those of earlier workers. Briquet regarded the Pogostemoneae as a tribe and his concept of the group is followed here. A brief synopsis of the nomenclature is given below. Tribe Pogostemoneae ( Benth. ex Endl. ) Briq. in Engl.& Prantl, Nat. Pflanzenfam. 4, 3a :326( 1897 )• Tribe Menthoideae subtribe Pogostemeae Benth. ex Endl., Gen. pi.: 612 ( 1838 ). Tribe Menthoideae subtribe Elsholtzieae Benth. ex Endl., Gen. pi. : 612 ( 1838 ). Tribe Menthoideae subtribe Mentheae Benth. ex Endl., Gen. pi. : 612 ( 1838 ) pro parte quoad Colebrookia sphalm. Tribe Satureieae subtribe Elsholtzieae Benth. in DC., Prodr. 12 : 149 ( 1848 ). Tribe Satureieae subtribe Pogostemoneae Benth. & Hook.f., Gen. pi. 2(2): 1162, 1164 ( 1876 ). Tribe Satureieae subtribe Menthoideae Benth. & Hook.f., Gen. pi. 2(2) : 1162, 1164 ( 1876 ) pro parte quoad Elsholtzia et Keiskea. Tribe Satureieae subtribe Pogostemoninae Kudo in Mem. Fac. Sci. Agric. Taihoku imp. Univ. 2 ( 2 ) : 45 ( 1929 )• 2.2. PREVIOUS TAXONOMIC TREATMENTS OF THE POGOSTEMONEAE. The Labiatae tribe Pogostemoneae:comprises^,some fcenr-currently 11 recognised genera, of which only three, Pogostemon, Dysophylla and Elsholtzia contain more than a dozen species. All genera within the tribe are relatively well-defined, with few new combinations of species being made during their history. However the taxonomic affinities of the genera, one to another within the Pogostemoneae, and in related tribes, have been subject to numerous assessments and interpretations. The early literature in particular is fragmentary and confused. Few authors agree on the importance that should be attached to the various characters, or indeed which characters should be used in the consideration of the affinities and subsequent groupings. The result has been a number of views at variance with each other and providing no clear picture of the structure of the tribe. The following account attempts to trace the history of the genera of the Pogostemoneae and the views of the authors in whose works these genera appear. The first currently recognised genus to be mentioned in the literature was Dysophylla to which Hermann ( 1717, 1726 ) gave the polynomial Veronica hirsuta latifolia Zeylanica aquatica. Linnaeus ( 17^7 ) used the generic name Alopecuro - Veronica. After publication of Species Plant arum ( 1753 ) Linnaeus did not recognise Dysophylla as a separate taxon but included it in his broad concept of Mentha, a heterogenous genus containing unrelated plants with naked stamens. He later ( 1767 ) included in Mentha two additional species, M. hirsuta and M. auricularia, of remote affinity and placed in synonomy of the latter the names Alopecuro - Veronica and Majoram foetidum ( a plant shown in an etching by Rumphius ( 1750 ) and amended his description by adding 11 filamenta pilosa 12 Roxburgh ( 1814 ) gave names to several Mentha species, all of which were nomina nuda. However, two of these were later taken up as M. verticillata and M. quadrifolia by Don ( 1825 )• In. the descriptions the stamens are again said to bearded. When Blume ( 1826 ) separated Dysophylla as a distinct genus based on Mentha auricularia L. he related it to, and placed it next to Mentha, but considered it to differ by the closure of the fruiting calyx, the fleshy swelling of the disc and the distinctly bearded stamens. Under the only species, D. auricularia, he cited Rumphius's illustration. ' Willdenow ( 1790 ) described a new genus, Elsholtzia, based on a Silesian plant, E. cristata. Although he gave a clear description and distinguished his genus from other Labiatae on corolla, calyx, stamen and inflorescence characters, a number of confused associations were made between this and other plants by later authors. Lamarck ( 1789 ) gave five Hyssopus species, but his rather vague limits of the genus allowed for inclusion of two species of Elsholtzia, one of which was Willdenow's type species. Persoon ( 1806 ) amended Willdenow's genus description to add 11 Flor.secundi, bracteati " and described two new species, E. paniculata ( referable to Pogostemon ) and E. ocymoides. Don ( 1825 ) considered Elsholtzia congeneric with Perilia and reduced the name to synonomy. The description differed from that of Perilla sensu Linnaeus (1764 ) on a number of points but closely matched that of Willdenow's Elsholtzia. Two new genera, Leucosceptrum and Colebrookea, were described by Smith ( 1805 ) based on Nepalese specimens. Leucosceptrum, because of its habit, corolla and deeply lobed disc was said to have 13 affinities with Verbenaceae, and because of the four-lobed corolla, exserted stamens and bilocular anthers, to be close to Mentha. Smith described the corolla as having four unequal segments, the upper deeply emarginate, the lower large and entire. However his illustration shows the opposite of this, the lower segment being emarginate and the upper entire. The second monotypic genus, Colebrookea, was;chiefly|characterised by its fruit morphology. The distinctive hairy calyx with its plumose teeth acts as a pappus for the single-seeded, dry fruit. Roxburgh ( 1815 ) published an account of a second species of Colebrookea, C. ternifolia, based on plants collected in Mysore. All later authors however, regard this merely as a variety of C. oppositifolia, leaving Colebrookea as a monotypic genus. C. oppositifolia is a shrub and very different from the herbaceous species of Elsholtzia. In fact they da share a number of floral characters. Poiret (1817} produced a rather confusing situation when he published a new generic name Elshotzia, and put into it Colebrookea oppositifolia, as a new combination. He even added the comment in the text 11 ce genre est le meme que le Colebrookea de Smith. II fait y reunir le barbula de Loureiro 11. The 11 barbula " of Loureiro ( 1790 ) is an earlier name for the Verbenaceae genus Caryopteris which is strikingly similar to the Pogostemoneae in calyx, corolla and anther characters. Desfontaines ( 1815 ) described a new genus, Pogostemon, to contain a species with bearded stamens, P. plectranthoides. Citing the shared similarity of calyx and corolla characters of Pogostemon and Hyssopus he claimed an affinity for his new genus; " Le genre Pogostemon a de I'affinite avec lfHyssope. La corolle renversee, les trois lobes de la levre superieure entiere et arrondis au sommet, les filets des etamines abaisses et barbus, sont les principaux characteres qui le distinguent Blume ( 1826 ) followed Desfontaines• description and published a second species from Java, P. menthoides, but which had naked filaments. The first comprehensive review of all the genera together was made by Bentham ( 1829 )• He considered the original descriptions of Dysophylla and Pogostemon unsatisfactory and suggested modifications for both. In Dysophylla he placed less emphasis on the connivence of the calyx teeth since it is a feature not common to all species. In Pogostemon he regarded the declination of the stamens to be so slight as to be of little significance* and he questioned Blume's inclusion of P. menthoides since it appeared anomalous with the absence of hairs on the stamens, hairy stamens being a constant feature of all the other species of both genera. The delimination of Dysophylla was expanded by Bentham to include those species of Mentha described by Roxburgh ( 1814, 18^2 ) and Loureiro ( 1790 ). Bentham reduced Elsholtzia to only one species, E. cristata Willd. He created three new genera, all closely related to Elsholtzia: Aphanochilus, very similar to Elsholtzia, but differing in the exserted stamens with anther-locules confluent, the shrubby habit and non-secund inflorescence ( the Per ilia species described by Don ( 1825 ), although unknown to Bentham, belonged to Aphanochilus ); the monotypic Cyclostegia, easily distinguished by its densely strobilate inflorescence, and Tetradenia, a Madagascan genus based on a plant labelled Mentha fruticosa in Hooker's herbarium, which differs from the other two genera in the irregular calyx and bright red, glandular 15 swellings around the nutlets for which it is named, A year later Bentham ( 1830 ) wrote a second account of the Labiatae. The only change from the earlier work was for Dysophylla, where he divided the genus into two parts on the basis of phyllotaxis. The first was characterised by opposite, paired leaves and agreed with Blume's original description, and contained three species, D. auricularia, D. strigosa and D. myosuroides. The second contained all of the Dysophylla species with verticillate leaves. The first complete treatment of the Labiatae was by Bentham ( 1832 — 1836 ) in which he repeated his earlier descriptions for most of the genera and formally recognised his own divisions of Dysophylla ( 1830 ) as sections Oppositifoliae. and Verticillatae. On the basis of inflorescence characters he divided Pogostemon into § Paniculatae and § Racemosae. Aphanochilus and Cyclostegia,however, were reduced to sections.of Elsholtzia, the distinctions being based primarily on morphological differences of the inflorescence and bracts: Section I. Aphanochilus :- Spicae saepius laxae aequales vel subsecundae folus florabilus lanceolatis vel subulatis. Section II. Cyclostegia :- Spicae densae. Folia floralia connata imbricata cyathiformia membranacea venosa margine ciliata. Section III. Elsholtzia :- Spicae et folia floralia lato-ovata secunda. Bentham considered Leucosceptrum Smith to be a section of the unrelated genus Teucrium in his tribe Ajugoideae, which it resembled in lacking the upper lip of the corolla, in the hairiness of its fruits and in its semi-shrubby habit. Bentham's division of Dysophylla into sections Opposit ifoliae and Verticillatae was taken further by Rafinesque ( 18^7 ) who published 16 the genus Eusteralis based on Mentha pumila Graham and M. verticillata Roxburgh which effectively raised section Verticillatae to generic rank. The problems posed by Dysophylla were also reflected by Hasskarl ( 1842 ), Miquel ( 1859 ) and Kuntze ( 1891 ) who were unable to distinguish it adequately from Pogostemon, and included it as a section of that genus. Miquel ( 1865 ) described four, new species of Elsholtzia, three of which he referred to Bentham's section Cyclostegia. Bentham ( in Bentham & Hooker, 1876 )., 0n the advice of Maximovicz referred them to Pogostemon (see Hooker 1896\ This arrangement was based on the similarity of the anther structure, the anthers being subglobose, imperfectly two-celled and two-valved as in Pogostemon, and not ovoid, distantly two-celled with the cells two-valved as in Elsholtzia. One year later S. Moore ( 1877 ) removed Miquel's four species of Elsholtzia and placed them in the new genus Comanthosphace. Moore's view was that Comanthosphace approached Elsholtzia more nearly than Pogostemon, particularly in the two-lipped, five-lobed form of the corolla. This arrangement was fairly reliable but Moore mistakenly described the verticillasters as being obscurely bracteate, an error pointed out by Hooker ( 1896 ). In fact the bracts are very large, another feature in common with Elsholtzia, but often caducous. •Hemsley ( 1890 ) described a Chinese plant which he provisionally placed in the genus Caryopteris ( Verbenaceae ) as C. ningpoensis, although he felt it resembled a species of Buddieia (* Buddiejaceae ) or Vitex ( Verbenaceae ) more than any other Caryopteris species. The confusion was largely removed when Handel-Mazzetti ( 1896 ) realised that it showed more affinities with Moore's genus Comanthosphace. 17 In the same paper as he described four Elsholtzia species Miquel ( 1865 ) proposed a new genus, Keiskea , to accomodate material collected in Japan. When considering its affinities he gave its position as close to Mentha although the description suggested closer affinities to Elsholtzia in that the corolla was sufybilabiate with an emarginate upper lip and three-lobed lower lip. The filaments were hairy at the base, forming an incomplete annulus. Miquel described one species, K. japonica. Keiskea remained monotypic until Diels C 1924 ) described a second species from China, K. sinensis. A third species, K. elsholtzioides was described by by Merrill ( 1937 )• He had difficulty in determining the affinities of this species and commented: 11 In making the preliminary examinations this was placed in Comanthosphace from which its calyx characters exclude it. In its characteristic persistent, broad bracts it resembles Elsholtzia but in spite of these, seems to belong in Keiskea." K. elsholtzioides differs markedly from the two previously known species of Keiskea which have narrower bracts and broadly campanulate calyces. Masamune ( 19^0 ) described a Formosan species of Keiskea, K. macrobracteata. The conspicuous characters, bilabiate calyx and broadly ovate bracts, which distinguishes it from K. .japonica and K. sinensis, prompted him to propose dividing the genus into two sections, Macrobracteatae and Eukeiskea: Section I. Macrobracteatae. Braetea ovata-rotundata. Carex ( sic ) bi-labiatus supra barbatus. Section II. Eukeiskea. Braetea linearis. Carex ( sic ) late campanulatus extus subglaber. Merrill's K. elsholtzioides, of which Masamune was unaware, would 18 belong to the first section. The next major work on the Labiatae after that of Bentham was written by Briquet ( 1897 )• His account of the Pogostemoneae had no major changes from the classification originally presented by Bentham ( 1832 - 36 ). However he described new subgeneric groupings in three genera. In Pogostemon he divided Bentham's § Racemosae into? A. Qlabriuscula with naked filaments and B. Barbata with hairy filaments.^Paniculatae Benth. was also divided into two groups designated only as A and B and distinguished by the density of the verticillasters ( see table 25). Dysophylla was divided into two sections, section Goniocalicinae, with strongly five-angled calyces and section Rhabdocalicinae, with cylindrical calyces. Section Rhabdocalicinae was further divided by annual or perennial habit ( see table 26). Two new series were described in Elsholtzia section Aphanochilus, series Platyelasmeae, containing only E. densa and E. eriostachya and distinguished by broad bracts and matt nutlets and series Stenelasmeae, containing the remaining species and distinguished by narrow bracts and shiny nutlets. In a later paper Briquet ( 1908 ) described a new genus from a Pamir plant collected by Paulsen, which he appropriately named Paulseniella pamirensis. The description and accompanying illustration, however, showed this to be a plant of Elsholtzia densa Benth., a fact pointed out by Fedschenko ( 1908 ) although he referred to the species as P. wakhanica. However the text makes it clear he is referring to the same plant as Briquet. Another Elsholtzia species of dubious position was described by Rehder ( 19^7 ) as E. dependens. " This species seems not closely related to any other species in the genus. According to its broadly 19 bracted spikes it ought to be placed in the section Aphanochilus Bentham, but it differs from the species of this group as from those of the other groups in the entire upper lip of the corolla, in the irregular ring of hairs at the mouth of the corolla formed by hairy disc-like excrescences at the base of the filaments and in a hairy crescent-shaped crest below the base of the lower lip, and in the rostrate nutlets. The drooping habit of the long and slender spikes is also very peculiar and, so far as I know, does not occur in any other Elsholtzia 11. Rehder's views were supported by Kudo in his monograph of the Chinese Labiatae ( 1929 ). Howeve^ he took it one stage further and removed it from Elsholtzia and established a new, monotypic genus, Rostrinucula. Kudo also revived Bentham's Aphanochilus, considering it to merit generic rank. This was not wholly accepted by other authors since many considered Elsholtzia and Aphanochilus to be congeneric, the calyx, anther and bract characters being unreliable for separating the two. During the last fifty years, very few major systematic changes have been made within the Pogostemoneae. Kitamura & Murata ( 1962 ) proposed the union of Leucosceptrum and Comanthosphace, citing shape of calyx and corolla, exsertion of stamens, anther shape, areole size and presence of stellate hairs on the leaves as common characters. As the authors did not examine all the species of either genus, their views have not been generally accepted. El-Gazzar & Watson ( 1967 ) made a study of Pogostemon and Dysophylla, using leaf characters, presence or absence of crystals in the calyx and presence or absence of stem aerenchyma. They concluded 20 that Bentham's Dysophylla section Oppositifoliae ( containing four species, D. auricularia Bl., D. rugosa Hook, f., D. salicifolia Dalz. ex Hook, f. and D. myosuroides Benth. in Wall*. ) should be sunk into Pogostemon since it shared the characters of opposite, broad, petiolate leaves, crystals present in the calyx and stem-aerenchyma absent. Dysophylla section Verticillatae ( containing all other Dysophylla species ) was characterised by verticillate, linear, glabrous, sessile leaves, crystals absent from the calyx and stem- aerenchyma present. This transfer of four species of Dysophylla, including the type species D. auricularia Bl., to Pogostemon raised a nomenclatural problem for which a number of solutions were proposed and subsequently rejected. The latest and most satisfactory proposal, suggested by Bakhuizen van den Brink & van Steenis ( 1968 ) and also made by Panigrahi ( 1976 ), was to revive the name Eusteralis Rafinesque for Dysophylla section Verticillatae. Keng ( 1978 ), however, gave Pogostemon and Dysophylla as congeneric taxa and placed species of Dysophylla section Verticillatae in Pogostemon section Eusteralis. Wu 8c Huang ( 197^ ) in their precursor account for the Flora Reipublicae Popularis Sinicae and later ( 1977 ) in the Flora itself gave a generic account of the Chinese species of Elsholtzia. This covered thirty three species and produced two new subsections and eight new series. Although new species, especially from China, continue to be described and partial treatments of genera appear in various floristic accounts, no major treatment of the group has appeared in recent times. The major changes in the taxonomy of the group to date are summarised in fig. 2 . 21 Fig. 2 . Diagrammatic scheme showing the major changes in the taxonomy of the Pogostemoneae, The right hand margin shows the current names and status of the taxa. Dates for descriptions of genera and sections and any changes of status are shown along the top margin. Authors to whom the dates refer are shown in the key below. Genera are represented by the shadings given in the key below. Authors. Genera. 1767 Linnaeus Mentha 1790 Willdenow 1805 Smith Dysophylla 1815 Desfontaines 1826 Blume Pogostemon 1829 Bentham 1833 Bentham Leucosceptrum 1865 Miquel 1877 S. Moore Comanthosphace 1917 Rehder 1929 Kudo Rostrinucula 1941 Masamune 1962 Kitamura & Murata Aphanochilus 1967 El-Gazzar & Watson 1978 Keng Cyclostegia V section Verticillatae Elsholtzia O section Oppositifoliae Tetradenia R § Racemosae Keiskea p § Paniculatae Colebrookea • denotes first description of a genus or section A denotes change of status Reading horizontally across the diagram shows any changes undergone by a genus or section. For example species of Comanthosphace were originally placed in Elsholtzia by Miquel ( 1865 ) then recognised by S. Moore ( 1877 ) as forming a new genus, Comanthosphace, which Kitamura & Murata ( 1962 ) considered to be congeneric with Leucosceptrum, Reading vertically down the table from a given date shows the genera recognised at that time, although the author referred to may not have cited all those genera in his work. For example by 1826, Mentha, Pogostemon, Leucosceptrum, Elsholtzia and Colebrookea were recognised as valid genera. Of these, Blume cited only Pogostemon but he also described a new genus, Dysophylla, based on a species formerly included within Mentha. p. oq Mentha ro section Eusteralis Pogostemon section Pogostemon j Leucosceptrum tr *>>>>>>>>>>>>>>>>>> > Rostrinucula • D • 1° ° Ql era ••aaoaaaaaaaaaaa section Aphanochilus ro ro 23 2.3. REVIEW OF POLLEN STUDIES. In addition to morphological studies, work has been carried out on a number of other aspects of the Labiatae which throw light on the taxonomic position of the Pogostemoneae. The most important of these is undoubtedly the study of pollen. A survey of this work does not fit easily with the preceding historical review and is treated separately below. The first large scale work on Labiatae pollen was carried out by Leitner ( 19^2 ). She stated that at anthesis some species of Pogostemoneae have bi-nucleate pollen grains while others have tri- nucleate grains. As these characters are constant in other, larger taxonomic groups, she suggested that a comprehensive survey might show the necessity of splitting the tribe into sub-tribes based on the pollen types. Erdtman ( 19^5? 1952 ) first pointed out the correlation between tri-nucleate and hexa-colpate grains ( Fritzsche 1832, and others ) and bi-nucleate and tri-colpate grains ( Engler & Diels 1936 ). By matching the pollen morphological features and the cytological facts presented by Leitner, the bi-nucleate / tri-colpate and tri-nucleate / hexa-colpate relationship was established. He used this evidence to divide the Labiatae into two groups, tentatively ranked as sub-families. Pogostemon was placed in the bi-nucleate / tri-colpate group ( group I ). The Pogostemoneae excluding Pogostemon were placed in the tri-nucleate / hexa-colpate group ( group II ). He also suggested that the group name Pogostemoneae be discarded since the group is heterogenous, and cited as further evidence the work of Mayer ( 1909 ) which showed Pogostemon to be anatomically different 2k from the other members of the Pogostemoneae. Leitner ( 19^2 ) had claimed to have found bi-nucleate pollen grains in some species of Colebrookea, Keiskea, Comanthosphace and Dysophylla. Other species of Comanthosphace and Dysophylla were said to have hexa-colpate ( and therefore by implication, tri-nucleate ) grains ( Risch 1956 ). Wunderlich ( 19^3 ) also investigated the number of nuclei and colpi in the pollen and her results agreed with those of earlier workers with three exceptions. Keiskea was found to be tri-nucleate / hexa-colpate and Comanthosphace and Dysophylla bi-nucleate / tri-colpate. She also found that Rostrinucula dependens ( Rehd. ) Kudo with bi-nucleate / tri-colpate grains differed from Elsholtzia species with tri-nucleate / hexa-colpate grains, thus confirming Kudo's separation of this species. Wunderlich refined Erdtman's division of the group, placing the Pogostemoneae excluding Elsholtzia, Keiskea and Tetradenia in group I, and the latter three genera in group II. Various later workers have counted the number of colpi and / or nuclei for species within the Pogostemoneae. In particular El-Gazzar & Watson ( 1968 ) sampled pollen from 550 species from 125 genera of Labiatae. All the results agree with the pattern pointed out by Wunderlich ( 19&3 )• Table 2 summarises the data. Table 2 • Numbers of nuclei and colpi in pollen grains of the Pogostemoneae. Genus Number of nuclei Number of colpi per cell per ytxin Elsholtzia 3 6 Keiskea 3 6 Tetradenia 3 6 Eurysolen - 3 Comanthosphace 2 3 Leucosceptrum - 3 Rostrinucula 2 3 Pogostemon 2 3 Dysophylla 2 3 Colebrookea 2 3 26 3. MATERIALS AND METHODS. 3.1. MATERIALS. All materials used in this study were obtained from herbarium specimens. No live material was available with the single exception of a specimen of Elsholtzia ciliata Benth. growing in the Chelsea Physic Garden, London. Specimens for the study were obtained from the following herbaria : British Museum ( Natural History ), London ( BM )*; Royal Botanic Garden, Edinburgh ( E ); Conservatoire et Jardins Botaniques, Geneve ( G ); Royal Botanic Gardens, Kew ( K ); Kunming station of the Botanical Institute, Kunming ( KTJN ); Museum Nationale d'Histoire Naturelle, Paris ( P ); the Herbarium, Department of Botany, National Taiwan University, Taipei ( TAI ); Botanical Institute, Tokyo ( TI ). Most species of Pogostemoneae are relatively small and tend to be collected in their entirety. However some of the shrubby species may be represented by partial specimens, particularly the young branches. These tend to have a dense indumentum and often lack mature leaves. Identification of the study material presented a number of problems. Much of it was either unnamed or misidentified. Four main works were used in identifying specimens; those of Bentham ( 1832 - 6 ), Hooker ( 1885 ), Briquet ( 1897 ) and Keng ( 1978 ). When these works proved inadequate, the specimens were compared with the types. Obscure or poorly understood taxa were scored wherever possible from authentic specimens. * Abbreviations follow those given by Holmgren & Keuken ( 1974 ) Index Herbariorum pt. 1, ed. 6. 27 3.2. METHODS. 3.2.1. Reconstitution of material. Flowers and fruiting calyces were too brittle to be dissected without prior softening. The material was soaked in sodium- orthophosphate solution for twelve hours, washed in distilled water and stored in absolute alcohol. The advantage of this method over conventional methods such as boiling is that large amounts of material can be prepared in minimum time. Storage in alcohol is permanent and allows easy reference at a later date. 3-2.2. Compilation of the data set. The number of species and bulk of material available precluded the recording of every specimen for computation. Nevertheless, all available species were included in the sampling and all material was examined. To obtain a full set of data for a species, three typical specimens were selected for scoring as a basis for each operational taxonomic unit ( OTU ). The species used in this study, their OTU numbers and acronyms are given in table 3 . The data were then averaged to produce a single data set for that species. Where only one or two specimens were available the data set was produced from those specimens. Those specimens selected for recording are given in appendix 1 . Flowers at anthesis, ripe infructescences and, in some cases such as Leucosceptrum canum, young inflorescences were required as well as stems and leaves. Since these were seldom all represented in a single specimen, wherever possible data from a number of gatherings were recorded for each species. 28 Table 3 . The OTUfs used in the analyses. KJAP 1. Keiskea japonica CSTE 2. Comanthosphace stellipila CBAR barbinervis CSUB 4. sublanceolata CJAP 5. japonica CNIN 6. ningpoensis LCAN 7- Leucosceptrum canum CTER 8. Colebrookea ternifolia COPP 9- oppositifolia ECON 10. Elsholtzia concinna ECIL 11. ciliata EKAC 12. kachinensis EPYG 13. pygmaea ESOU 14. soulei EARG 15. argyi ELUT 16. luteola EBOD 17. bodinieri EHET 18. heterophylla ESTR 19. strobilifera EERI 20. eriostachya EDEN 21. densa EAQU 22. aquatica EINT 23. integrifolia EHUN 24. hunanensis EPEN 25. penduliflora Table 3 ( continued )• ERUG 26. Elsholtzia rugulosa ESTA 27. stauntoni EFLA 28. flava EFRU 29. fruticosa EPUB 30. pubescens ECOM 31- communis EALO 32. alopecuroides EGRI 33- griffithii EGLA 34. glanduligera EELA 35« elata EWIN 36. winitiana ESTA 37. stachyodea EBLA 38. blanda EMYO 39- myosurus EOCH 40. ochroleuca EPIL 41. pilosa ECAP 42. capituligera DTRI 43. Dysophylla trinervia DGLA 44. glabrata DTOM 45. tomentosa DPEN 46. pentagona DSTO 47. stocksii DSAM 48. sampsoni DPEG 49. peguana DGRI 50. griffithii DYAT 51• yatabeana DCRA 52. crassicaulis Table 3 ( continued )• DSTE 53- Dysophylla stellata DLIN 54. linearis DCRU 35- cruciata DQUA 56. quadrifolia DAUR 57- auricularia DSAL 58. salicifolia DRUG 59- rugosa DMYO 60. myosuroides PBEN 61. Pogostemon benghalei PPAR 62. parviflorus PPAN 63. paniculatus PTUB 64. tuberculosus PGLA 65. glaber PHEY 66. heyneanus PCAB 67. cablin PELS 68. elsHoltziodes PAMA 69. amarant ho ide s PFOR 70. formosanus PMEN 71. menthoides PFRA 72. fraternus PBRA 73. brachystachyus PMIC 74. micagensis PMUT 75. mutamba PNIG 76. nigrescens PPHI 77- phillipensis PREF 78. reflexus PRET 79. reticulatus Table 3 ( continued ). PMOL 80. Pogostemon mollis PTRA 81- travancoricus PATE 82. atropurpureus PSPE 83• speciosus PVEL 84. velatus PW1L 85. williamsii PPUR 86. purpurascens PPAL 87. paludosus PVIL 88. villosus PWIG 89. wightii PEOT 90. rotundatus PSTR 91. strigosus PHIS 92. hispidus PPUB 93. pubescens CFOE 94. Comanthosphace formosana EDEP 95. Eostrinucula dependens BSIN 96. sinensis LPLE 97. Leucosceptrum plectranthoideum ENIP 98. Elsholtzia nipponica EOLD 99. oldhami ECHI 100. chinense EPSE 101. pseudocristata DMAI 102. Dysophylla mairei DFAU 103. faurei DAND 104. andersoni DKOE 105. koehneana DGEA 106. gracilis Table 3 ( continued )• DLYT 107. Dysophylla lythroides DHEL 108. helferi DPUM 109. pumila PNEL 110. Pogostemon nelsoni PBAT 111. battakianus PWAT 112. wattii PHIR 113. hirsutus PRUP 114. rupestris PMAC 115. macgregori PGAR 116. gardneri PCHA 117. chaixii PDIE 118. dielsianus PGRI 119. griffithii PNIL 120. nilagiricus PLIT 121. litigiosus TGOU 122. Tetradenia goudotii THIL 123. hildbrantii TFRU 124. fruticosa EMAN 125. Elsholtzia manshurica EGRA 126. Eurysolen gracilis EFED 127. Elsholtzia feddei EELE 128. elegans EBED 129. beddomei KELS 130. Keiskea elsholtziodes KGLA 131. glandulosa KSIN 132. sinensis KSZE 133. szechuanensis Table 3 ( continued )• CNAN 134, Comanthosphace nanchuanensis PBRE 135. Pogostemon brevicorollus DTSI 136. Dysophylla tsiangii DFAL 137. falcata DSZE 138. szemacensis 34 Data on pollen were obtained from the literature except for Leucosceptrum canum and Eurysolen gracilis* Pollen samples from these two species were acetolysed and examined by light microscopy. 3-2.3- Measurement of characters. The characters used in the study, their states and character type according to Gower's coefficient of similarity ( Gower 1971 ) are given in table 4 • Petiole length ( character 2 ) was measured to the nearest millimetre. Bracteole and pedicel lengths ( characters 15 and 64 ) were measured to the nearest 0.3 millimetre. The lengths of the longest and shortest calyx tooth, calyx tube at anthesis and in fruit, corolla tube and upper and lower corolla lips ( characters 17, 18, 21, 22, 30, 32 and 33 respectively ) were all measured to the nearest 0.05 millimetre. The number of leaves per whorl ( character 4 ) was recorded only for those species with more than 2 leaves at a node ( character 3 )• Wherever possible the leaves were counted from whorls in the middle of the stem. Characters 6, 7 and 8 refer to overall shape, basal shape and apical shape respectively in the leaves. Each of the states of these characters represents a broad class of shape e.g. ovate. Each specimen was assigned to the most appropriate class and coded accordingly. A similar procedure was adopted with bract and bracteole shape ( characters 11 and 14 ). The coded data for each OTU is given in appendix 2. * denotes missing or non-applicable data. 3-3. NUMERICAL METHODS. Numerical methods are a relatively recent innovation and have only lately become widespread in phenetic taxonomy. A phenetic classification is one in which the groups are considered natural 35 in the sense of Gilmour ( Gilmour 19^0, 1961, see Farris 1977 ). That produced by numerical methods 11 reflects the characteristics of the organism as faithfully as the requirements of ultrametric representation and a classificatory simplicity permit " ( McNeil 1979 )• However there are a number of pitfalls for the numerical pheneticist e.g. choice of characters, arbitrary grouping methods and problems in clustering. 3*3-1. Selection of feharacters. ' A common misconception is that for a classification to be natural in the sense of Gilmour, all observable characters must be used ( cf.McNeil 1979 )• The inclusion of every character of every unit in a single study is impractical, even impossible. Moreover it is undesirable since some characters will be common to all units and therefore irrelevant or will show no pattern that can be associated with the groups produced in the analyses. The character states to be measured must be selected. However as Everitt ( 1977 ) concisely stated " it is important to bear in mind that the initial choice of variable is itself a categorization of the data which has no mathematical or statistical guidelines, and which reflects the investigator's judgement of relevance for the purpose of the classification ( this of course could also be saia of the entities chosen for study.)". McNeil's ( 1979 ) view is that " it is those characters which do show association with recognisable groups that are then emphasised in clarifying the groups ". Such emphasis of the characters is an a posteriori weighting and cannot influence the initial choice of characters. 36 Table The characters scored, their states and character types according to Gower's coefficient of similarity. Characters Type, ' 1. Leaves : type 1 0 homophyllus 1 heterophyllus 2. Leaves : petiole length 3 in millimetres 3. Leaves : phyllotaxis 2 0 opposite 1 verticillate Leaves : number per whorl 3 Leaves : comparative size of 2 members of a pair or whorl 0 equal 1 unequal 6. Leaves : shape 2 0 linear 1 ovate 2 orbicular 3 trifid 7. Leaves : basal shape 2 0 cuneate 1 attenuate 2 rounded 3 truncate 37 Table 4 ( continued ) 8- Leaves : apical shape 0 acute 1 acuminate 2 obtuse 9. Leaves : margin 0 entire 1 dentate 2 doubly dentate 10. Bracts : type Onot DiembraaoiJjs 1 membranous 11. Bracts : shape 0 linear 1 ovate 2 at least as broad as long 12. Bracts : fusion 0 free 1 connate for at least part of their length 13. Bracts : persistence 0 persistent 1 deciduous 14. Bracteoles 0 absent 1 linear 2 lanceolate 15. Bracteoles : length in millimetres Table 4 ( continued ). 16. Calyx : symmetry of teeth 0 regular 1 irregular 17* Calyx : length of longest tooth in millimetres 18. Calyx : length of shortest tooth in millimetres 19. Calyx : annulus in throat 0 absent 1 present 20. Calyx : number of main veins 21. Calyx : tube length at anthesis in millimetres 22. Calyx : tube length in fruit in millimetres 23. Calyx : tooth form in fruit 0 not plumose 1 plumose 24. Calyx : tooth position in fruit 0 incurved 1 erect 2 spreading 25. Calyx : teeth hairs 0 absent 1 present Table 4 ( continued ). 26. Calyx : tube hairs 0 absent 1 present 27. Calyx : angles 0 more or less terete 1 strongly angled 28. Corolla : symmetry 0 lower lip with 1 lobe 1 lower lip with 2 lobes 29. Corolla : division 0 upper lip not emarginate 1 upper lip emarginate 30. Corolla : tube length in millimetres 31. Corolla : exsertion 0 exserted 1 not exserted 32. Corolla : length of upper lip in millimetres 33• Corolla : length of lower lip in millimetres 34. Corolla : colour 0 not white 1 white 35• Corolla : colour 0 not yellow 1 yellow Table 4 ( continued ). 36. Corolla : colour 0 not purple 1 purple 37- Corolla : annulus 0 absent 1 complete ring of hairs 2 interrupted ring of hairs 38. Corolla : invagination 0 invagination absent ' 1 invagination present 39- Corolla : shape 0 corolla not gibbous above annvlus 1 corolla gibbous above annulus £ 40. Stamens : exertion 0 exserted 1 not exserted 41. Anthers : size 0 all anthers equal 1 lower pair reduced 42. Stamens : length 0 equal 1 lower pair longer 2 upper pair longer 43. Anthers : locule number 0 unilocular 1 biloc"lar-fused 2 bilocular-free Table 4 ( continued ). 44. Filaments : hairs 0 glabrous 1 hairy towards base 2 hairy towards middle 45. Filaments : base 0 not bulbous 1 bulbous 46. Style : type 0 gynobasic 1 terminal 47. Style : lobes 0 plain 1 clavate 48. Style : basal shape 0 not bulbous 1 bulbous 49. Disc : number of tumescent glands 0 none 1 one 2 four 50. Nutlets : hairs 0 absent 1 present 51. Nutlets : surface 0 not verrucose 1 verrucose Table ^ ( continued )• 52. Nutlets : apex 0 not rostrate 1 rostrate 53- Nutlets : number at maturity 0 one 1 four Hairs : septate, eglandular 0 absent 1 present 35- Hairs : septate, glandular 0 absent 1 present 36. Hairs : branched, stalked 0 absent 1 present 57- Hairs : branched, sessile 0 absent 1 present 58. Indumentum : adaxial leaf surface 0 glabrous 1 sparsely hairy 2 densely hairy 59* Indumentum : abaxial leaf surface 0 glabrous 1 sparsely hairy 2 densely hairy Table 4 ( continued ). 60. Indumentum : stem 0 glabrous 1 sparsely hairy 2 densely hairy 61• Inflorescence : number of whorls or spikes 62. Inflorescence : form of verticils 0 not secund 1 sub-secund 2 secund 63 • Inflorescence : number of flowers per whorl 64. Flowers : length of pedicel in millimetres 3.3.2. Coding of character-states. When selecting the character-states care must be taken to ensure that those states allow accurate expression of the data one wishes to record. Thus the two characters ••••••• a) septate hairs absent 0 present 1 b) hairs eglandular 0 glandular 1 are inadequate for coding data from a plant which possesses only septate hairs, but whose hairs may be both glandular« and eglandular, since both states of the second character apply. If the characters are given as a) septate hairs absent 0 present 1 b) glandular hairs absent 0 present 1 the difficulty over the second character is overcome and the plant is coded 1 for each character. If however, the characters are given as a) septate eglandular hairs absent 0 present 1 b) septate glandular hairs absent 0 present 1 then the code for each character remains 1, but the information content is doubled and the data are much more valuable. Inadequate or imprecise expression of character states may result in inadvertant weighting of a character,either by repetition or by misapplication of the instruction for computing similarity. A major problem is the coding of quantitative characters. No satisfactory method has yet been devised for removing size as a factor in analyses. Quantitative characters may be taken as absolute measurements, or converted to ratios or some other suitable form. Some conversion is usually necessary if only to ensure that measurements of different characters are read on similar scales. 3*3«3« Similarities : match and mis-match. The computer must be instructed on how the contribution of each character to the overall similarity is to be calculated. This is done on the basis of match and mis-match and is calculated in one of the four ways explained in table 5. An inappropriate instruction can materially affect the contribution of a character. For example if the character corolla white 0 yellow 1 is read as a type one, then two OTU's with white corollas will be treated as being dissimilar, since 0-0 is a mis-match and contributes nothing to the overall similarity. The appropriate instruction here would be to read the character as a type two. If the character is expressed by the states corolla not white 0 white 1 then it must be read as a type one or OTU's with purple corollas and OTU's with yellow corollas will be treated as similar. These instructions to the computer should be 46 Table 5• Designation of character types according to Gower's coefficient of similarity. The overall similarity between each pair of OTU's is the average of the similarity calculated on each character separately. The characters may be treated in different ways as follows ( means the value of the Kth character for the ith value ) Qualitative ( type 2 ). Xik = Xjk similarity = 100$ Xik ^ Xjk similarity = 0$ Qualitative ( type 1 ). As type 2 but ignore the Kth character when X., = X., = 0 lk jk Quantitative ( type 3 )• similarity = 1 - (x x } 100# ik - .ik range ( K ) i.e. one, minus the difference between the two values divided by the range of the values that the Kth character takes in the data set, expressed as a percentage. Quantitative ( type 4 ). As type 3 but ignore the Kth character when X., = X =0 lk jMk Comparisons with missing values are ignored. 47 considered when deciding on the character states and their codings. 3.3*^« Assessment of results. This is generally treated as a cerebral process. For example in clustering techniques the main problem is delimitation of the clusters. To achieve this one must first decide what constitutes a cluster. In single-linkage cluster analysis a similar problem is seen in the use of phenon-?lines. These are lines drawn across the dendrogram at arbitrary levels of similarity to denote taxonomic levels. How should the levels be chosen ? Again Everitt (1977 ) explains it well :" ... there is no universal agreement on what constitutes a cluster; in fact it has even been suggested ( Bonner 1964 ) that the ultimate criterion for evaluating the meaning of such terms as a cluster is the value judgement of the user. If using the terms produces an answer of value to the investigator, that is all that is required. " 3«3«5« Missing values. Perhaps the greatest single cause of distortion in numerical analyses is missing values in the initial data matrix. In those methods which are able to cope with an incomplete data matrix the missing values are commonly dealt with in one of two ways. A missing value can be replaced with an average figure calculated from the data for the other OTU's or by some other arbitrary figure. This is obviously unsatisfactory and in any case can only be provided for quantitative characters. This is particularly limiting in such methods as principal components analysis where the characters form the axes in multidimensional space on which the OTU's are located as points. 48 The second method is to ignore the character whose value is missing and calculate the overall similarity from a reduced number of characters for that OTU only. If many values are missing the similarity must be regarded with suspicion, but for few missing values the result is a small information loss but little or no distortion. This approach is used in methods such as principal co-ordinates analysis. 3.3.6. Sampling. l£he need for adequate sampling is well known. Inadequate samples may provide atypical data or fail to portray the full range of variation of a character between the OTU's. In morphological studies it is not always possible to obtain a large sample and frequently only a single specimen is available. In addition, and equally important, a restricted sample may not provide a full data set when ( say ) flowering and fruiting characters are required. 3.3.7. Dependant characters. Missing values also occur when dependant characters are used. The character ••••••• number of leaves per whorl is dependant on the character ••••••• leaves alternate 0 whorled 1 An alternate leaved plant will have a missing value for the character concerning the number of leaves. 3.4. THE ANALYSES. There has been such a proliferation of methodology in numerical taxonomy that choosing suitable analyses is not easy. The investigator must be aware of the strengths and weaknesses of the various methods. There are particular limitations to, and distortions produced by, each method of analysis. My data was analysed using the CLASP program. This is a package which produces a similarity matrix, a nearest neighbours table, single1-linkage analysis, principal co-ordinates analysis and clustering to maximise within-group mean similarity analysis. 3.4.1. Similarity matrices. The coded data were used to construct a similarity matrix using the measure of similarity described by Gower ( 1971 ) ( see table 5 ). A similarity matrix gives the percentage similarity of each OTU with every other OTU. For a large matrix the information contained in the mass of numbers is difficult to assimilate. The range of percentage similarities can be divided into arbitrary classes and a different symbol assigned to each class. By replacing the percentage similarities with the appropriate symbols, a simplified similarity matrix can be produced. This allows greater facility in handling the data and makes grouping of similar OTU's within the matrix more easily visible. 3.4.2. Nearest neighbours. For each OTU a list is produced of those OTU's with the highest similarity to it i.e. the nearest neighbours for each OTU. Such a simple treatment of the data can be a useful tool, since checking the position of an OTU relative to a number of 50 others in the full similarity matrix is tedious work. Use of the nearest neighbours list eased the checking considerably. 3.4.3. Single-linkage cluster analysis. This is an agglomerative, non-overlapping clustering method in which an OTU has a similarity to an existing cluster equal to its similarity to the closest member of that cluster. The links between OTU's and clusters and between two clusters are the single- links between pairs of OTU's ( see Sneath and Sokal 1975 for full explanation ) . All agglomerative non-overlapping cluster analyses are based on similarities of individuals and these similarities can be in error for reasons outlined above. In addition, because of the one-to-one linkage of OTU's there is a tendency to add individuals to existing clusters instead of initiating new clusters. Often the result is a phenomenon well known in single-linkage cluster analysis, the production of long, straggly clusters or " chaining ". This can result in loss of definition in the overall picture of the data, especially for distant relationships or where intermediate OTU's are present. However, single-linkage cluster analysis has a number of advantages in a study such as this one. It is a simple and easily-generated analysis readily available in a number of programs. It gives a good representation of close relationships and as with any clustering method, produces discrete groups. The results can be portrayed visually in the form of a dendrogram and any distortion or misplacement of OTU's is easily checked using a nearest neighbours list. Dendrograms represent two-dimensional heirarchical displays 51 of similarity data and are not themselves very practical classifications. They can however be converted into name hierarchy classifications by two principal rank ordering procedures called box-in-box and phenon-line methods. Box-in-box classifications ( see Heywood 1973 ) are useful because they are sequential, each branching point in the dendrogram representing a hierarchy but severe problems arise in taxa where the same rank is applied to different hierarchy levels. In addition the rank at which the hierarchy begins ( or ends ) is arbitrarily selected by the investigator. In a complex dendrogram the many levels of branching may produce an equally complex and unwieldy classification ( see pp. 126 - 129, fig. 26 and table 16 ). Classifications produced by the use of phenon-lines may also be overly complex if all the levels of branching are shown and usually only a small number of phenon-lines at selected levels are used as arbiters of rank. Although this use of few phenon-lines involves oversimplification of some parts of the dendrogram with a comensurate loss of information McNeil ( 1979 ) has recently pointed out the practical necessity of simplification when converting complex dendrograms into name classifications. The main fault with such a method is the arbitrary selection of the similarity levels and, to some extent, the ranks assigned to them. The whole procedure is subjective and reflects the investigator's a priori concepts of the taxa. 3*4.4. Principal co-ordinates analysis. This method utilises eigenvalues and eigenvectors of a matrix of distances between units, derived from the similarity matrix. It 52 provides co-ordinates for each OTU as a point in multi-dimensional space. The number of dimensions is determined by the number of OTU's minus one, or by the number of characters, whichever is the lowest. In this case there were sixty-four dimensions i.e. the number of characters used. The distance between OTU's is inversely related to the similarities between those OTU's. This method places the axis corresponding to the largest eigenvalue in the direction of maximum variance of the cloud of points representing OTU's, the axis corresponding to the next largest eigenvalue in the direction of maximum possible variance at right angles to the first, and so on. It is convenient to represent the data referred to these axes as a series of two-dimensional plots. The size of eigenvalue gives an indication of how much variation is accounted for. Usually only the first three or four eigenvalues are important, subsequent values accounting for very little of the total variation. This consideration of only the largest eigenvalues brings an immediate problem such as representation of three or four dimensions not being adequate, apparent clusters being split if further dimensions are considered. OTU's which are apparently close neighbours in any one projection may in reality be quite distant when all other projections are considered. The delimination of clusters in ordination diagrams is usually done " by eye " - a very subjective method. The formation of unusual or oddly shaped clusters invites interpretation according to individual whim. Despite these disadvantages, principal co-ordinates analysis is a very useful method for taxonomic study and has a number of advantages over other ordination tecniques. Principal 53 co-ordinates analysis was devised by Gower ( 1967 ) as a method free from the distortions found in other ordination techniques. Since it begins with a similarity matrix it is able to cope with missing values and multistate characters, unlike principal components analysis for example which is unable to cope with either ( see 3.3.5. missing values,pp. 47-48) By using a suitable coefficient of similarity in the production of the similarity matrix it can also use qualitative and quantitative characters. This makes principal co-ordinates analysis a very flexible method for ordination, well suited to taxonomic investigations. In addition the ordination diagrams are good visual displays and are easily studied. Although ordination techniques can be unreliable in the representation of close relationships ( due to consideration of only the largest eigenvalues ) they are extremely useful in presenting an overview of relationships. 3.4.5. Clustering to maximise within-group mean similarity ( WGMS ). In this method, all OTU's are randomly assigned to a number of groups. An OTU is then transferred from one group to another if by / doing so the WGMS is increased ( and conversely the between-group mean similarity ( BGMS ) is decreased ). This process is continued until further transfers no longer increase the WGMS. The selection of the number of groups to be used is largely a matter of personal choice although the number of groups seen in the similarity matrix may be used as an indicator. The random assignment of OTU's to the groups can be used to provide an objective check of the groups subjectively seen in the similarity matrix. Since the number of groups is preselected, and every OTU must be assigned to a group, one may encounter the " rag-bag " effect in which a number of unrelated OTU's are clustered to give a group with a very low WGMS. Repetition of the program does not necessarily place all individual OTU's in the same groups as previous runs, although the overall composition of the groups will probably remain unchanged. Thus;the technique does not always give good representations of close relationships but is useful for the examination of more distant or general relationships. 3.4.6. Second runs of analyses. The position of apparently neighbouring OTU's in the principal co-ordinates analysis can be checked by providing further two- dimensional plots using different eigenvalues. If a subset of OTU's from the original program is used in a second run of the analysis the largest eigenvalues will differ from those of the first run. This provides another view of the OTU's and may separate previously neighbouring OTU's. Similarly new data concerning the groupings in the clustering to maximise WGMS analysis may be found by a second run using a subset of OTU's. The OTU's were divided into two subsets, species of Pogostemon and Dysophylla ( subset B ), and all other OTU's except species of Rostrinucula and Colebrookea ( subset A ), this being the main division shown by the analyses of run 1 ( see pp. 107 - 108 ). The programs for the simplified similarity matrix, principal co-ordinates and clustering to maximise WGMS analyses were then re-run for these subsets. 55 3.5- SUMMARY. The number of taxa and characters involved in this study- provided a very large data set ( sixty-four characters for each of 138 OTU's ). Numerical methods were employed to facilitate the handling of the data and the production of a classification. The analyses I have used are diverse but complementary methods of analysis. Principal co-ordinates analysis and the clustering to maximise WGMS analysis are useful for indicating any clusters of OTU's which may be present : they provide an overview of the OTU's. Single-linkage cluster analysis and, in its limited way, the nearest neighbours table provide OTU : OTU comparisons. 56 CHARACTER VARIATION. The Labiatae are usually considered to be a " natural " group in a Linnean sense. Most taxa are very similar to each other in general appearance and morphological variation tends to occur in relatively few characters. Some of these characters have long been considered of great taxonomic importance within the Pogostemoneae; particularly calyx and corolla shape, arrangement of stamens and anther-locule number feature prominently in the works of earlier authors. A survey of sixty four characters ( see table 4 ) and descriptions of variation within those which appear to be of taxonomic significance in the Pogostemoneae is given below. 4.1. LEAVES. All taxa have leaves which are dorso-ventrally flattened to form a distinct lamina. Three basic lamina shapes are found within the group; linear, ovate to orbicular and lobed. The most common is the ovate to orbicular type, where seven of the ten genera ( Colebrookea, fig. 3 , Comanthosphace, Eurysolen, fig. 32 , Keiskea, Leucosceptrum, Rostrinucula, fig. 29 , and Tetradenia ) only have this type. Most species of Elsholtzia ( e.g. E. ciliata ) some species of Pogostemon ( e.g. P. mollis ) and Dysophylla trinervis ( all shown in fig. 3 ) also have leaves of this shape. The distinction between an ovate and an orbicular leaf is a fine one and the two characters tend to grade into each other. Linear leaves are confined to the genus Dysophylla C see fig. 3 ) and to Pogostemon nilagiricus and Elsholtzia pygmaea, although the latter is somewhat dubious since 57 the observation is based on one specimen the leaves of which may- better be considered as narrowly ovate. Three-lobed leaves occur only in Elsholtzia integrifolia. Variation in the shape of the leaf apex is limited and in most species the apex is acute or acuminate, although obtuse apices are found in some species of Dysophylla ( e.g. D. trinervis. fig. 3 ) Elsholtzia ( e.g. E. katchinensis, fig. 3), Pogostemon ( e.g. P. mollis, fig. 3 ) and Tetradenia ( e.g. T. fruticosa ). The shape of the leaf base can be divided into four character states; attenuate, cuneate, rounded and truncate. Species of keiskea, Comanthosphace, Colebrookea ( cf. fig. 3 ) and Rostrinucula ( cf. fig. 29) are exclusively cuneate, Tetradenia species all have rounded leaf bases and Leucosceptrum canum and Eurysolen gracilis ( fig.32 ) have attenuate leaf bases. Species of Elsholtzia and Pogostemon usually have cuneate ( e.g. E. heterophylla, fig. P. glaber, fig. 3 ) or attenuate ( e.g. E. stachyodea, fig. 3 ) leaf bases, although a few species have rounded leaf bases ( e.g. E. katchinensis, fig. 3 , P. speciosus, fig. 3 ). Truncate leaf bases are confined to species of Dysophylla ( e.g. D. stellata, fig. 3 ) although D. trinervis ( fig. 3 ) and D. koehneana have cuneate leaf bases and in D. quadrifolia ( fig. 3 ) the leaf base is usually attenuate. The leaf margins usually show varying degrees of incision, which may be conveniently scored as entire, dentate, or doubly dentate. Species of Keiskea, Comanthosphace, Colebrookea ( cf. fig. 3 ), Rostrinucula ( cf. fig.29 ), Leucosceptrum canum and Eurysolen gracilis ( fig. 32) all have dentate margins. In species of Pogostemon and 58 Tetradenia the margin may be dentate ( e.g. Pogostemon mollis, fig. 3, Tetradenia fruticosa ) or doubly dentate ( e.g. Pogostemon paniculatus, fig. 3, Tetradenia goudotii ). In Dysophylla the margins may be entire ( e.g. Dysophylla tomentosa, fig. 4 ) or dentate ( e.g. Dysophylla stellata, fig. 3 )• Species of Elsholtzia exhibit the whole range from entire ( e.g. Elsholtzia integrifolia ) to dentate ( e.g. Elsholtzia densa, figs 3, 31 ) and doubly dentate ( e.g. Elsholtzia ciliata, fig. 3 ). 4.2. HETEROPHYLLY. Heterophylly is rare but is found in two species of Elsholtzia ( E. heterophylla, fig. 4, and E. bodinieri ). Unlike most taxa these two species possess leaves on the stolons which are much smaller, usually broader and much hairier than the cauline leaves. 4.3. LEAP SIZE. Members of the Labiatae normally have one pair of leaves at each node of the stem; each leaf of a pair is identical to the other. Pogostemon paniculatus ( fig. 4 ) and P. gardneri are unusual in having one leaf of a pair much smaller than the other. This feature is occasionally found in P. purpurascens but is absent from the remainder of the group. 4.4. PHYLLOTAXIS. The Pogostemoneae ( excluding Dysophylla section Verticillatae Benth. ) usually follow the general L^biatae pattern of opposite, decussate leaf pairs. Colebrookea ternifolia is supposed by some authors to have ternate leaves but I have never found this. Species of 59 Fig. 3 • Leaf shapes, margins and petioles in the Pogostemoneae. a. Pogostemon mollis x 1 b. Dysophylla trinervis x 2 c. Elsholtzia katchinensis x 1 d. Pogostemon glaber x 1 e. Pogostemon speciosus x 1 f. Pogostemon paniculatus x 1 g- Elsholtzia stachyodea x 1 h. Colebrookea oppositifolia x 0.5 i. Elsholtzia ciliata x 1 j- Elsholtzia densa x 1 k. Dysophylla stellata x 1 1. Dysophylla quadrifolia x 1 60 Fig. 3 . Phyllotaxis in the Pogostemoneae. Pogostemon paniculatus x 0.5 Leaves in opposite pairs. Note the disparity in size of the members of the pairs of leaves. Dysophylla tomentosa x 1 Leaves in whorls of six. Elsholtzia densa x 1 Leaves in opposite pairs. Dysophylla linearis x 1 Leaves in whorls of four. Elsholtzia heterophylla x 1 e. Leaves in opposite pairs. f. Stolons bear smaller, more rounded leaves than those of the stems. 62 Fig. k 63 Fig. 5 • Bract types in the Pogostemoneae. a. Eurysolen gracilis x 5 b. Elsholtzia luteola x 5 c. Elsholtzia ciliata x 5 d. Dysophylla auricularia x 10 e. Elsholtzia katchinensis x 5 f. Rostrinucula dependens x 5 g- Elsholtzia flava x 5 h. Elsholtzia densa x 5 i. Pogosteraon paniculatus x 5 j- Tetradenia fruticosa x 10 k. Comanthosphace japonica x 5 1. Elsholtzia pilosa x 5 m. Keiskea elsholtzioides x 5 n. Leucosceptrum canum x 5 65 Dysophylla section Verticillatae Bentll. ( fig. k) are very distinct in having whorls of leaves at each node. The number of leaves per whorl varies between, but is usually constant within, each species. The most common numbers are three, four and five but D. stocksii may have fourteen or more leaves per whorl. if.5. PETIOLES. Petiole length is sometimes difficult to ascertain, especially in species with attenuate leaf bases, but when a distinct petiole is present it varies from 1mm ( e.g. Elsholtzia heterophylla, fig. k ) to 80mm ( e.g. Elsholtzia fruticosa ) long. In most Dysophylla species the leaves are sessile, but D. quadrifolia ( fig. 3 ) and a few other species have short but quite distinct petioles. *f.6. BRACTS. The Pogostemoneae show considerable variation in the bracts, particularly in colour, shape, fusion and persistence. Six combinations of these four characters can occur: Bracts brown, broader than long, free, deciduous: this combination is found in all species of Comanthosphace, Rostrinucula and Leucosceptrum canum ( cf. figs 29 )• The bracts often fall before the flower buds open. Bracts brown, broader than long, free, persistent: this combination is found in species of Tetradenia and Elsholtzia sections Elsholtzia and Aphanochilus series Platyelasmeae ( cf,. figs 31.). In the species of Elsholtzia section Elsholtzia the bracts are membranous. 66 Bracts brown, broader than long, fused, persistent: in species of Elsholtzia section Cyclostegia and E. luteola ( figs 5, 30.) the bracts are again membranous and in addition each pair is connate at the margins, forming a cyathium. Bracts green, broader than long, free persistent: in Elsholtzia kachinensis ( fig? 5 ) and E. concinna the bracts are green and never membranous. Bracts green, ovate, free, persistent: this type includes the following species ( all shown in fig. 5 ), Elsholtzia flava, Eurysolen gracilis, some Pogostemon species ( e.g. P. paniculatus ), some Dysophylla species ( e.g. D. auricularia ) and all species of Keiskea. Bracts green, linear, free, persistent: the remaining species of Pogostemon ( e.g. P. fraternus ) and Dysophylla ( e.g. D. peguana ) and Elsholtzia ( e.g. E. pilosa, fig. 3 ) together with species of Colebrookea all have this character combination. Jf.7. BRACTEOLES. Bracteoles may be present or absent. When present they are usually shorter, narrower and more hairy than the bracts. Bracteoles are absent in species of Keiskea, Tetradenia, Elsholtzia sections Elsholtzia and Cyclostegia and section Aphanochilus series Platyelasmeae In species of Comanthosphace, Rostrinucula and Leucosceptrum canum the bracteoles are caducous and, like the bracts, often fall before the flower buds open. 67 A-.8. INDUMENTUM. , Four hair types ( fig. 6 ) are found, the indumentum being composed of one or more types: Septate eglandular hairs; these are simple hairs each at least several cells long. The cells are laterally compressed and arranged so that the narrow sides alternate along the length of the hair, and the terminal cell is usually somewhat pointed. Septate glandular hairs; these are structurally identical with the septate eglandular hairs except for the terminal cell, which is a globular, single-celled gland. Branched stalked hairs; these are the 11 stellate " hairs of earlier authors, each hair in fact having a central axis bearing irregular branches along its length. The hairs are multicellular, the septa clearly visible. Branched sessile hairs; similar to branched stalked hairs, but distinguished by the lack of a central axis, the branches radiating irregularly from a central point. Most species have septate, eglandular hairs only. A number of species e.g. Pogostemon brachystachyus and Dysophylla linearis have a mixture of glandular and eglandular septate hairs. Branched stalked hairs are less common, being found in five genera. Species of Comanthosphace, Leucosceptrum and Rostrinucula bear branched stalked hairs with a few septate eglandular hairs. The three species of Tetradenia and a number of Elsholtzia species e.g. E. capituligera and E. stachyodea have a mixture of branched stalked and septate eglandular hairs. Other species e.g. Elsholtzia eriostachya and E. fruticosa have branched stalked, septate eglandular and septate glandular hairs. Branched 68 sessile hairs are found in two species of Pogostemon; in P. tuberculosus the hairs have a central boss from which the short stiff branches radiate; in P. velatus there is no* central boss and the branches are long and flexuous. All species possess some hairs, although these may be restricted to the inflorescence. The abaxial leaf surface is always more hairy than the adaxial; the veins more hairy than the surface of the lamina. The density of hairs on the stems decreases with age, so that the lowest parts of the stem are frequently glabrous. The density of the indumentum varies between individuals as well as between species. Only Pogostemon glaber and sbm« species of. Dysophylla e.g. D. yatabeana have glabrous stems and leaves. All other species have some indumentum, ranging from very thin, e.g. Elsholtzia luteola to very dense, e.g. Pogostemon mollis. 9. CALYX. The calyx is generally campanulate, five toothed with the teeth sometimes much elongated. In all species of Keiskea ( cf. fig. 7 )» most Elsholtzia species ( e.g. E. densa, figs 7> 31 ) and E. ciliata, ( fig. 7 ) and some species of Pogostemon ( e.g. P. fraternus, fig. 7 ) the teeth are unequal, in species of Tetradenia ( cf. fig. 7 ) the upper tooth is much the broadest and overlaps the lateral teeth. The calyx tube is strongly veined, the most prominent veins being generally referred to as the main veins. Three species, Elsholtzia integrifolia, Comanthosphace ningpoensis and Elsholtzia chinense have fifteen main veins; species of Comanthosphace 69 ( excluding C. ningpoensis ) and Leucosceptrum canum ( fig. 7 ) have twelve. The remaining taxa have five or ten main veins, although in some species of Dysophylla ( e.g. D. tomentosa ) the number is difficult to determine. In species of Colebrookea ( cf. fig. 7 ), Dysophylla pentagona, D. stocksii and D. griffithii the main, central veins of the teeth are thickened to form five strong ribs. The calyx is usually hairy on the outer surface, and the teeth are fringed with cilia. Three species, Elsholtzia kachinensis, E. penduliflora and Dysophylla glabrata (fig.32 ) have a completely glabrous calyx. Keiskea .japonica ( fig. 7 ) has a hairy calyx tube but glabrous teeth, while several species including Pogostemon amaranthoides and P. paludosus have a glabrous calyx tube and hairy teeth. In species of Colebrookea the teeth are plumose. The presence of an annulus of stiff hairs in the throat of the calyx is confined to species of Keiskea and some species of Pogostemon ( e.g. P. fraternus and P. litigiosus, both in fig. 7 )• The calyx may be accrescent at the fruiting stage. In species of Elsholtzia series Platyelasmeae the increase in,size is striking, the fruiting calyx ( cf. figs 7i 31 ) being up to eight times as large as at anthesis. In species of Colebrookea the calyx teeth are plumose and become greatly elongated but not much widened in fruit. The calyx teeth are normally erect during the fruiting stage but in Dysophylla stellata and Pogostemon litigiosus they are spreading and in some species of Dysophylla ( e.g. D. pentagona ) and Pogostemon ( e.g. P. nelsoni ) the teeth are strongly incurved. Hair types in the Pogostemoneae. septate glandular hair, septate eglandular hair, branched stalked hairs, branched sessile hair, the branches short and stiff, branched sessile hair, the branches long and flexuous. 72 Fig. 7 . Calyx shapes and vernation in the Pogostemoneae. a. Dysophylla trinervis x 5 b. Elsholtzia flava x 5 c. Leucosceptrum canum x 5 d. Pogostemon parviflorus x 5 e. Elsholtzia densa ( fruiting ) x 5 f. Colebrookea oppositifolia ( fruiting ) x 10 g. Tetradenia goudotii x 5 h. Elsholtzia ciliata x 5 i. Pogostemon litigiosus x 5 j. Pogostemon fraternus x 5 k. Keiskea japonica x 5 Only main veins are shown. 73 Fig. 7 • Fig. 8 . Corolla structures in the Pogostemoneae. a. Keiskea japonica x 5 b. Eurysolen gracilis x 5 c. Elsholtzia stauntoni x 5 d. Rostrinucula dependens x 5 e. Tetradenia fruticosa x 5 f. Comanthosphace .iaponica x 5 g- Colebrookea oppositifolia x 20 h. Pogosteraon tuberculosus x 5 i. Leucosceptrum canum x 5 75 Fig. 8 . 76 4.10. COROLLA. The corolla is zygomorphic, although in some species of Dysophylla weakly so and not obviously bilabiate. However, the corolla lobes are generally arranged to form an upper group of three and a single lower lobe. Elsholtzia integrifolia is.clearly exceptional in having a bifid lower lobe. In Pogostemon, Dysophylla and Rostrinucula ( cf. fig.29 ) the upper central lobe is entire; in the remaining six genera it is shallowly to deeply emarginate. The corolla tube shows little variation except in length. It may reach 9*5mm in length ( e.g. Elsholtzia bodinieri ) or be as little as 0.5mm long ( e.g. Tetradenia hildbrandtii ) and is usually rather slender at the base widening gradually to the throat. In species of Pogostemon the upper lip is longer than or equals the lower lip. In all other genera the lower lip is longer than or equals the upper lip. An annulus of hairs in the throat of the corolla is found in six genera. In species of Keiskea, Comanthosphace and Tetradenia ( cf. fig. 8 ) the annulus is a complete ring at, or slightly below the level of insertion of the staminal filaments. In a number of Elsholtzia species e.g. E. stauntoni ( fig. 8 ), E. hunanensis and E. capituligera the annulus is an open ring, the lines of hairs projecting dorsally and ending under the upper lip. All species of Rostrinucula ( cf. figs 8, 29 ) also have a partial annulus composed of clusters of hairs at the insertion point of each staminal filament. In addition to these hair-clusters there is an invagination on the ventral surface of the corolla forming a crescent-shaped papilla in the throat, itself also hairy and forming part of the annulus. Eurysolen gracilis ( figs 8, 32 ) has a corolla which is gibbous 77 slightly above the base. Immediately below the gibbous curve of the corolla is an invagination similar to that in Rostrinucula, forming a hairy papilla which all but closes the corolla tube. Three corolla colours are known: yellow, white and rose-pink to purple, the latter being the most common. 4.11. STAMENS. In common with most Labiatae, the Pogostemoneae have four stamens arranged into two pairs; an upper and a lower pair. These may be equal in length or one pair longer than the other. In species of Colebrookea, Rostrinucula ( cf. fig. 29) and Tetradenia the stamens are equal; in Eurysolen gracilis ( fig.32 ) the upper stamens are longest; in species of Comanthosphace, Keiskea and Leucosceptrum canum the lowest stamens are longer • In the remaining genera all three states can occur. The anthers are almost always exserted, the only exceptions being Elsholtzia aquatica and species of Elsholtzia series Platyelasmeae ( cf. fig. 31 )• The filaments are inserted toi/ards the top of the corolla tube. In Rostrinucula dependens,,R> sinensis and Eurysolen gracilis ( cf. figs 29,32) each filament base is marked by a bulbous swelling below the point of insertion, which bears annular hairs. Species of Dysophylla, Pogostemon and Leucosceptrum ( cf. fig. 9 ) have hairs on the filaments. In the first two genera the hairs are long, thread-like and usually purple due to pigments in the crosswalls of the cells. In Leucosceptrum canum the hairs are short and white. In all species of Dysophylla and most species of Pogostemon the hairs are 78 borne towards the middle of the filament; in Leucosceptrum canum ( fig. 9 ) and some species of Pogostemon, ( e.g. P. travancoricus, fig. 9 , and P. hispidus ) the hairs are borne towards the base of the filament. The anthers are usually equal in size. However Elsholtzia pilosa has two smaller, lower anthers often reduced to half the size of the upper ones. The anthers are bilocular in species of Keiskea, Elsholtzia (figs 9j30,31) and Tetradenia and unilocular in the remaining genera. Species of Tetradenia and most species of Elsholtzia have locules confluent through partial fusion. Species of Keiskea, Elsholtzia hunanensis and E. chinense have distinctive free locules. \ 4.12. STYLE. Two types of style occur in the Labiatae; gynobasic, when the style arises from the base of,and between,the deeply-divided lobes of the ovary and terminal, when the ovary is shallowly-lobed and the style is not basal. Gynobasic styles are found throughout the family except for the Ajugoideae which have terminal styles. In Leucosceptrum canum ( fig. 10) the style usually has the appearance of being terminal but in some specimens it more nearly approaches the gynobasic condition while the reverse seems true of Elsholtzia flava.( fig. 10 ). All others .in the Pogostemoneae have gynobasic styles. The style is typically long, slender and straight or slightly curved, with a bifid tip. The lobes at the tip are subulate and equal. In species of Elsholtzia the style may vary in the shape of the base 79 and of the lobes. In some species ( e.g. E. stachyodea, fig. 9 , E. ochroleuca ) the style bears a bulbous swelling at the base, just above the point of emergence from between the nutlets. In species of Elsholtzia series Platyelasmeae ( cf. fig. 9 ) the style lobes are clavate at the tips. This is also found in Pogostemon litigiosus but in no other species. 4.13. DISC. The disc may be regular or may bear one to several tumescent lobes. In all species of Keiskea and Elsholtzia (figs 9,30,31) there is a single, somewhat elongated lobe on the posterior edge of the disc. In all Tetradenia species ( cf. fig. 9 ) there are four lobes, spaced regularly around the disc. They are bright-red and quite large, overtopping the young nutlets. 4.14. NUTLETS. Nutlet characters can be difficult to verify as ripe fruits are not always available. However variation has been observed with respect to number, shape, hairiness and ornamentation. In the Labiatae as a whole there are usually four nutlets, a feature also true of the Pogostemoneae. Exceptions to this general rule are Keiskea japonica ( fig. 11), Elsholtzia kachinensis, Dysophylla stocksii and Colebrookea oppositifolia ( fig. 11). In these species there is only one nutlet at maturity, the other three apparently undergoing early abortion. In Shape the nutlets: are generally obovoid and blunt at the apex. However Rostrinucula species (figs 11, 29 ) have nutlets equipped 80 with a rostrate tip, very clearly visible at maturity. Hairy nutlets are found in all species of Comanthosphace except C. nanchuanensis, Rostrinucula and Colebrookea, and in Elsholtzia chinense where the hairs tend to be confined to the apex of the nutlet. All other species have glabrous fruits. Ornamentation of the nutlets is limited. Most species have nutlets with a smooth to slightly rugulose surface. By contrast those of Elsholtzia series Platyelasmeae ( figs 11, 31 ) are distinctly verrucose towards the apex. In Keiskea japonica the nutlet has ridges which form an almost reticulate pattern. In species of Colebrookea the single ripe nutlet does not separate from the calyx, the two structures acting as a single dispersal unit. 4.15. INFLORESCENCE. The inflorescence is usually racemose ( although it is paniculate in some species of Pogostemon ) and varies greatly in length and in the number of whorls of flowers, or verticels. The verticels are secund in all species of Keiskea and many species of Elsholtzia ( e.g. E. stauntoni, E. ciliata and E. luteola, fig. 30 ). In the paniculately branched species of Pogostemon the verticels are sub- secund, i.e. the flowers are secundly arranged in each verticel but the verticels themselves are not secund on the stem. The flowers in all genera may be pedicellate or sessile. 9 • Stamen types in the Pogostemoneae. a. Keiskea .iaponica b. Colebrookea oppositifolia c. Comanthosphace sublanceolata d. Elsholtzia ciliata e. Leucosceptrum canum f. Dysophylla linearis Pogostemon parviflorus h. Pogostemon atropurpureus All stamens x 10 All anther details x 20 83 Fig. 10 . Style and disc types in the Pogostemoneae. a. Leucosceptrum canum b. Elsholtzia densa c. Elsholtzia stachyodea d. Elsholtzia fruticosa e. Comanthosphace .japonica f. Pogostemon tuberculosus R- Keiskea .iaponica h. Tetradenia fruticosa All x 10 8k 11 • Nutlet types in the Pogostemoneae. a. Keiskea japonica b. Colebrookea oppositifolia c. Eurysolen gracilis d. Dysophylla stellata e. Pogostemon mollis f. Leucosceptrum canum g- Comanthosphace barbinervis h. Elsholtzia flava i. Elsholtzia densa 3- Rostrinucula dependens Profile and inner face of nutlet are shown. All x 10 86 Fig. . 87 5. RESULTS OF THE ANALYSES. 5.1. THE SIMILARITY MATRICES. The full similarity matrix from the coded data in appendix 2 is given in appendix 3 • For ease of study a simplified similarity matrix ( appendix 4 ) was produced ( see also P- )• A preliminary examination of this matrix suggests a number of possible groups of species. They are shown in figs 12 and 13. Three groups are quite discrete, formed by species of Keiskea, species of Tetradenia and species of Colebrookea respectively. Species of Comanthosphace, Rostrinucula and Leucosceptrum canum form a fourth distinct group, although L. canum ( OTU 7 ) and Comanthosphace nanchuanensis ( OTU 13^ ) are only weakly linked with the other species and the two species of Rostrinucula form a subgroup within the main group. The two groups of species of Elsholtzia are formed roughly by section Aphanochilus series Stenelasmeae and sections Elsholtzia and Cyclostegia. Sections Elsholtzia and Cyclostegia form a close-knit group, each species sharing high similarities with the other members of the group. Section Aphanochilus series Stenelasmeae is more diffuse, with some species e.g. Elsholtzia beddomei ( OTU 129 )» E. integrifolia ( OTU 23 ) and E. penduliflora ( OTU 25 ) sharing very few high similarities with the other group members. However there is considerable overlap between the two groups, patricularly between species of the E. communis and E. ciliata complexes ( OTU's 31» 32, 33» 3k and 13» 15 respectively ). It is interesting to note here the position of E. chinense ( OTU 100 ), which shares a roughly equal number Pogostemon Elsholtzia section Aphanochilus series Stenelasmeae Elsholtzia sections Elsholtzia and Cyclostegia Keiskea y^f 89 Fig. 13 . Diagram showing six possible groups indicated in the simplified similarity matrix. 90 of high similarities with species of Elsholtzia section Aphanochilus series Stenelasmeae and species of Pogostemon. The three species of Elsholtzia section Aphanochilus series Platyelasmeae form a reasonably discrete group. Excluding other members of the group E. eriostachya ( OTU 20 ) seems most closely linked with species of section Aphanochilus series Stenelasmeae while E. densa ( OTU 21 ) and E. manshurica ( OTTJ 125 ), although sharing high similarities with members of both groups, seem more closely linked with species of sections Elsholtzia and Cyclostegia. The groups formed by species of Pogostemon and Dysophylla are the most difficult to define clearly. Two groups are formed by species of Pogostemon and a third group by species of Dysophylla. However the members of each group share so many high similarities with members of the other two groups that they can as easily be considered as one, large, loose group. A number of OTU's do not fit easily into any of the groups outlined above e.g. Dysophylla glabrata ( OTU ), Pogostemon litigiosus ( OTU 21 ) and most notably Eurysolen gracilis ( OTU 126 ) which is not similar to any other OTU. Three other OTU's are outstanding in their positions, Dysophylla mairei ( OTU 102 ) and Leucosceptrum plectranthoideum ( OTU 97 ) are grouped with species of Elsholtzia section Aphanochilus series Stenelasmeae and Elsholtzia aquatica ( OTU 22 ) is grouped with species of Dysophylla. The matrices produced in runs 2A and 2B show very similar results ( appendices 5 and 6 respectively ). In run 2A, Elsholtzia bodinieri ( OTU 17 ) and E. heterophylla ( OTU 18 ) form a subgroup within E. section Elsholtzia but neither of the other members of section Cyclostegia ( i.e. E. luteola, OTU 16 and E. strobilifera, OTU 19 ) show 91 any indications of being separate from section Elsholtzia, A more detailed picture of the Dysophylla and Pogostemon groups is given by run 2B. Species of Dysophylla section Verticillatae form a well-defined block in the matrix but species of Pogostemon form a much looser group. Species of Dysophylla section Oppositifoliae ( OTU's 571 58, 59, 60.and OTU 104 ) form a transition group. D. salicifolia, D. rugosa, D. myosuroides and D. andersoni ( OTU's 58, 59? 60 and 104 ) have strong and obvious links with the Dysophylla block but also share high similarities with a large selection of Pogostemon species. To a lesser extent the same applies to Dysophylla auricularia ( OTU 57 ) and D. glabrata ( OTU kk ) which share fewer high similarities with other species. Only two subgroups within the Pogostemon species can be defined with any clarity, the benghalensis / paniculatus complex ( sixteen species ) and the fraternus complex plus P. at r opur pur en s and P. speciosus ( six species )• Even these however show considerable overlap with species outside the groups and are not sufficiently clear to allow confident recognition. There are several other minor groupings but all so diffuse as to defy adequate definition. Consideration of these various groups of species within the similarity matrix suggests two alternative schemes, recognising either nine groups or six groups. Simplified diagrams of the similarity matrix for each scheme are given in figs12 and 13- In the nine group scheme, the groups are :- Keiskea, Elsholtzia sections Elsholtzia and Cyclostegia, Elsholtzia section Aphanochilus,Pogostemon.. , pro majore, Dysophylla, Pogostemon pro minore, Comanthosphace / Leucosceptrum canum / Rostrinucula, Tetradenia and Colebrookea. In the 92 six group scheme the groups are Keiskea, Elsholtzia, Pogostemon / Dysophylla, Comanthosphace / Leucosceptrum canum / Rostrinucula, Tetradenia and Colebrookea. 5.2. CLUSTERING TO MAXIMISE WITHIN-^GROUP MEAN SIMILARITY ( WGMS ). Some attempt can be made to find which of the two schemes from the similarity matrices ( pp 91-92), i.e. nine groups or six groups, best fit the data by using a method which maximises the WGMS for the groups. The WGMS and BGMS values for the six group scheme and the nine group scheme are given in tables 6 and 7 respectively. In each case some anomalies are to be expected since not all species in the similarity matrix fit easily into any of the main groups. Table 6 shows that allocation of species among six groups does not produce clear divisions. Group k comprising species of Comanthosphace, Leucosceptrum canum and Rostrinucula has a relatively low WGMS of 85.8/6 but also rather depressed values for the BGMS ranging from 67«7% to 72.1$. The species of Pogostemon, Dysophylla and Colebrookea in groups 1, 2, 5 and 6 are obviously more similar to each other than to either of the remaining groups ( groups 3 and k ). The relatively high BGMS values compared to the WGMS values suggest that none of the six groups is particularly discrete or homogenous. What is clear from the results is that with the exception of group k the groups formed hardly match those expected from examination of the similarity matrix. In particular species of Pogostemon are divided among three groups ( not two ) and species of Elsholtzia, Keiskea and Tetradenia are placed together in a single group ( not three separate groups ). The nine group analysis ( table 7 ) gives a result much closer to 93 Table 11 • WGMS and BGMS values for four groups in the clustering to maximise WGMS analysisrun 2A . 1 84.8 2 82.4 88.9 3 73-7 72.0 79.7 4 71.6 70.4 72.1 85.8 5 82.3 81.4 70.9 71.2 87.9 6 80.8 78.8 70.1 67.7 76.4 88.3 1 2 3 4 5 6 Key to groups 1 - species of Colebrookea, Pogostemon and Dysophylla section Opposit ifoliae 2 - species of Pogostemon 3 - species of Keiskea, Tetradenia, Elsholtzia and Eurysolen 4 - species of Comanthosphace, Rostrinucula and Leucosceptrum canum 5 - species of Pogostemon 6 - species of Dysophylla section Verticillatae ( for exact distribution of OTU's see appendix 8 ) 94 Table 11 • WGMS and BGMS values for four groups in the clustering to maximise WGMS analysisrun 2A . 85.8 67.9 90.8 70.3 70.2 94.7 67.9 64.2 65.8 88.8 71.8 69.3 70.4 80.0 85.8 72.6 74.8 74.0 71.8 74.3 83.1 70.7 70.0 68.0 78.9 83.0 72.2 89.6 67.2 65.8 66.7 75.4 73.8 71.3 76.5 79.3 72.8 77.3 76.4 70.8 72.2 78.3 73.0 68.5 86.7 Key to groups 1 - species of Comanthosphace, Rostrinucula and Leucosceptrum canum 2 - species of Keiskea 3 - species of Tetradenia 4 - species of Dysophylla section Verticillatae 5 - species of Pogostemon and Dysophylla section Oppositifoliae 6 - species of Elsholtzia section Aphanochilus and Eurysolen 7 - species of Pogostemon 8 - species of Colebrookea, Dysophylla glabrata and Pogostemon • amarantho ide s 9 - species of Elsholtzia sections Elsholtzia and Cyclostegia ( for exact distribution of OTU's see appendix 8 ) 95 Table 11 • WGMS and BGMS values for four groups in the clustering to maximise WGMS analysisrun 2A . 1 88.5 2 69-5 90.8 3 71.3 70.2 94.7 4 68.1 64.2 65.8 88.8 5 68.9 69.3 70.4 80.0 85.8 6 74.0 74.8 74.0 71.8 74.3 83.1 7 67.4 70.0 68.0 78.9 83.0 72.2 89.6 8 68.5 65.4 70.3 65.4 71.6 72.2 69.4 9 73.9 77.3 76.4 70.8 72.2 78.3 73.0 10 80.5 61.5 70.3 65.7 65.8 72.2 72.3 .8 10 Key to groups 1 - species of Comanthosphace and Leucosceptrum canum 2 - species of Keiskea 3 - species of Tetradenia 4 - species of Dysophylla section Verticillatae 5 - species of Pogostemon and Dysophylla section Oppositifoliae 6 - species of Elsholtzia section Aphanochilus and Eurysolen 7 - species of Pogostemon 8 - species of Colebrookea 9 - species of Elsholtzia sections Elsholtzia and Cyclostegia 10 - species of Rostrinucula 96 the expected one, only two features differing from the similarity matrix ( appendix ^ ). In group 8 Pogostemon amaranthoides ( OTU 69 ) and Dysophylla glabrata ( OTU 44 ) are included with the species of Colebrookea ( see appendix 8 ). In group 9 Elsholtzia sections Elsholtzia and Cyclostegia now include two of the three species which comprise Elsholtzia section Aphanochilus series Platyelasmeg.e, i.e., E. densa ( OTU 21 ) and E. manshurica ( OTU 125 )• The third species, E. eriostachya ( OTU 20 ) remains in Elsholtzia section Aphanochilus series Stenelasmeae ( see appendix 8 )• The increase in the number of groups is attained by joining groups 1 and 5 of the six group schemes and splitting group 3 into four new groups. Apart from this and the movement of OTU's 44, 69, 21 and 125 noted above, the only other change is the movement of Dysophylla andersoni ( OTU 104 ) from a Pogostemon group in the six group scheme to the Dysophylla group in the nine group scheme. In every other respect the composition of the groups is remarkably stable ( see appendix 8 )« The table of similarities for the nine group version ( table 7 ) shows groups 2 and 3? species of Keiskea and Tetradenia respectively, to have high WGMS values and low BGMS values and both are noticeably more similar to groups 6 and 9i sections of Elsholtzia, than to the other groups. Group 1, species of Comanthosphace, Rostrinucula and Leucosceptrum canum, has a relatively low WGMS of 85.8$ but the BGMS values are also rather low and, as in the six group scheme, this group seems quite acceptable. Group 4, species of Dysophylla, and groups 5 and 7> species of Pogostemon, share low BGMS values with the remaining groups but high values with each other. The WGMS of 85.8% for group 5 is little higher than the BGMS of 83% and 80% shared with 97 groups 4 and 7- Although the WGMS of groups 4 and 7 are higher, neither these nor group 5 are convincingly separable from each other. Groups 6 and 9 also have relatively low WGMS values. Group 6, species of Elsholtzia section Aphanochilus series Stenelasmeae, shares a relatively high BGMS with group 9, species of Elsholtzia sections Elsholtzia and Cyclostegia ,but lower values with other groups. However group 9 shares relatively high BGMS values with groups 2 and 3 as well as with group 6. Group 8 is an anomal^y, containing the unlikely combination of the two species of Colebrookea with one species each of Pogostemon and Dysophylla. The heterogeneity of this combination is reflected in the very low WGMS value of 79-3$ and the obvious similarity with species of Pogostemon in groups 5 and 7 and the species of Dysophylla in group 4. Slight rearrangement of the groups to remove the anomal y in group 8 results in the similarities shown in table 8 . In this table the species of Pogostemon and Dysophylla in group 8 have been removed, Pogostemon amaranthoides ( OTU 69 ) now being placed in group 7 and Dysophylla glabrata ( OTU 44 ) in group 4. In addition the two species of Rostrinucula ( OTU's 95 and 96 ), which are rather peripherally linked with the species of Comanthosphace in group 1, have been removed and now form a tenth group. The addition of Pogostemon amaranthoides to group 7 and Dysophylla glabrata to group 4 produces only small changes in the similarity values for these groups. However the effect of their removal on group 8 is striking. The WGMS rises to 95-2% and the links with groups 4, 5 and 7 disappear, the BGMS values falling to levels comparable with the 98 other groups. The effect of removing the species of Rostrinucula from group 1 is less marked. The WGMS for group 1 rises to 88.5$ and the BGMS values remain almost unchanged. The new group 10 has a WGMS of 94.8$ and is most similar to group 1 with which it shares a BGMS of 80.5%. Its BGMS values with the other groups are low, ranging from 61.5% to 72.3%. Like the species of Rostrinucula, Leucosceptrum canum ( OTU 7 ) and Comanthosphace nanchuanensis ( OTU 134 ) appear to be rather distant members of the Comanthosphace group as shown in the simplified similarity matrix. The WGMS and BGMS values for each of these species with Comanthosphace and Rostrinucula are shown in tables 9 and 10. it is noticeable that Leucosceptrum canum ( table 9 ) is more similar to Comanthosphace pro majore than is Comanthosphace nanchuanensis ( table 10). Leucosceptrum canum is much less similar to Rostrinucula than either Comanthosphace nanchuanensis or Comanthosphace pro ma .j ore. In both the six and nine group schemes Eurysolen gracilis ( OTTJ 126 ), Dysophylla mairei ( OTU 102 ) and Leucosceptrum plectranthoideum ( OTU 97 ) are grouped with species of Elsholtzia section Aphanochilus series Stenelasmeae and Elsholtzia aquatica ( OTU 22 ) is grouped with species of Dysophylla ( see appendix 8 ). Like the simplified similarity matrix of run 1, that of run 2A ( appendix 5 ) suggests alternative sets of groups. The first alternative recognises four groups; species of Keiskea, species of Tetradenia, species of Comanthosphace and Leucosceptrum and species of Elsholtzia. The second alternative recognises the same groups but with the species of Elsholtzia divided into two groups formed by sections 99 Table 9 • WGMS and BGMS values for species of Comanthosphace, Rostrinucula and Leucosceptrum canum. 1 93.7 2 81.6 94.8 3 84.3 72.3 100 1 2 3 Key to groups 1 - species of Comanthosphace 2 - species of Rostrinucula 3 - Leucosceptrum canum Table 10. WGMS and BGMS values for species of Comanthosphace, Rostrinucula and Comanthosphace nanchuanensis. 1 93.7 2 81.6 94.8 3 81.6 82.6 100 1 2 3 Key to groups 1 - species of Comanthosphace 2 - species of Rostrinucula 3 - Comanthosphace nanchuanensis 100 Table 11 • WGMS and BGMS values for four groups in the clustering to maximise WGMS analysis run 2A. 1 84.2 2 72.9 88.5 3 77.5 74.1 83.6 4 76.8 71.9 76.0 81.4 1 2 3 4 Key to groups 1 - species of Keiskea and Elsholtzia sections Elsholtzia and Cyclostegia 2 - species of Comanthosphace and Leucosceptrum canum 3 - species of Elsholtzia section Aphanochilus series Stenelasmeae and Eurysolen 4 - species of Tetradenia and Elsholtzia section Aphanochilus series Platyelasmeae ( for exact distribution of OTU's see appendix 9 ). 101 Table 11 • WGMS and BGMS values for four groups in the clustering to maximise WGMS analysis run 2A. 1 83.6 2 76.0 81.4 3 74.9 71.2 90.8 4 78.4 78.6 77.9 88.1 5 74.1 71.9 69.5 74.1 88.5 1 2 3 4 5 Key to groups 1 - species of Elsholtzia section Aphanochilus series Stenelasmeae and Eurysolen 2 - species of Elsholtzia section Aphanochilus series Platyelasmeae and Tetradenia 3 - species of Keiskea 4 - species of Elsholtzia sections Elsholtzia and Cyclostegia 5 - species of Comanthosphace and Leucosceptrum canum ( for exact distribution of OTU's see appendix 9 ) 102 Table 11 • WGMS and BGMS values for four groups in the clustering to maximise WGMS analysis run 2A. 1 83.6 2 71.1 94.7 3 74.9 70.2 90.8 k 78.4 76.6 77.9 88,1 5 74.1 71.3 69.3 74.1 88.5 6 78.9 74.9 72.3 75.2 72 A 8 7.5 1 2 3 k 5 6 Keys to groups 1 - species of Elsholtzia section Aphanochilus series Stenelasmeae and Eurysolen 2 - species of Tetradenia 3 - species of Keiskea k - species of Elsholtzia sections Elsholtzia and Cyclostegia 5 - species of Comanthosphace and Leucosceptrum canum 6 - species of Elsholtzia section Aphanochilus series Platyelasmeae 103 Elsholtzia and Cyclostegia and section Aphanochilus series Stenelasmeae. In this scheme species of Elsholtzia section Aphanochilus series Platyelasmeae would be rather difficult to place as they have similarities with members of both groups of Elsholtzia species. A third alternative is to segregate the species of series Platyelasmeae as a sixth group. The composition of the groups and the WGMS and BGMS values for the four and five group schemes are given in appendix 9 and tables 11 and 12 . It is apparent from these results that neither scheme is satisfactory. In the four group scheme ( table 11) only group 2 ( species of Comanthosphace and Leucosceptrum canum ) has a sufficiently high WGMS to merit recognition as a discrete unit. The other groups have low WGMS values and share high BGMS values. These groups show a marked deviation from the expected distribution of species ( appendix 5 )« Group 1 contains species of Keiskea and Elsholtzia sections Elsholtzia and Cyclostegia, group 3 contains species of Elsholtzia section Aphanochilus series Stenelasmeae and group 4 contains species of Elsholtzia section Aphanochilus series Platyelasmeae and Tetradenia. The greater similarity of species of Elsholtzia sections Elsholtzia and Cyclostegia to species of Keiskea rather than to species of Elsholtzia section Aphanochilus series Stenelasmeae is unexpected, while the juxtaposition of Elsholtzia section Aphanochilus series Platyelasmeae and Tetradenia in group 4 is also surprising. The five group scheme ( table 12) more nearly approaches the arrangement suggested by the simplified similarity matrix ( appendix 5 )• Species of Elsholtzia sections Elsholtzia and Cyclostegia, species of Elsholtzia section Aphanochilus series Stenelasmeae, species of Keiskea 104 and species of Comanthosphace and Leucosceptrum are all quite discrete groups, having high WGMS values compared to their BGMS values shared with the other groups. The species of Elsholtzia sections Elsholtzia and Cyclostegia have a high WGMS, that of species of Elsholtzia section Aphanochilus series Stenelasmeae is much lower. They share high BGMS values with each other and the other groups. The fifth group is again formed by species of Elsholtzia section Aphanochilus series Platyelasmeae and species of Tetradenia. As in the four group scheme the WGMS of 81.4% is very low and barely exceeds the BGMS of 78.6% shared with species of Elsholtzia sections Elsholtzia and Cyclostegia. A scheme for six groupis can be produced simply by removing the species of Elsholtzia section Aphanochilus series Platyelasmeae from group 2 and placing them in a separate sixth group ( table 13 )• The species of Tetradenia now forming group 2 have a WGMS of 94.7% and low values for the BGMS values shared with other groups. They are most similar to species of Elsholtzia sections Elsholtzia and Cyclostegia but form an obviously separate group. The new group 6 has a WGMS of 87.5% but has quite high BGMS values with group 2, species of Tetradenia, group 4, species of Elsholtzia sections Elsholtzia and Cyclostegia and particularly with group 1, species of Elsholtzia section Aphanochilus series Stenelasmeae. The WGMS of 83.6% for group 1 is only a little higher than the BGMS of 78% shared with group 6. Throughout the analysis Eurysolen gracilis ( OTTJ 126 ), Leucosceptrum plectranthoideum ( OTU 97 ) and Dysophylla mairei ( OTU 102 ) are grouped with species of Elsholtzia section Aphanochilus series Stenelasmeae. The simplified similarity matrix of run 2B ( appendix 6 ) is 105 Table 11 • WGMS and BGMS values for four groups in the clustering to maximise WGMS analysis run 2A. 1 90.2 2 82.4 86.3 3 82.4 82.4 90.1 4 78.4 77.5 85.5 88.5 5 87.1 82.6 81.4 78.6 88.4 1 2 3 4 5 Key to groups 1 - species of Pogostemon 2 - species of Pogostemon 5 - species of Dysophylla sectiors Oppositifoliae and Verticillatae 4 - species of Dysophylla section Verticillatae 5 - species of Pogostemon ( for exact distribution of OTU's see appendix 10) 106 Table 11 • WGMS and BGMS values for four groups in the clustering to maximise WGMS analysis run 2A. 1 89.3 2 82.2 86.3 3 78.3 77.9 89.3 k 83.1 82.8 83.4 87.1 1 2 3 4 Key to groups 1 - species of Pogostemon 2 - species of Pogostemon 3 - species of Dysophylla section Verticillatae k - species of Dysophylla section Oppositifoliae and Pogostemon ( for exact distribution of OTU's see appendix 10) 107 rather confused but five groups can be distinguished, although there is a considerable amount of overlap between them. Appendix 10 and table 14 show the groups and their WGMS and BGMS values. The WGMS values are high, particularly for groups 1 and 3, but each group shares a high BGMS value with at least one other group ( tahle 14 ). For example group 3 with a WGMS of 90.has a BGMS of 85.5% with group 4. Group 1 has a WGMS of 90.2% but has a BGMS of 87.1% with group 5 and 82.4% with groups 2 and 3- As with the simplified similarity matrix ( see p. 90 ), although each group seems to be tight-knit, none is completely separable from all the others. If the number of groups is reduced to four, similar results are obtained ( table 15 and appendix 10). Groups 1 and 2 are formed by species of Pogostemon ( benghalensis complex and brachystachyus / mollis complex respectively ) and group 3 by species of Dysophylla section Verticillatae. Group 4 contains species of Dysophylla section Oppositifoliae and seven Pogostemon species. However it is interesting to note that in both schemes the group containing species of Dysophylla section Oppositifoliae ( group 3 in the five scheme, group 4 in the four scheme ) share relatively high and roughly equal values for BGMS with all other groups. This group appears to occupy an intermediate position between the other groups. In both schemes Elsholtzia aquatica ( OTU 22 ) is included in the same group as species of Dysophylla section Vert ic illat ae. 5.3. PRINCIPAL CO-ORDINATES ANALYSIS. A general view of similarities can be obtained by looking at two- dimensional plots using the first four eigenvectors of the principal 108 co-ordinates analysis ( figs 14-16 ). The first four vectors account for 42.7% of the total variation and, although low, this percentage is sufficiently high to provide a basis for comparing the groups formed. The figures generally show two large, well-separated clusters; species of Elsholtzia, Comanthosphace, Leucosceptrum, Keiskea, Eurysolen and Tetradenia form one cluster, species of Pogostemon and Dysophylla form the other, with an intermediate position occupied by the two species of Colebrookea. The two species of Rostrinucula are quite isolated from other points, particularly in fig. 14, although less distinctly in figs15»l6. In the Pogostemon / Dysophylla cluster, the species of Dysophylla are concentrated at one end of the cluster and the species of Pogostemon at the other. Similarly within the Elsholtzia / Comanthosphace cluster the species of Comanthosphace and Leucosceptrum canum are concentrated at one end of the cluster; in fig.16 they form a completely separate group. Neither the species of Keiskea, Eurysolen nor those of Tetradenia can be separated from species of Elsholtzia. A clearer picture of the features shown by the main clusters can be obtained from the second runs of the analyses. Figs 17 - 19 are two-dimensional plots for the Elsholtzia / Comanthosphace cluster and clearly demonstrate the separation of the various groups. Species of Comanthosphace and Leucosceptrum canum form a separate group in fig.17 and occupy a well-defined region at the edge of the main cluster in figs 18 and 19. The fact of Leucosceptrum canum ( OTU 7 ) as a member of the group is unequivocal. Comanthosphace nanchuanensis ( OTU 134 ) is also a member of the group in fig. 17 although it is rather distant 109 Figs. Two-dimensional plots of the principal co-ordinates analysis using the four largest eigenvalues as axes. Key. species of Pogostemon • species of Dysophylla O species of Elsholtzia A species of Comanthosphace • species of Keiskea A species of Tetradenia ® species of Leucosceptrum ® species of Rostrinucula (D species of Colebrookea © species of Eurysolen (?) 110 Fig. 14 . vector 1 yt •v Am A at (r)ia A" ^ A*" "ffAA"* A'° ©,lv A" A" A®^T) A z» A* ' A- Obo A" A* AA« ©©' AJ A" A* o O** A*4 A^A'V An O,ov ^ A3' A* O^ aA3« 03 OS«> O "5 ,0k o'* *5o O" „ 0,I?0 A" O31 CFO* o,c o1*0 cT vector 2 111 Fig. 14 . vector 1 A0,A'J A" A'3' A^A* A"* A^A"* A' A.» _ A A' o 10 #iT #»< o o " a'n A' os^cFd*0 o JW O" (T)- A- r A* vir A15 o,01> 'iktfd* A- w or A» (R) A» A39 A* A- 1 A' A* (RJ* vector 2 112 Fig. 14 . vector 1 3 A * Q.oi A» n A Aj» A" An A* "A A a* A» w A*1 A' A". (T)* A" A'" i 13 A • fV A'» A*' L IS 8 9 H •> ~ AI A"10 a»5 A A^ • • Oo»"« r i3b AIX3 5 1 A"* A'- A "* s A« o o o* O" O A" mw?* nmi (sfr vector 2 113 Figs 17i18,19 . Two-dimensional plots of the principal co-ordinates analysis of run 2A using the four largest eigenvalues as axes Key. species of Elsholtzia • species of Comant ho sphac e • species of Tetradenia (?) species of Leucosceptrum (T) species of Eurysolen (?) species of Keiskea A species of Dysophylla o 114 Fig.14 . vector 1 A,5A" AI? A1** AJ0 A1" A31 1 A** A« A" A'1" A (ST iia A' A' A (T) A"- A A,„ ^ A* aV A'*' A1'A13 • l J1* s • df I"1 • k vector 2 115 Fig. 14 . vector 1 ®'2 ® A'5 | A*®'" A" A*" , A2* A" A'vAq,A1 A,? A'"A' ? 3 A" A* ®< A * A" AIfc A*° Q.oi A I. A** A,J M«t A*? A*** a ®,2fc aV, A's A23 » 10 A" AW A A'51 A' A180 vector 2 116 Fig. 19 • vector 1 GT (t^TT A,Si A' AJ5 A'7* 5 A'" A'5 A ' A"* A" A3* A'* A '•» A 1 I Pt A*' A23 aV A" vector 4 117 Figs 20,21,22 • Two-dimensional plots of the principal co-ordinates analysis of run 2A showing the distribution of species of Elsholtzia Key. species of section Elsholtzia • species of section Cyclostegia O species of section Aphanochilus series Stenelasmeae a species of section Aphanochilus series Platyelasmeae a 118 Fig. 19 • vector 1 A1" a3V A** A1* Jt> £ A* Au A" •< A1' A'" i •V o • •V* O" O" vector 2 119 Fig. 14 . vector 1 A'1 O" i cr •V o" r A1 A3' A" A14* a31 a- A?5 A* A- A14 vector 2 120 Fig. 19• vector 1. A1* A« A" A» A Ai: O o,T A* vector 4 121 Figs 23, 24, 25 . Two-dimensional plots of the principal co-ordinates analysis of run 2B using the four largest eigenvalues as axes Key. species of Pogostemon q species of Dysophylla q species of Elsholtzia a 122 Fig.19 • vector 1 „0 bS lib •"2 o5T o50 "'od ScT 6 ow O" o5 « • • fK vector 2 123 Fig. Zb . vector 1 O O O1* #13 o Aw •V O* O" O^O'* o,M db* JW O1* •"if5® o,7t o"3 OP- •V vector 3 124 Fig. 14 . vector 1 3 A" oSfc •"•W o5' 105 (To* O o** Q» 0>vf 05> tf O,0,< o,n ®.35 ^ o"* O" O* o o' O O" • his vector 2 125 from the other members in figs 18 and 19. Spec ies of Keiskea form an obvious group in fig.18 • In figs17 and 19 they occupy an intermediate position within the main cluster, although with a distinct tendency to align with species of Elsholtzia sections Elsholtzia and Cyclostegia. Similarly species of Tetradenia form an obvious group in fig. 19. In figs17 andl8 they too occupy an intermediate position within the main cluster. Eurysolen gracilis ( OTTJ 126 ) does not separate away from the species of Elsholtzia section Aphanochilus series Stenelasmeae in any of the plots, always remaining within the group. Species of Elsholtzia are divided into two groups. The first represents sections Elsholtzia and Cyclostegia and occupies that end of the main cluster furthest away from species of Comanthosphace. The species forming these two sections are quite intermixed in all three plots. The second group represents section Aphanochilus series Stenelasmeae and occupies that part of the main cluster nearest to species of Comanthosphace. Leucosceptrum plectranthoideum ( OTU 97 ) and Dysophylla mairei ( OTU 102 ) lie within this group. The two groups are separated in the plots by species of Tetradenia, Keiskea, Elsholtzia section Aphanochilus series Platyelasmeae and E. concinna ( OTU 10 ). This division is more easily seen when only the.species of Elsholtzia are marked on the plots ( figs 20 - 22 ). E. eriostachya ( OTU 20 ) consistently segregates with species of section Aphanochilus series Stenelasmeae whilst E. densa ( OTU 21 ) segregates with species of sections Elsholtzia and Cyclostegia. The third member of section Aphanochilus series Platyelasmeae, E. manshurica ( OTU 125 ) occupies an intermediate position between the groups although in fig.20 it 126 is part of the sections Elsholtzia and Cyclostegia group. Figs 23 - 25 show the two-dimensional plots for the Pogostemon / Dysophylla group in run 2B. In all three plots the points form a rather straggly cluster. In fig.23 there are two outlying points, Dysophylla helferi ( OTU 108 ) and D. glabrata ( OTU kk ). The species of Dysophylla and Elsholtzia aquatic a ( OTU 22 ) are clustered on the left hand side of the plot while the species of Pogostemon always occupy the right hand side. Although the bulk of the species lie in one or the other half of the cluster, there is no sharp discontinuity but rather a gradation between them. 5.4 SINGLE-LINKAGE ANALYSIS. Fig.26 shows a dendrogram and the box-in-box classification derived from it. In this example the heirarchy begins at subtribe and ends with species; the dendrogram is a small portion ( approximately one-seventeenth ) of the single-linkage analysis dendrogram in fig. 27 The box-in-box classification derived from this position is already complex; that derived from all the full dendrogram would be inc omprehensible. The application of similar ranks to different heirarchy levels is well illustrated by the two species of Colebrookea. In the dendrogram ( fig. 26 ) these two species link at a second dichotomy; therefore the rank applied to this level must be the same as that applied to Rostrinucula, Eurysolen and Tetradenia, which also link at a second dichotomy. Table 16 shows the name-heirarchy represented by the box- in-box classification of fig. 26 . Colebrookea oppositifolia and C. ternifolia rank as genera as do Rostrinucula, Eurysolen and Tetradenia. 127 Fig. 26 . A single-linkage dendrogram and the box-in-box classification derived from it. • • •8CTIR(H) •9COPP(G) -122TGOU(F) •123 THIL(E) 124TFRll(D) 126EGRAIC) 95RDEP(B) 96RSIN (A) • • J i OJ 1 »i #1 i • • E -d- o to -P f>» s to o fl a o Sb o ^ 0) o •H o •H o bO •p -P -p O •H o O o CD t1 ' uoei ** & i- w z i — u u0 wo 0 o» « fig o ?1< 3 « O"^ 53ae$ n <•> ti Table 16 . Name hierarchy for eight taxa represented in the box-in-box classification shown in fig. 26 . Tribe Subtribe Genus Section Species ABCDEFGH ABCDEF AB A A B B C C DEF DE D E F F GH G G H H The letters in the table correspond to the taxa shown in fig. 26 129 However the two species of Colebrookea link at the 95% similarity level while the other three " genera 11 link at the 81% similarity level. This method of producing a classification is inaccurate and inadequate. Table 17 shows the classification produced when three phenon - lines are used to indicate ranks in the dendrogram shown in fig. 27. The ranks, at 79%i 82% and 86% similarity levels, were selected to represent tribe, subtribe and genus respectively. This much simplified classification shows a single tribe containing five subtribes and seventeen genera, nine of which are monotypic. Placing the tribe rank at the 79% similarity level is a fairly safe choice, since this study encompassed a single tribe. ( The taxa might in fact represent one or several tribes but, without external comparisons, this cannot be demonstrated ). The selection of the 86% similarity level for generic rank would also seem reasonable since at this level the taxa divide into a number of fairly discrete " chunks 11 ( cf. McNeil 1979 ) while below 86% there is no level at which obvious divisions of the taxa occur. However, use of the 86% similarity level does yield a rather large number of monotypic genera ( nine out of seventeen ). The data for five of these " monotypic " genera, Elsholtzia beddomei ( OTU 129 )i E. integrifolia ( OTU 23 ), E. penduliflora ( OTU 25 ), Pogostemon litigiosus ( OTU 121 ) and Comanthosphace nanchuanensis ( OTU I3*f ), were coded from single specimens. In view of such a restricted sample, the positions of these taxa must be treated with caution. It is feasible to designate the rank of subtribe which must lie between the 79% and 86% similarity levels ( tribe and genus respectively ). From the dendrogram, the best choice would appear to 687 921 S6 96 o n t m 2 {' S c 9 fz 52 21 oz IZ sei V2 I 62V 00 V <59 22 oil it ll ( U c 91 t Zi <' 28 ( 91 (...., IS C 9 8 ^S 021 ( • i ( SZ • • (• • Ul (• • OJ < 6S ( SOI { 9*1 c 05 I 8V ( cs ( Jot ( 25 I *0l ( 6oV ( 15 ( 6*t ( VOV ( ss ( 9S ( < 8£t c S* ( 901 ( { SOl 9S (• 06 ( 06 (• 89 (• f 9 (• 19 (• Sit ( *9 ( 19 ( 29 ( £6 ( £9 ( c •" 90 , ( 611 ( 99 ( 9t t { Zlt 111 c Zl I ( 18 (• o<5 26 ill SB ll 62 SU 16 fu 16 62 «Z 01 i2 201 I* 92 6£ 05 IS 0-i 81 9C S£ £c % 2Z l£ a s- Bl c\•j 91 hO 6 l •H 82 V 11 CI (. Si < 11 X f< 66 { 86 'tO I l£t ££l oci 2(1 I : < ( ( ( ( ((((((((((({ f 6i 0' t8 0 ' (9 O'SB 0'Z8 0*68 O'U 0" £6 0' S6 0'16 o '66 0' 08 28 o'ys o 98 0* 88 0* 06 0* 26 O't 6 0'96 0'8 Table 17. Classification of the Pogostemoneae produced by use of three phenon-lines in the s^gle-linkage dendrogram, <1) cd tQ i w PQ H § M PQ « tD EH 03 132 be 81% giving four subtribes, 82% giving five subtribes or 83% giving six subtribes. Intuitive knowledge of the Pogostemoneae suggests that perhaps the 82% level is marginally the best but no one level can be shown empirically to be a better choice than the others. The level to be designated for the rank of species is the most debatable. Several levels are possible candidates but no one level presents an obvious choice. Several taxa which have long been considered conspecific were included in the study in the hope that they could be used as indicators in deciding on a species level. However such taxa had to be coded from authentic material and this was usually restricted in each case to a single specimen ( the type ). The result was that while averaged data for one conspecific taxon was derived from several representative specimens, data for the other conspecific taxon was derived from a single specimen. This seems to have introduced sufficient distortion so that the presumed conspecific taxa e.g. Colebrookea oppositifolia ( OTU 8 ) and C. ternifolia ( OTU 9 ), Elsholtzia densa ( OTU 21 ) and E. manshurica ( OTU 125 ), link at relatively high levels of similarity. Again it is familiarity with the taxa which suggests that the 97% similarity level is the most suitable for the rank of species C see fig. 27 ). In view of the doubts as to the accuracy of this, units which link at or below the 97% similarity level are indicated as only possibly conspecific in the formal treatment ( see pp. 219 - 231 ). 133 6. DISCUSSION OF RESULTS. In interpreting the results of the various analyses it has been necessary to apply some concept of rank to the clusters of OTU's. The OTU's used in this study were species and the study was confined to a single tribe. Thus the rank of any aggregate of OTU's must be higher than species but lower than tribe. It would, of course, be possible to divide the units into a number of smaller tribes but, as explained in the previous section ( p. 129), this is impossible without reference to taxa representing other tribes. I have recognised only three supra-specific ranks,: section, genus and sjibtribe. 6.1. RANKS. Genera were taken to be clusters of OTU's which were clearly separable from other OTU's or clusters of OTU's. In the single- linkage analysis they are defined by the genus phenon-line ( see fig. 27). In the clustering to maximise WGMS they are groups which share low BGMS's relative to their WGMS's. In the principal co- ordinates analyses they are clusters which are completely separated in at least one of the two-dimensional plots ( see figs Ik - 25 ). Sections were taken to be clusters of units which form at higher levels than genera. In the single-linkage analysis they link above the genus phenon-line. In the clustering to maximise WGMS they share high BGMS's relative to their WGMS's. In the principal co- ordinates analyses they cannot be completely separated in any of the two-dimensional plots. Subtribes were taken to be clusters of clusters formed at lower levels of similarity than genera. In the single-linkage analysis they 691 are defined by the subtribe phenon-line ( see fig. 27 )• In the principal co-ordinates analyses they are separable in at least one of the two-dimensional plots. In the clustering to maximise WGMS the BGMS's shared by member clusters of one subtribe may be low but must obviously be higher than BGMS's shared by member clusters of differing subtribes. Member clusters of the same subtribe generally share WGMS values in excess of ?k%* 6.2. SUBTRIBES. A comparison of the subtribes which can be derived from the various analyses is shown in table 18. Principal co-ordinates analysis ( figs - 16 ) would provide a very neat division into Pogostemon / Dysophylla versus all other groups were it not for species of Colebrookea and Rostrinucula occupying the middle ground where the division would occur. As it is there are three alternative subtribe divisions ( labelled PCA 1, 2 and 3 in table 18 ). However PCA 2 and 3 are unlikely divisions, not supported by any of the other analyses and may be discounted. In the clustering to maximise WGMS ( table 8) the BGMS values indicate that the genera form four " groups of groups " which match those of PCA 1. This arrangement of subtribes is partially repeated in the single-linkage analysis ( fig. 27 ) which shows subtribes Comanthosphacineae and Colebrookineae. Elsholtzia and Keiskea are again placed together in subtribe Elsholtzineae but this time with Pogostemon and Dysophylla, while Tetradenia is the only genus of subtribe Tetradenineae. Eurysolen is also removed from subtribe Elsholtzineae to a subtribe of its own, Eurysolenineae. 135 Table 18 . Six arrangements of subtribes within the Pogostemoneae derived from different analyses. 5 Fh cd cd Cd •p cd a> •H d •H O ft^ H M a> a H o -P H a> u -p cd PJ H a) H o S to H o fl o o tQ •P 5 H CD O CQ I O O O tQ CD ft 04 hi CD It is possible to produce a third arrangement of subtribes incorporating groupings from both the single-linkage and principal co-ordinates analyses ( table 18 ). In this third arrangement Elsholtzia, Keiskea and Tetradenia form one subtribe as in the principal co-ordinates and clustering to maximise WGMS analyses. The nearest neighbours for each species of Tetradenia ( excluding other species of Tetradenia ) are species of Elsholtzia with which they share relatively high similarities ( see appendix 3 ). The only exception is Pogostemon rotundatus ( OTU 90 ) which is one of the five nearest neighbours of Tetradenia goudotii ( OTU .122 ). The high similarities shared by Tetradenia and Elsholtzia suggest that they should be placed in the same subtribe and not in separate subtribes as in the single-linkage analysis. Eurysolen is placed on its own in a separate subtribe as in the single-linkage analysis ( fig. 27 ). This is supported by the simplified similarity matrix ( appendix k ) which shows Eurysolen as being not closely similar to any other OTU. Pogostemon and Dysophylla form a subtribe as in the principal co-ordinates and clustering to maximise WGMS analyses. The subtribes common to both single-linkage, principal co-ordinates and clustering to maximise WGMS analyses, i.e. Colebrookineae and Comanthosphacineae, are retained. In an attempt to resolve the problem of choosing one arrangement of subtribes comparative tables showing WGMS and BGMS values of the various subtribes were drawn up for the three schemes ( tables 19, 20, 21 ). Despite some radical differences in the composition of the subtribes all three arrangements show a very similar range of values for both WGMS and BGMS. For example in 137 arrangement one ( table 19 ) subtribe Elsholtzineae contains Elsholtzia, Keiskea, Tetradenia and Eurysolen; in arrangement two ( table 20 ) it contains Elsholtzia, Keiskea, Pogostemon and Dysophylla; in arrangement three ( table 21 ) it contains only Elsholtzia, Keiskea and Tetradenia. The WGMS values are 79-6%, 79-6% and 79-7% respectively, a difference of only 0.1%. This illustrates one of the weaknesses of phenetic methods. For any problem it may be possible to produce several hypotheses but have no empirical method of demonstrating which, if any, is the " best " solution. Although the groupings of genera within the Pogostemoneae cannot be resolved by the present evidence clearly there are suprageneric groups present which should be recognised. I have selected the subtribes of arrangement three ( table 21 ) as those which most accurately reflect the distinguishing characters of the taxa and therefore have most practical value. This also has the merit of being the arrangement closest, although at a lower rank taxonomically, to that of Bentham ( 1832 - 36 ) ( and later Endlicher, 1838, see table 1 ) who was the only major monographer of the Labiatae to designate ranks between genus and tribe ( Briquet, 1897, and Kudo, 1929» simply placed all these genera in the tribe Pogostemoneae). Support for my choice of subtribes also comes from studies of pollen and stomata. There has been no complete survey of pollen in the Pogostemoneae so it was not possible to score this data for computation. However sufficient data is available to make consideration of the evidence worthwhile. The pollen of Labiatae is conservative in its characters 138 Table 19 • WGMS and BGMS values for subtribes derived from principal co-ordinates and clustering to maximise WGMS analyses ( arrangement one ). 1 79.6 2 72.1 85.8 3 72.2 70.1 8^.3 k 69.7 68.0 72.0 95.2 1 2 3 k Key to groups. 1 - Elsholtzineae ( Keiskea, Elsholtzia, Tetradenia, Eurysolen ) 2 - Comant ho sphac ineae ( Comanthosphace, Rostrinucula, Leucosceptrum canum ) 3 - Pogostemonineae ( Pogostemon, Dysophylla ) - Colebrookineae ( Colebrookea ) 139 Table 20 • WGMS and BGMS values for subtribes derived from single-linkage analysis ( arrangement two ). 1 79-6 2 71. 85.8 3 70.6 71.3 9^.7 k 71.1 70.9 70.8 95.2 5 7^.5 72.2 69.** 71.1 100 1 2 3 k 5 Key to groups. 1 - Elsholtzineae ( Keiskea, Elsholtzia, Pogostemon, Dysophylla ) 2 - Comanthosphaoineae ( Comanthosphace, Rostrinucula, Leucosceptrum canum ) 3 - Tetradenineae ( Tetradenia ) k - Colebrookineae ( Colebrookea ) 5 - Eurysolenineae ( Eurysolen ) 1*fO Table 21 . WGMS and BGMS values for subtribes derived from all analyses ( arrangement three ). 1 79.7 2 72.1 85.8 3 72.2 70.1 k 69.7 68.0 72.0 95.2 5 73.8 72.2 72.8 71.1 100 1 2 3 5 Key to groups. 1 - Elsholtzineae ( Keiskea, Elsholtzia, Tetradenia ) 2 - Comanthosphacineae ( Comanthosphace, Rostrinucula, Leucosceptrum canum ) 3 - Pogostemonineae ( Pogostemon, Dysophylla ) k - Colebrookineae ( Colebrookea ) 5 - Eurysolenineae ( Eurysolen ) 1VI and only two features, the number of colpi and the number of nuclei, show taxonomically useful variation; the pollen grains are either bi-nucleate / tri-colpate or tri-nucleate / hexa-colpate ( see pp. 23 - 2k ). A number of authors ( e.g. Wunderlich 19631 and EL-Gs#zar'&Watson 1968 ) have advanced the view that these characters have a high taxonomic value and have used them to good effect in studies of the relationships within the Labiatae and closely related groups. Pollen from eight genera in the Pogostemoneae has been examined by other workers and to provide a full sample of the genera I examined pollen from specimens of Leucosceptrum canum and Eurysolen gracilis under the light microscope. Since the species of any one genus ( as far as is known ) only produce one type of pollen the data can be summarised as in table 2 • Using this summary a number of points can be made. Confirmation of Kudo's ( 1929 ) separation of Rostrinucula with bi-nucleate / tri-colpate pollen, from Elsholtzia with tri-nucleate / hexacolpate pollen, has already been pointed out by Wunderlich ( 1963 ) ( see p. 2k)* The pollen type also confirms my placing Rostrinucula in the Comanthosphacineae since Comanthosphace also has bi-nucleate / tri-colpate pollen. The position of Leucosceptrum canum in this subtribe is not necessarily supported since species of the Ajugoideae, to which L. canum might perhaps belong,also have bi-nucleate / tri-colpate pollen. Tetradenia seems to be correctly placed with Elsholtzia and Keiskea in the Elsholtzineae, these three genera being the only ones to have tri-nucleate / hexa-colpate pollen. Pogostemon, Dysophylla 142 and Colebrookea, with which some early authors ( e.g. Bentham & Hooker 1876 ) have placed Tetradenia,have bi-nucleate / tri-colpate pollen. The separation of Eurysolen and Elsholtzia into two subtribes, for which there is perhaps least evidence from my analyses, is also confirmed. The only species of Eurysolen, E. gracilis, has tri- colpate ( and by implication bi-nucleate ) pollen and not tri- nucleate ( and hexa-colpate ) pollen which is found in all the species of the Elsholtzineae which have been examined. It should also be said perhaps, that as well as supporting these specific points, the pollen data lend weight to the general argument for a division of the Pogostemoneae into subtribes, with Elsholtzia, Keiskea and Tetradenia forming one group and the remaining genera forming at least one other group. Studies of stomata from the lower epidermes of leaves carried out by El-Gazzar& Watson ( 1968 ) provide similar results to those of pollen data. ( No data is available for stomata in Eurysolen gracilis ). Although El-Gazzar & Watson believe stomatal type to be a less reliable character than pollen type it nevertheless lends further support to my division of the Pogostemoneae. The polygon in fig. 28 gives a diagramatic representation of the BGMS values for the five subtribes. The thickness of the line joining any two subtribes is proportional to their BGMS value but the lines have been exaggerated to make the relationships more obvious. Species of Comanthosphace and Rostrinucula were formerly included in Elsholtzia ( see p. 16, fig. 2 ) and as might be expected, the Comanthosphac ineae are most closely allied to the Elsholtzineae. The Eurysolenineae too, 143 Fig. 28 . Diagrammatic representation of the BGMS values for subtribes derived from a combination of analyses. Elsholtzineae Eurysolenineae Comanthosphac ineae Colebrookineae Pogostemonineae The width of the line joining two subtribes is proportional to the similarity shared by those subtribes C x - v ) width of line joining a & b = *__ mm where x = BGMS of subtribes a & b y = lowest BGMS of any subtribe are closely allied to the Elsholtzineae but also show a strong link with the Pogostemonineae. In turn the Pogostemonineae have roughly equal affinities with the Elsholtzineae and Colebrookineae as well as the Eurysolenineae; they are much less similar to the Comanthosphacineae. Bentham ( 1832 - 36 ) described Pogostemon and Dysophylla as forming an isolated group whose affinities placed it between the tribes Pogostemoneae and Ocimoideae. Without reference to members of the Ocimoideae the validity of this cannot be checked. However the Pogostemonineae are not isolated from the other subtribes of the Pogostemoneae. The most isolated of the subtribes is the Colebrookineae which is relatively weakly linked with the Elsholtzineae and very weakly so with the Comanthosphac ineae. The most appropriate linear sequence for the subtribes is Comanthosphac ineae, Elsholtzineae, Eurysolenineae, Pogostemonineae and Colebrookineae and they are discussed in this order below. 6.3. SUBTRIBE CQMANTHQSPHACINEAE. 6.3.1. Comanthosphace and Leucosceptrum. As a single group the seven species of Comanthosphace, together with Leucosceptrum canum ( OTU 7 ) are readily separated from other taxa in each of the analyses ( Leucosceptrum plectranthoideum is discussed on p. 173 ). The two species of Rostrinucula may also be regarded as part of this group and are discussed later ( pp. 148 -152 ), The relationships between species of Comanthosphace and Leucosceptrum canum are complex. Six of the species of Comanthosphace are morphologically very similar to each other and form an acceptable group. The seventh species, C. nanchuanensis ( OTU 134 ) is much less 14-5 similar to the other six, and less obviously a member of the group. Leucosceptrum canum is most similar to species of Comanthosphace but is sometimes separated from them. For example the single-linkage dendrogram (fig. 27) shows Comanthosphace pro ma.jore and Leucosceptrum canum as distinct genera, not linking until the 85% level* Comanthosphace as a genus can be recognised by a number of characters and character combinations. The most obvious and consistent are the broad, membranous, deciduous bracts and narrow, deciduous bracteoles, the closed annulus of hairs within the corolla, the hairy nutlets and the dense pubescence of branched and unbranched hairs on all parts of the plant. In addition the flowers are large with relatively short, more or less equal calyx teeth; the upper lobe of the corolla is emarginate and the anthers are unilocular. Comanthosphace nanchuanensis appears to lack bracteoles although this may be recorded in error since both bracts and bracteoles fall very early and are not present on every specimen. Similarly the glabrous appearance of the nutlets may be erroneous since no fully mature nutlets were available. The annulus within the corolla is incomplete. All the other characters distinguishing species of Comanthosphace are shared by C. nanchuanensis and the inclusion of this species within the Comanthosphace group seems intrinsically correct. Certainly the difference in overall similarity between Comanthosphace pro ma.jore and C. nanchuanensis is not sufficiently great to justify the removal of the latter in a separate genus. Leucosceptrum canum differs from species of Comanthosphace in two of the same characters as C. nanchuanensis. Bracteoles are present, but therfe are no annular hairs within the corolla, and the nutlets 146 are glabrous. In addition Leucosceptrum canum has a terminal style whereas all species of Comanthosphace, including C. nanchuanensis, have gynobasic styles. With the exception of the style character Leucosceptrum canum is scarcely less similar to Comanthosphace than is C. nanchuanensis, as shown by the similarities in tables 9 and 10 and the present concept of the genus Comanthosphace might easily be expanded to include Leucosceptrum canum. The gynobasic versus terminal style character is considered by most authors to be one of a number of characters which distinguish taxa at the tribal level in the Labiatae. Most authors attach greater importance to these characters than to others; in effect they give a weighting to the tribal characters. The analyses used here are all unweighted methods and cannot reflect this importance, whether or not it is taxonomically valid to do so.' In a weighted method giving greater consideration to characters of high taxonomic value, such as the style character is usually considered to be, the augmented contribution of this character would reduce the similarity between Leucosceptrum canum and species of Comanthosphace. Many authors accept the absolute veracity of the two states of the style character, i.e. gynobasic versus terminal, and much use of it has been made in studies of the Labiatae and Verbenaceae. I remain unsure of the delimination of these character states. My own experience is that the style type and nutlet attachment are not easily or clearly interpreted, at least for some species of the Pogostemoneae. The style character is linked with a nutlet character so that in the gynobasic condition the nutlets have a small, basal attachment 147 scar; in the terminal condition the nutlets have a large lateral attachment scar. A terminal style and lateral attachment scar are usually regarded as characteristic of the subfamily Ajugoideae, and on the basis of these characters, Leucosceptrum canum is usually placed in this group. In most species this pairing of style and nutlet characters holds true, but in others the link appears to break down. In Elsholtzia densa ( OTU 21 ), for example ( figs 10,11.31 ) the style is gynobasic but the nutlets have a relatively large, somewhat oblique scar; in E. flava ( OTU 28 ) the style is apparently gynobasic and the nutlet scars small but clearly lateral ( figs 10,11 ). In Leucosceptrum canum(fig.10)the style is not always obviously terminal, sometimes appearing to rise almost directly from the disc. This divergence from the defined condition is reinforced by the nutlet scars which are small and basal. In this respect they are very similar to the nutlets of species of Comantho sphac e ( cf. fig.11 ). The nutlet characters may have been wrongly assessed in Leucosceptrum canum,a claim which has already been made by Kitamura & Murata ( 1962 ) . However, they failed to take into account the apparent disparity of style characters between the tribe Ajugeae and the Satureieae, the tribe to which they transferred it. A more detailed study of gynobasic versus terminal styles in a wide range of species is required to assess the variation and importance of this character in the Labiatae. Should the style character prove to be less important than supposed, Leucosceptrum and Comanthosphace show sufficient overall morphological similarity to be considered congeneric. However, until the style character is reassessed, no conclusive statement can be made. 148 6.3.2* Rostrinucula. The simplified similarity matrix ( appendix 4 ) suggests that the two species of Rostrinucula ( OTU's 95 and 96 ) may form part of the Comanthosphace / Leucosceptrum group. The clustering to maximise WGMS analysis also places the two species of Rostrinucula in this group which shares low BGMS values with other groups but also has a low WGMS ( see tables 6, 7 ), suggesting that the group is not homogenous. If the two species of Rostrinucula are removed as a separate group, the WGMS values of both this and the Comanthosphace / Leucosceptrum group rise sharply without any significant rise in the BGMS values ( see table 8 ). The BGMS shared by the Rostrinucula group and the Comanthosphace / Leucosceptrum group is also sufficiently low to support this division of taxa. The principal ' co-ordinate plots similarly emphasise the link between species of Rostrinucula and species of Comanthosphace. The species of Rostrinucula lie between the two main clusters in each plot ( figs 14, 15, 16). However they are closest to, although always separate from, species of Comanthosphace. In the single-linkage dendrogram ( fig. 27 ) the designated genus level is at 86% similarity. The species of Rostrinucula do not link until well below this, at the 83% similarity level, and then only with species of Comant ho sphac e and Leucosceptrum canum. Rostrinucula dependens ( OTU 95? fig- 29 ) was originally described as Elsholtzia dependens by Rehder ( 1917 ) who stated that it seemed " not closely related to any other species of the genus My own investigations also refute any strong link with Elsholtzia. Instead there is a clear indication here that while Rostrinucula is 149 Fig, 29. Rostrinucula dependens. a. habit x \ b. calyx x 10 c. corolla x 5 d. dissected flower x 5 e. stamen x 10 f. annular hairs at base of stamen filament x 40 g. nutlet, inner face and profile x 5 151 sufficiently distinct to be upheld as a genus, its phenetic affinities lie with species of Comanthosphace and Leucosceptrum canum, these genera being constantly linked with it. Indeed Rostrinucula sinensis ( OTU 96 ) was originally described by Hemsley ( 1890 ) as Leusceptrum sinense, and only recently recognised by Wu ( 1965 ) as being a species of Rostrinucula. Species of Rostrinucula bear a striking overall resemblance to species of Comanthosphace, particularly in the long flower spikes with their broad, membranous, deciduous bracts and in the whitish indumentum of unbranched and branched hairs which cover all parts of the plant. The calyces are of similar size and shape, as are the corollas, and species of both genera have unilocular anthers with glabrous filaments, hairy nutlets and lack a tumescent gland on the disc. Since these are all characters which distinguish Comanthosphace and Leucosceptrum from other genera within the Pogostemoneae the phenetic affinities of Rostrinucula must lie hare. Rostrinucula, however, does not share all the characters of the Comanthosphace / Leucosceptrum group. Some features found in Rostrinucula are unique within the Pogostemoneae while others, although r occulting elsewhere in the tribe, are not found in species of Comanthosphace or Leucosceptrum. Rostrinucula dependens was named for its nutlets which each possess a long curved beak when mature ( figs11,29) This character is unique within the Pogostemoneae. Unfortunately the specimens of Rostrinucula sinensis available for examination had no mature nutlets so the character could not be assessed for this species but Wu ( 1965 ) described the nutlets as rostrate. Within the corolla of both species 152 of Rostrinucula ( cf. figs 8,29 ) is an annulus of hairs which, unlike that in species of Comanthosphace, is incomplete. The hairs are borne on swollen disc-like excrescences at the base of the anther filaments and on the inner surface of a crescent-shaped invagination below the base of the lower lip. An identical arrangement of filament-discs and invagination is found in Eurysolen gracilis ( figs 8,32 )• Rostrinucula dependens and R. sinensis differ from species of Comanthosphace, Leucosceptrum and Elsholtzia by the entire upper lip of the corolla ( cf. fig. 8 ). In this respect only the species of Rostrinucula more closely resemble species of Pogostemon and Dysophy11a. Although so closely linked to Comanthosphace / Leucosceptrum there can be no doubt as to 'the status of Rostrinucula as a separate taxon, distinguished by the nutlet and corolla characters given above. 6.4. SUBTRIBE. ELSHQLTZINEAE. 6.4.1. Elsholtzia. The results of the analyses show that the species of Elsholtzia can be regarded as forming one group embracing all but one taxon, E. aquatica ( OTU 22 ), or alternatively they can be regarded as two or three groups sharing high BGMS values. For the sake of clarity E. aquatica, which is now considered to be a species of Pogostemon ( see pp. 208 -210) is omitted from the following discussion. A brief summary of the number of groups formed by each analysis is given below. Simplified similarity matrix ( appendices 4, 5 ) - one main group but within this two or perhaps three subgroups. Clustering to maximise WGMS, ( . tables 6,7,8^" one or gr°ups. Clustering to maximise WGMS, run 2A. (tables 11-13) - three groups. 153 Principal co-ordinates analysis run 2A ( figs 17-22 ) - one main group containing two subgroups. Single-linkage analysis ( fig. 27 ) - two groups and three isolated species. When one group is formed it corresponds to the genus Elsholtzia as described by Bentham ( 1832 - 36 ) and other authors. When two groups are formed thay correspond to section Elsholtzia with section Cyclostegia of Bentham and section Aphanochilus series Stenelasmeae of Briquet. The section Aphanochilus series Platyelasmeae Briquet containing three species becomes divided up. Two species, E. densa ( OTU 21 ) and E. manshurica ( OTU 125 ), become included within the group comprised of sections Elsholtzia and Cyclostegia and the third species, E. eriostachya ( OTU 20 ), becomes included with the group comprised of section Aphanochilus series Stenelasmeae. When three groups are formed they correspond to sections Elsholtzia and Cyclostegia Bentham, section Aphanochilus series Stenelasmeae Briquet and section Aphanochilus series Platyelasmeae Briquet. 6.4.2. Elsholtzia sections Elsholtzia and Cyclostegia. The group corresponding to -Benthamfs sections Elsholtzia and Cyclostegia appears as a single close-knit group in every analysis. Even E. densa ( OTU 21 ) and E. manshurica ( OTU 125 ) are sometimes included within this group as in the clustering to maximise WGMS analysis ( see .table 7* appendix 8 ). Bentham ( 1829 ) described two genera, Aphanochilus and Cyclostegia, which he later reduced to sections of the genus Elsholtzia, distinguishing the sections on the basis of variation in bract and 154 inflorescence characters. Section Aphanochilus has spikes equal with lanceolate or ovate bracts or spikes secund with lanceolate bracts. Section Cyclostegia has dense spikes and connate, imbricate, cyathiform, membranous, veined bracts with ciliate margins. Section Elsholtzia has broadly ovate spikes and broadly ovate, secund bracts. These sections were accepted by all subsequent authors with the exception of Kudo ( 1929 )i who reverted to Bentham's original concept of Aphanochilus and Elsholtzia as separate genera. Kudo assigned two species of formerly unknown sectional affinities to Elsholtzia, E. luteola ( OTU 16 ) and E. heterophylla ( OTU 18 ), which were later ascribed by Wu and Huang ( 1974 ) to section Cyclostegia. However,the type species of section Cyclostegia, E. strobilifera ( OTU 19 ), was not included in Kudo's work and he did not make clear whether or not he considered Elsholtzia and Cyclostegia to be con-sectional. Several new species were ascribed to both sections by Wu and Huang ( 1974 ) but the sectional definitions remained those of>Bentham. There seems to be no reason for recognising two sections since in all of my analyses sections Elsholtzia and Cyclostegia form a single, close- knit group. Table 22 shows Bentham's characters scored for all the species of sections Elsholtzia and Cyclostegia. All species have more or less dense inflorescences and imbricate, ciliate bracts. All species have veined bracts although in S. conclniia ( OTU 10 )-..and E. kachinensis ( OTU 12 ), which have green, non-membranous bracts, the veins are less obvious than in other species which have membranous and often brown bracts. E. concinna and E. kachinensis are the only species with non- membranous bracts ( see fig. 5 )• Four combinations of the three characters are represented: Table 22. Bract characters in Elsholtzia sections Elsholtzia and Cyclostegia Bentham. SPECIES CHARACTERS 1 2 3 4 5 6 7 8 E. concinna + + + + - - - - E. kachinensis + + + + - - - - E. hunanensis + + + + + - - - E. ciliata + + + + + — - + section E. pygmaea + + + + + - - + Elsholtzia E. soulei + + + + + - - + E. argyi + + + + + - - + E. oldhami + + + + + - - + E. nipponica + + + + + - - + E. pseudocristata + + + + + - - + E. feddei + + + + + - - + E. elegans + + + + + - - + E. luteola + + + + + + - + section E. strobilifera + + + + + + + - E. heterophylla + + + + + + + - Cyclostegia E. bodinieri + + + + + + + - Key to the characters. 1. dense + / lax - 2. imbricate + / not overlapping - 3. clearly veined + / obscurely veined - margins ciliate + / margins glabrous - 5. membranous + / non-membrabous - 6. connate + / free - 7. cyathiform + / not cyathiform - 8. secund + / cylindrical - ( see also figs 30, 31 )• 156 Bracts free, non-cyathiform, inflorescence cylindrical; E. concinna ( OTU 10 ), E. kachinensis ( OTU 12 ), E. hunanensis ( OTU 2k ). Bracts free, non-cyathiform, inflorescence secund; E. ciliata ( OTU 11 ) E. pygmaea ( OTU 13 ), E. soulei ( OTU 14 ), E. argyi ( OTU 15 ), E. oldhami ( OTU 99 )» E. nipponica ( OTU 98 ), E. pseudocristata ( OTU.101 ), 31.-feddei.l.OTU-127 ), E. elegans ( OTU 128 ). Bracts connate, non-cyathiform, inflorescence secund; E. luteola ( OTU 16 ). Bracts connate, cyathiform, inflorescence cylindrical; E. strobilifera ( OTU 19 ), E. heterophylla ( OTU 18 ), E. bodinieri ( OTU 17 ). Whichever combination of these three characters is used to divide the species into groups there is always some degree of overlap and at least one species, E. concinna, E. kachinensis, E. hunanensis or E. luteola will be an intermediate between the groups. Since four of Bentham's characters i.e. bracts dense, bracts imbricate, bracts veined and bracts with ciliate margins have proved to be identical throughout both sections ( see table 22 ) and the other characters, in any combination, always show intermediate species, the original distinction between the two sections is no longer valid. My data has produced no evidence for a division of these species into two or more higher taxa. Indeed the fourteen species representing sections Elsholtzia and Cyclostegia sensu Bentham consistently form a single homogenous group. Therefore I have placed all fourteen species in a single group, section Elsholtzia, which is characterised by orbicular, membranous bracts ( except in E. kachinensis and E. concinna ); the inflorescence may be cylindrical, strobilate or secund but always with imbricate bracts ( see ppQ 221 - 222 ). 157 Fig. 30. Elsholtzia luteola. a. habit x 1 b. pair of fused bracts x 5 c. pair of bracts opened out x 5 d. corolla x 10 e. dissected flower x 10 f. stamen x 20 g. nutlet, inner face and profile x 10 158 159 6.4.3. Elsholtzia section Aphanochilus series Stenelasmeae. The group corresponding to Briquet's section Aphanochilus series Stenelasmeae also appears to be distinctive in the results of each analysis eventhough E. eriostachya ( OTU 20 ) of series Platyelasmeae is sometimes included within it,as , for example, in the clustering to maximise WGMS analysis C see table 7 appendix 8 ). Section Aphanochilus series Stenelasmeae is a much less well-defined assemblage of OTU's than either section Aphanochilus series Platyelasmeae or section Elsholtzia sensu mihi. Some OTU's share many high similarities with other OTU's e.g. E. communis ( OTU 31 ), E. griffithii ( OTU 33 ), E. alopecuroides ( OTU 32 ) and E. glanduligera ( OTU 34 ). Others e.g. E. flava ( OTU 28 ) and E. fruticosa ( OTU 29 ) share high similarities with only a few other OTU's of the group. This results in a group which is well-defined at one end but rather diffuse at the other. This is very well illustrated in the simplified similarity matrices (appendices 4,5). In the clustering to maximise WGMS analysis section Aphanochilus series Stenelasmeae seems to suffer more from the " rag-bag 11 effect than any other group; loosely associated species such as Eurysolen gracilis ( OTU 126 ) and Elsholtzia integrifolia ( OTU 23 ) are included in the group together with the species of section Aphanochilus series Stenelasmeae simply because every species in the analysis must be placed in a group. However the make-up of the group is consistent, the same species being included within it each time. The principal co-ordinates analysis of run 2A ( figs 17-22 ) group the species of section Aphanochilus series Stenelasmeae as a large, open but well- defined area within the main species cluster. Despite the associated species in the clustering to maximise WGMS 160 analyses and the rather loose nature of the cluster in the simplified similarity matrix and the principal co-ordinates analysis, section Aphanochilus series Stenelasmeae is clearly and consistently distinguishable as a group in each of the analyses. Bentham ( 1829 ) and Kudo ( 1929 ) considered Aphanochilus to be a genus distinct from Elsholtzia, although Bentham qualified his view with the statement " perhaps my Aphanochilus might be united with it ( Elsholtzia ) as a second section " and later ( Bentham 1832 -36 ) did just that. Kudo distinguished Aphanochilus by the verticillate, often long and lax flower spikes, the lanceolate or sublanceolate bracts, the divergent, finally confluent anther locules, the equal disc and the shining nutlets. He distinguished Elsholtzia ( including species of Bentham's section Cyclostegia and Briquet's section Aphanochilus series Platyelasmeae ) by the many-flowered verticels in secund spikes, the broadly ovate, densely imbricate bracts, the emarginate upper lip of the corolla, the lower pair of stamens longer than the upper, the divergent anther-locules and the disc with a swollen nectary. However in all of Kudo's species of Aphanochilus and Elsholtzia the upper lip of the corolla is emarginate, the lower pair of stamens is longer than the upper, the anthers are bilocular and the locules are partially fused and the disc has a swollen gland or nectary. Species in both genera may have long, dense, secund flower spikes. Thus the only credible distinguishing character for separating Aphanochilus and Elsholtzia sensu Kudo is the bracts lanceolate or subulate versus the bracts broadly ovate and densely imbricate. Bentham ( 1829 ) similarly distinguished Aphanochilus, Elsholtzia and a third genus, Cyclostegia, by inflorescence, bract, corolla and 161 anther characters. Later ( Bentham 1832 - 36 ) he reassessed these characters and commented that 11 The three sections of Elsholtzia differ, in many respects, from each other in habit; but on closer examination of their characters, these distinctions do not appear to be of sufficient importance to preserve the genera Aphanochilus and Cyclostegia which I had originally established, but which I have now considered as mere sections of Elsholtzia. " Bentham characterised sections Aphanochilus and Elsholtzia as follows: section Aphanochilus. Spikes equal, bracts lanceolate to ovate or secund with bracts lanceolate, section Elsholtzia. Spikes and bracts broadly ovate, secund. The characters match those used by Kudo after the characters common to both taxa have been removed from Kudo's descriptions. Bentham's reassessment of his taxa is in close accord with the results presented here although, as discussed above, section Cyclostegia cannot be differentiated from section Elsholtzia and it is the species of section Aphanochilus series Stenelasmeae Briquet,not species of section Aphanochilus sensu lato,which form a subgroup within Elsholtzia and thus correspond with Bentham's " mere section Section Aphanochilus series Stenelasmeae was first described by Briquet in 1897; the bracts were described as small, linear-lanceolate or stiffly pointed and the nutlets shiny. Briquet created the series to differentiate the majority of the species in section Aphanochilus from E. densa and E. eriostachya which he placed in section Aphanochilus series Platyelasmeae, a series characterised by very short, broadly ovate or rounded bracts and dark, dull nutlets. The group shown in my analyses corresponds exactly to Briquet's concept of section 162 Aphanochilus series Stenelasmeae ( although it contains several species unknown to Briquet ) and is comparable to the group formed by species of section Elsholtzia and Cyclostegia. Since section Elsholtzia sensu milii is regarded here as a section, section Aphanochilus series Stenelasmeae has been raised to the rank of section ( see p. 223) The distinguishing features of section Aphanochilus sensu stricto given by Briquet are rather slight. E» flava ( OTU 28, fig. 10), for example, has broadly ovate, not linear-lanceolate bracts and there is no direct comparison with section Elsholtzia. Bentham's description, although referring to section Aphanochilus sensu lato, provides a more suitable basis for characterising section Aphanochilus sensu stricto. In order to accomodate some of the more recently described species of Elsholtzia Wu & Huang ( 197^ ) put forward eight new series within their section Aphanochilus subsection Stenelasmeae. However they confined their attentions to Chinese species and although the isolation of E. flava ( OTU 28 ) and E. penduliflora ( OTU 25 ) in separate series agrees with the positions shown in my analyses, on the whole their divisions are not supported by my results. Only three of Wu's & Huang's series, Fruticosae, Blandae and Communes, contain more than one species. Series Communes does show as a fairly clear group in the single-linkage dendrogram ( fig. 27) but the other two series are not evident in the analyses. In fact series Fruticosae appears to be a heterogenous group, five of the six species sharing their highest similarities with species of other series e.g. E. winitiana ( OTU 36 ) with E. blanda ( OTU 38 ) in series Blandae ( see appendix 7 ). There seems to be insufficient evidence for maintaining Wu's & Huang's series which are, anyway, unsatisfactory for a number of reasons. The 163 characters given by the authors do not adequately differentiate between series. As an example, series Capituligerae is described as " frutices parvi; spicae sphaeroideae; corolla 4-fida, labio superiore integra ". The entire upper lip of the corolla is incorrectly observed; all species of Elsholtzia ,including E. capituligera ( OTU 42 ),have an emarginate upper lip. All species have a 4-fid corolla, not just those of this series. Similarly series Communes is described as " herbae erectae; spicae cylindriceae compactae; corolla 4-fida, labio superiore integro 11. Again the description of the upper lip is incorrect and the 4-fid corolla is common to all species in the genus. Cylindrical, compact flower spikes are also found in many species in other series e.g. E. pilosa of series Pilosae and E. winitiana of series Fruticosae. 6.4.4. Elsholtzia section Aphanochilus series Platyelasmeae. The third group produced in the analyses corresponds to Briquet's section Aphanochilus series Platyelasmeae and contains three species; E. eriostachya ( OTU 20 ), E. densa ( OTU 21, fig. 31 ) and E. manchurica ( OTU 125 )• It is most clearly seen in the single-linkage analysis ( fig. 27 ) and the clustering to maximise WGMS of run 2A ( table 13 ) where the three species form a distinct group separate from, but sharing high similarities with, sections Aphanochilus sensu stricto and Elsholtzia sensu mihi. In the simplified similarity matrices ( appendices 4,5 ) series Platyelasmeae might arguably be placed with section Elsholtzia sensu mihi, E. manshurica especially sharing a number of high similarities with species in this section. E. densa shares high similarities with E. elegans ( OTU 128 ) of section 164 Elsholtzia sensu mihi and with E. glanduligera ( OTU 34 ) of section Aphanochilus sensu stricto. E. eriostachya does not share high similarities with any species from section Elsholtzia sensu mihi. E. densa, E. manshurica and E. eriostachya are however most similar to each other and may be regarded as forming a separate group. In the clustering to maximise WGMS (table 6,7»see appendix 8 ) and the principal co-ordinates analysis ( figs 14 - 16 ) the species of series Platyelasmeae do not form a separate group. E. eriostachya is included within section Aphanochilus sensu stricto while E. densa and E. manshurica are included within section Elsholtzia sensu mihi. If E. densa and E. manshurica are considered in isolation from E. eriostachya then on the basis of overall similarity they could acceptably be placed in section Elsholtzia sensu mihi. Similarly, if E. eriostachya is considered with sections Elsholtzia sensu mihi and Aphanochilus sensu stricto only, then it would undoubtedly be placed in section Aphanochilus. This is exactly the treatment given in some earlier works. Bentham ( 1832 - 36 ) placed E. eriostachya (and later, in 1848, E. densa )in his section Aphanochilus since, although the bracts were broad, like those of species of his section Elsholtzia, the inflorescence was not secund. Kudo ( 1929 ) placed E. densa ( as E. janthina Dunn ) in Elsholtzia, which he regarded as a genus in the narrow sense, although he described the inflorescence of Elsholtzia pro ma.iore as secund and that of E. densa as cylindrical ( see fig. 31 '). However, E. eriostachya and E. densa are much more similar to each other than they are to any of the species of either section Elsholtzia sensu auct. or Aphanochilus sensu stricto. This is quite clear from the nearest neighbours list ( appendix 7 ) as well as in the clustering to 165 maximise WGMS of run 2A ( table 13 ) and the single-linkage analysis ( fig. 27 ). The latter analysis shows that these three species form a distinct genus. This is similar to the view of Briquet ( 1897 ) who placed E. densa and E. eriostachya in a separate group characterised by having short, broadly ovate or rounded bracts and dark, matt nutlets. He considered this group to merit only the rank of series, section Aphanochilus series Platyelasmeae, apparently on negative evidence; although series Platyelasmeae has broad bracts, these are neither secund as in section Elsholtzia nor connate as in section Cyclostegia. In fact, the bracts of species of series Platyelasmeae in some ways resemble those of section Aphanochilus sensu stricto and section Elsholtzia sensu mihi ( cf. fig 5 )• Table 23 gives a comparison of these characters in each group. Species of section Aphanochilus sensu stricto generally have narrow, non-membranous, non-imbricate and usually green bracts. Species of section Elsholtzia sensu mihi generally have orbicular, membranous, imbricate and usually brown or bi-coloured bracts. Species of series Platyelasmeae have orbicular, non-membranous, non- imbricate and usually bicoloured or brownish bracts. In addition E. eriostachya, E. densa and E. manshurica resemble section Elsholtzia sensu mihi in lacking bracteoles, which are found in all species of section Aphanochilus sensu stricto. Examination of series Platyelasmeae reveals three distinctive features within Elsholtzia sensu lato. The fruiting calyx is not simply accrescent but becomes greatly inflated ( cf. fig. 7 )• The style lobes have clavate tips ( cf. fig. 10 ) and the mature nutlets are distinctly verrucose ( cf. fig. 11 ). E. densa, E. eriostachya and E. manshurica are almost exactly similar to one another with respect to their 166 Table 23• Bract characters in Elsholtzia. sect. Aphanochilus sensu ser. Platyelasmeae Briquet sect. Elsholtzia eensu! stricto. mihi. not as broad as long at least as broad as long at least as broad as long non-membranous non-membranous membranous ( excejt for E. concinna and E. kachinensis ) non-imbricate non- imbr ic at e imbricate usually green usually brown or usually brown or bicoloured bicoloured ( excejt for E. concinna and E. kachinensis ) ( see also figs 5, 30 & )• 167 qualitative character states. They differ significantly only in the variation of ten quantitative characters. These characters were scored as simple measurements and not converted into proportions ( see p. ^5 ). Consequently there is the possibility of distorted separation, particularly in the principal co-ordinates analyses, due to the size factors. This would explain the grouping of E. eriostachya with the generally small flowered species of section Aphanochilus sensu stricto and the larger flowered E. densa and E. manshurica with species of section Elsholtzia sensu mihi. Of previous treatments the closest to my results is that of Kitagawa ( 1935 ). He considered the calyx, style and nutlet characters highly significant and recognised Briquet's series Platyelasmeae as a distinct genus, Platyelasma, and added a new species, P, manshurica, to it. He not only recognised Platyelasma as a taxon distinct from Elsholtzia and Aphanochilus, but also at the same rank. Although the single-linkage analysis places E. densa, E. eriostachya and E. manshurica in a separate genus, all the other analyses show these species forming a group at similarity levels ( and thus ranks ) roughly equal to those at which sections Elsholtzia sensu mihi and Aphanochilus sensu stricto are formed. This, and the high similarity levels shared by sections Elsholtzia sengu mihi, Aphanochilus sensu stricto and Aphanochilus series Platyelasma does not justify recognition of Platyelasma as a genus. The most suitable treatment of E. densa, E. eriostachya and E. manshurica is to place them in a third section; Elsholtzia section Platyelasmeae. It is interesting to note here the monotypic genus Paulseniella described by Briquet ( 1908 ) from a plant collected in the Pamir, it has a campanulate calyx, becoming inflated in fruit, a barely exserted 168 Fig. 31• Elsholtzia densa. a. habit x 1 b. calyx at anthesis x 10 c. calyx in fruit x 5 d. corolla x 10 e. dissected flower x 10 f. stamen x kO g. style disc and young nutlets x 10 h. nutlet, inner face and profile.x 5 169 Fig. 31. 170 subequally five-lobed corolla, style lobes with globular swellings at the tips and tuber^ulate-rugose nutlets. P. pamirensis was later recognised by Fedschenko ( 1908 ) as being based on a specimen of Elsholtzia densa. Although mistaking the identity of the plant, Briquet thought it sufficiently distinct to warrant its recognition as a new genus. 6.4.5. Relocated OTU's. A number of species of Elsholtzia appear to be somewhat divorced from the bulk of the genus, especially in the single-linkage analysis ( fig. 27 ) and the simplified similarity matrix ( appendix ^ ). In almost every case the data were obtained from a single specimen and the resulting distortion appears to have affected the placing of some species. However, most can be accurately placed by referring to the nearest neighbours list ( appendix 7 ). 6.4.6. E. beddomei ( OTU 129 ) and E. kachinensis ( OTU 12 ). These species are both shown as monotypic genera in the single- linkage analysis ( fig. 27 )• Although sharing few high similarities with other species of Elsholtzia there is no doubt as to where their phenetic affinities lie. Reference to the nearest neighbours list ( appendix 7 ) and the simplified similarity matrix ( appendix ^ ) confirms that E. beddomei belongs to section Aphanochilus sensu stricto and E. kachinensis to section Elsholtzia sensu mihi. 6.4.7. E. penduliflora ( OTU 2$ ). This species,t00,appears as a monotypic genus in the single- 171 linkage analysis ( fig. 27 ) and the simplified similarity matrix ( appendix 4 ) also shows it to be isolated from other OTU's. It is rather surprising to find Pogostemon brevicorollus ( OTU 135 ) as the nearest neighbour to Elsholtzia penduliflora ( see appendix 7 ). However the data set for Pogostemon brevicorollus is incomplete and the similarity between it and Elsholtzia penduliflora may be considered as dubious. The similarity values shared by Elsholtzia penduliflora with its remaining four nearest neighbours, E. elata ( OTU 35 )i and E. blanda ( OTU 38 ), Pogostemon travancoricus ( OTU 81 ) and P. amaranthoides ( OTU 69 ) give no conclusive pointer to where the phenetic affinities of Elsholtzia penduliflora lie. However both the principal co-ordinates ( ®ee figs 14-22 ) and the clustering to maximise WGMS analyses ( appendices 8 & 9 ) place it in Elsholtzia section Aphanochilus sensu stricto, and it seems best placed here. 6.4.8. E. hunanensis ( OTU 24 ), E. flava ( OTU 28 ) and E. fruticosa ( OTU 29 ). In the single-linkage dendrogram ( fig. 27 ), E. hunanensis, E. fla^a and E. fruticosa are not referable to any section of Elsholtzia. As witli the previous species both the nearest neighbours list ( appendix 7 ) and the simplified similarity matrix ( appendix ^ ) can be used to locate all three species. E. hunanensis belongs to section Elsholtzia sensu mihi, E. fruticasa and E. flava to section Aphanochilus sensu stricto. However E. flava warrants further investigation. In this species the style is not always obviously gynobasic ( see p. 78 ) and the nutlets have a small but laterally displaced attachment-scar ( see fig. 11 and p. 147 ). The bracts are unusually broad for section 172 Aphanochilus sensu stricto and the very large, broadly-ovate leaves and coarse habit help to mark out this species. Should the style and nutlet-attachment characters prove to have greater value than they have been accorded here ( see discussion of these characters in Leucosceptrum, pp. 146-147) the position of E. flava will need to be re-examined. 6.4.9. E. concinna ( OTU 10 ). This species presents a more difficult situation and the evidence from the various analyses is conflicting. In the simplified similarity matrix ( appendix 4 ) and the principal co-ordinates analysis of run 2A ( figs 17-22 ) E. concinna might be included within either section Aphanochilus sensu stricto or within section Elsholtzia sensu mihi. In the clustering to maximise WGMS ( see appendices 8,9' is grouped with species of section Aphanochilus sensu stricto. In the single-linkage dendrogram ( fig. 27 ) it links on at the 86% level and is not referable to any section, although the nearest neighbours list ( appendix 7 ) shows it to be most similar to Dysophylla mairei ( = Elsholtzia yunnanensis, see pp. 173 - 174 ) which is a member of section Aphanochilus. The raw data were obtained from two rather poor specimens and should perhaps be regarded with some suspicion. Vautier ( 1959 )i when describing E. concinna ( fig; 5 ) stated that, although differing in bract and inflorescence characters, E. concinna and E. strobilifera ( OTU 19 ) were similar in general appearance; E. concinna and E. ciliata ( OTU 11 ) differed in leaf and inflorescence features. The clear implication was that E. concinna belonged to section Elsholtzia sensu mihi. E.- concinna certainly shares the characters which' define this 173 section ( see table 23 ) rather than those which define section Aphanochilus sensu stricto, although it in fact resembles E. ciliata more than E. strobilifera. In view of this, I have placed E. concinna in section Elsholtzia sensu mihi but the availability of more study material may necessitate reconsideration of its position. 6.4.10. Leucosceptrum plectranthoideum ( OTU 97 ). This species is placed in Elsholtzia section Aphanochilus sensu stricto in all the analyses except the single-linkage dendrogram ( fig. 27 ) where it links with Elsholtzia but is not referable to any particular section. Originally described by Leveille ( 1915 - 16 ) 'as Buddie ja plectranthoidea it was transferred to Leucosceptrum by Marquand ( 1930 ). Having examined the type I have identified it as a specimen of Elsholtzia fruticosa ( OTU 29 ). E. fruticosa and Leucosceptrum plectranthoideum are not, as might be expected, inseparable, or at least very close, units in the analyses. Once again this seems due to inadequate sample size, only the type of Leucosceptrum plectranthoideum being available. This specimen did not fall completely within the variation range of Elsholtzia fruticosa as represented by the specimens used to generate the averaged data set C see p. 27 ). 6.4.11. Dysophylla mairei ( OTU 102 ), This taxon was described by Leveille ( 1912 ) as a species of Dysophylla. However all of the analyses clearly show it to belong to Elsholtzia and not to Dysophylla. Dunn ( 1915 - 17 ) gave Dysophylla mairei as a synonym of Elsholtzia 174 pilosa and indeed the type appears to be a specimen of E. pilosa ( OTU 41 )• However Dysophylla mairei and Elsholtzia pilosa lie adjacent only in the single-linkage analysis ( fig. 27 ) and E. pilosa does not appear in the list of nearest neighbours for Dysophylla mairei ( appendix 7 )• In the principal co-ordinates analysis of run 2A ( figs 17-22 ) there are several units as close to Dysophylla mairei as is Elsholtzia pilosa, for example E. glanduligera ( OTU 34 ) and E. ochroleuca ( OTU 40 ). Dysophylla mairei could equally well be included in any of these taxa. This throws doubt on Dunn's selection of Elsholtzia pilosa as the appropriate" species in which to include Dysophylla mairei. My analyses show D. mairei to be a separate OTU and not so similar to any other OTU as to warrant conspecificity. Certainly D. mairei is phenetically allied to Elsholtzia pilosa, E. glanduligera, E. ochroleuca and perhaps several other species e.g. E. capitfrligera (OTU 4l) but there is insufficient evidence to justify the inclusion of Dysophylla mairei in any of these species. D. mairei must, therefore, be regarded as a species of Elsholtzia and needs to be formally transferred from Dysophylla to Elsholtzia. The name Elsholtzia mairei was applied by Leveille ( 1915 - 16 ) to another plant ( conspecific with E. rugulosa, OTU 26 J and the combination E. mairei(Lgveill§) mihi would be a later homonym. A new name is required and I have chosen Elsholtzia yunnanensis ( see p. 223 6.4.12. E. integrifolia ( OTU 23 ). All previous authors to have considered E. integrifolia have placed it in Elsholtzia section Aphanochilus. This is based on Bentham's original concept ( 1832 - 36 ) since none of them appears to have seen 175 the type specimen, a sheet only recently relocated by me in the herbarium at the British Museum ( Natural History ). As a supposed member of Elsholtzia section Aphanochilus, E. integrifolia was included in this study, but the analyses show it to be an isolated taxon, sharing an overall similarity of only 84.1$ with its nearest neighbour, E. griffithii ( OTU 33 ) ( see appendix 7 )• In the recent Flora Reipublicae Popularis Sinicae volume 66 ( 1977 ) Wu 8c Huang cite E. integrifolia as a synonym for Schizonepeta tenuifolia ( L. ) Briq. in the tribe Nepetae, apparently on evidence obtained from a photograph. When this work became available to me in 1979i after completion of the computor runs, I compared the type of E. integrifolia with available material and descriptions of Schizonepeta, although unfortunately not of Schizonepeta tenuifolia. Elsholtzia integrifolia does share a greater overall similarity to other species of Schizonepeta than to species of Elsholtzia. I take Wu & Huang ( 1977 ) to be correct in assuming these taxa to be conspecific and therefore exclude Elsholtzia integrifolia from the Pogostemoneae. 6-4.13. E. chinense ( OTU 100 ). This was thought to be a previously undescribed species represented by two specimens in the L6veille herbarium at the Royal Botanic Garden, Edinburgh. Both specimens were collected by d'Argy in the Kiangsu province of China, and were labelled as E. .japonica. Since this name already exists for another species ( E. japonica Miq. ) and to avoid confusion I labelled the OTU as E. chinense. In the simplified similarity matrix ( appendix 4 ) E. chinense shares high similarities with species of both Elsholtzia and Pogostemon. 176 In the clustering to maximise WGMS ( appendices 8, 9 ) and the principal co-ordinates analyses ( figs 14-22 • ) it clusters with species of Elsholtzia. In the single-linkage analysis ( fig. 27 ) it clusters with species of Pogostemon. However, E. chinense lacks certain constant features possessed by species of Elsholtzia and Pogostemon. Species of Pogostemon have an entire upper lip to the corolla, hairy stamen-filaments and unilocular anthers. E. chinense has an emarginate upper lip to the corolla, naked stamen-filaments and bilocular anthers. Species of Elsholtzia have bilocular-fused anthers and a tumescent lobe on the disc. E. chinense has bilocular-free anthers and no lobe on the disc. It also has hairy nutlets while those of species of Elsholtzia and Pogostemon are invariably glabrous. These characters suggest that E. chinense might be misplaced and a brief survey of genera outside the Pogostemoneae confirms that E. chinense is more similar to species of Agastache ( tribe Nepetae ) and, in particular, to A. rugoaus Fish. 8c Mey. with which it may be conspecific. I have •therefore excluded it from the Pogostemoneae. 6.4.14. Distribution. The distribution of Elsholtzia as accepted by other authors was rather striking. Most species are confined to eastern Asia and Malesia with the weedy species, E. ciliata ( OTU 11 ), extending over a wider area as far as central Europe ( where it may be a relict of cultivation ). A single, very isolated species E. aquatica ( OTU 22 ) occurred in southern Malawi. However, the removal of this species to Pogostemon also removes the disjunction in the distribution of the genus. 177 6.4.15. Keiskea. The species of Keiskea in the study form a discrete and homogenous group. This is so in each of the analyses and is well illustrated in the simplified similarity matrices ( figs 12,13 and appendices 4, 5 ) and the clustering to maximise WGMS ( tables 7, 8, 12, 13 )• However both these analyses show species of Keiskea to be considerably more similar to species of Elsholtzia than to any other group. This relationship is also shown by the principal co-ordinates analyses ( figs 14 - 16.) where the species of Keiskea consistently cluster with species of Elsholtzia. In the same analysis of run 2A ( figs 17, 18, 19 ) this link is further refined where the species of Keiskea cluster with species of Elsholtzia section Elsholtzia sensu mihi, although in fig. 18 they separate cleanly to form another group. The single-linkage analysis ( fig. 27 ) emphasises the discreteness of Keiskea as a group, not linking with the Elsholtzia and Pogostemon / Dysophylla groups until the 86$ level. This obvious similarity of species of Keiskea to species of Elsholtzia confirms the view of earlier authors ( e.g. Bentham & Hooker 1876, Ohwi 1965 ) who placed Keiskea next to Elsholtzia. The two genera were said to differ mainly in features of the calyx, that of Keiskea being deeply divided and sometimes described as bilabiate, while that of Elsholtzia is shallowly toothed and not bilabiate. During this study, several other characters which distinguish the genera have been determined. As well as being deeply divided, the calyx in each species of Keiskea ( cf. fig.7)h&a:an annulus of hairs in the throat. No species of Elsholtzia possess such an annulus. Similarly each species of Keiskea cf.fig-.8has a complete annulus of hairs within the corolla. Some species 178 of Elsholtzia e.g. E. rugulosa ( fig 8 ) also possess an annulus within the corolla but these annuli are always incomplete, with the ends of the open circle of hairs extending dorsally under the upper lip. In species of Elsholtzia the number of mature nutlets is always four. Fertilised flowers of all species of Keiskea have four very young nutlets. Keiskea .japonica is the only species in which mature nutlets are known to me. The specimens examined had only one large mature nutlet per flower (fig.11)a character also noted by Ohwi ( 1965 )• However my sample was very small and non-development of one, two or three nutlets is common in Labiatae. It may be that a larger sample will show the normal number of mature nutlets in species of Keiskea to be four. The confluence of the anther locules is an ambivalent character. Species of Keiskea have bilocular anthers in which the locules are free and separated by a short connective. Species of Elsholtzia have bilocular anthers in which the locules are confluent, although still recognisably bilocular ( cf. figs 11,30 ). The single exception to this is E. hunanensis ( OTU 2k ) which has anthers similar to those of species of Keiskea. Thus, although this appears to be a distinguishing feature between the two genera, the exceptional Elsholtzia species show it to be a linking character too. The clear phenetic similarity of Keiskea with Elsholtzia is established by a number of characters. The common characters of a ( sometimes weakly ) bilabiate corolla with an emarginate upper lip and spreading, three-lobed lower lip were recognised by earlier authors ( Bentham & Hooker 1876, Briquet 1897, Ohwi 1965 ). In addition species of both genera have a single tumescent gland on the disc, a character within the tribe possessed only by these two genera ( cf. fig. 10 ). In 179 Keiskea the bracts are similar in shape and texture to those of a number of Elsholtzia species e.g. E. flava ( fig.. 5 ). The few- flowered verticils forming a loose, secund inflorescence in species of Keiskea are also typical of some species of Elsholtzia, particularly those of Elsholtzia section Elsholtzia sensu mihi e.g. E. ciliata ( OTU 11 ). Only one species of Keiskea, K. macrobracteata, is omitted from this study for lack of available material. From the description however, all the comments made above regarding species of Keiskea would apply equally to K. macrobracteata. The description also suggests that it may be the species of Keiskea most similar to species of Elsholtzia section Elshollzia sensu mihi, particularly in respect of the bract characters. Masamune ( 19^0 ) in his original description of this species considered that its conspicuous features distinguished it from other species of Keiskea and divided the genus into two sections, section Macrobracteatae, represented solely by K. macrobracteata, and section Eukeiskea. The omission of K. macrobracteata prevents any assessment of these sections from being made here. 6.4.16. Tetradenia. The three species of Tetradenia, like species of Keiskea, form an obvious group in all analyses. Again like Keiskea, the species of Tetradenia are most similar to species of Elsholtzia. The clustering to maximise WGMS analysis ( see tables 7, 13) suggests a greater similarity to Elsholtzia section Elsholtzia sensu mihi than to Elsholtzia section Aphanochilus sensu stricto but this is not borne out by the principal co-ordinates analyses. In run 2A ( figs 17, 18 ) the species of Tetradenia lie more or less between the Elsholtzia sections. The single-linkage 180 analysis ( fig. 27 ) emphasises the isolation of Tetradenia from other taxa since it does not link on until the 81$ similarity level. In his original description of the genus Bentham ( 1829 ) recognised Tetradenia by the four tumescent lobes on the disc and placed it " immediately before Elsholtzia " . In a later work ( 1832 - 36 ) he expanded this statement considerably: " it ( Tetradenia ) is intermediate between Elsholtzia and Colebrookea; it differs from the first by the calyx and the more regular corolla; from Colebrookea by the calyx not plumose at the maturity of the fruit, by the style less deeply cleft; and from both by the remarkable glands of the ovarium " . Notwithstanding the link with Elsholtzia Bentham ( 1848 ) and other authors ( e.g. Bentham & Hooker 1876 ) grouped Tetradenia with Pogostemon, Dysophylla and Colebrookea; the group was characterised particularly by unilocular anthers or the anther-locules confluent, the corolla tube rarely longer than the calyx, the lobes equal or the anterior longer. Elsholtzia was placed in another group characterised by distant or sometimes confluent anther-locules. The evidence of this study supports the link between Tetradenia and Elsholtzia while refuting that with Colebrookea, Pogostemon and Dysophylla ( although from the principal co-ordinates analyses, figs 14 - 16 , Colebrookea would appear to be more similar to Tetradenia than are either Pogostemon or Dysophylla ). The proximity of Tetradenia and Elsholtzia in all of the analyses except the single-linkage analysis C fig. 27 ) is clear evidence of the phenetic similarity between the genera. This proximity is reflected by a number of characters. Perhaps the most obvious are the shape of the corolla, the shape and texture of the 181 bracts and. the bilocular ( not unilocular ) anthers. Species of both Tetradenia and Elsholtzia have an emarginate upper lobe to the corolla. The corolla in most species of Elsholtzia, as observed by Bentham, is more markedly bilabiate than that of species of Tetradenia but some species of Elsholtzia, e.g. E. myosurus and E. densa, have similar corollas ( see figs 8, 31 )• Species of Pogostemon and Dysophylla have an entire upper lip to the corolla. The link between species of Tetradenia and species of Elsholtzia section Elsholtzia sensu mihi is emphasised by the similarity of the bracts. In species of both groups the brac.ts are orbicular or somewhat broader than long, usually brown ( or bi- coloured ) and membranous. Species of Elsholtzia section Platyelasmeae sensu mihi also have broad, brown or bi-coloured bracts but they are not membranous. Similar bracts are found in Elsholtzia flava ( OTTI 28 ) ( cf. fig. 5 )• Like species of Elsholtzia section Elsholtzia sensu mihi and Elsholtzia section Platyelasmeae sensu mihi, species of Tetradenia lack bracteoles. This contrasts markedly with species of Pogostemon, Dysophylla and Elsholtzia section Stenelasmeae sensu stricto in which all species possess bracteoles. The anthers of species of Tetradenia are bilocular and confluent through fusion, not unilocular as given by Bentham ( 1848 ) and Bentham & Hooker ( 1876 ) Two other characters help to link species of Tetradenia with species of Elsholtzia while divorcing them from species of Pogostemon, Dysophylla and Colebrookea. Like species of Elsholtzia, species of Tetradenia have glabrous anther-filaments; all species of Pogostemon and Dysophylla have hairy anther-filaments. Again like species of Elsholtzia, species of Tetradenia possess tumescent glands on the disc; no species of Pogostemon, Dysophylla or Colebrookea has such glands ( cf. fig. 10 ). 182 These tumescent glands provide one of two unique features of Tetradenia as a genus. Species of Tetradenia have four bright red glands encircling the disc. The young nutlets rise above the glands, but as they mature the glands enlarge greatly, finally overtopping the nutlets. The second unique feature of species of Tetradenia is a calyx character. The upper tooth of the calyx is very broad, much broader than, and overlapping, the teeth on either side of it. Similar calyces are unknown in other species of the Pogostemoneae ( cf. fig. 7 )• The corolla in species of Tetradenia has a complete annulus of hairs within its throat. Although some species of Elsholtzia e.g. E. rugulosa ( OTU 26 ) have annuli, they are never complete circles of hairs. Other than in species of Tetradenia, complete annuli are found only in species of Keiskea and Comanthosphace, ( cf. fig. 8 ) 6.5. SUBTRIBE EURY50LENINEAE. 6.5*1- Eurysolen. The taxonomic position of Eurysolen is open to conjecture. Prain ( 1898, 1901 ) described the only species, E. gracilis, and tentatively placed it near Gomphostemma in his tribe Prasieae citing only the character of anthers one-celled from a very early stage as supporting evidence for his choice. Briquet ( see Prain 1901 ) suggested that Eurysolen might be placed in the Prasieae or the Ajugoideae. However the lack of mature fruits prevented the exact determination of its position. Later authors were similarly hampered by lack of fruiting material. Kudo ( 1929 ) placed Eurysolen in the Prasiae; Mukerjee ( 19^0 ) placed Eurysolen in his tribe Ajugoideae. Wu ( 1959 ) placed Eurysolen in the 183 Ajugoideae but noted an apparent relationship with Pogostemon. Unfortunately he did not amplify this in any way. Chermisirivathana ( 1963 ) examined a number of fruiting specimens and described the mature fruits as dry and with a small basal attachment. On this evidence she placed it with Pogostemon, Dysophylla, Elsholtzia and Colebrookea in her subfamily Stachyoideae. This was supported by Keng ( 1969? 1978 ) who considered that, in general habit, inflorescence and flower structure Eurysolen was identical with another member of the Stachyoideae, Achyrospermum. My study is restricted to the tribe Pogostemoneae and the exact position of Eurysolen cannot be ascertained without reference to the other tribes to which it might belong. However if Chermisirivathana1 s and Kengfs reasons for rejecting the Ajugoideae and Prasieae are accepted, Eurysolen would seem well-placed with the other genera of the Pogostemoneae. My own examination of mature fruits of Eurysolen ( fig* 11 ) confirms Chermisirivathana^ observations; they are dry and not fleshy as in the Prasieae and the attachment scar is small and basal, not large and lateral as in the Ajugoideae. In general habit and inflorescence Eurysolen gracilis resembles some species of Pogostemon, e.g. P. wightii ( OTU 89 ) and in floral structure resembles species of Elsholtzia and Rostrinucula ( fig. 8 ). My analyses give conflicting results for the affinities of Eurysolen within the Pogostemoneae. In the simplified similarity matrix ( appendix 4 ) Eurysolen gracilis ( OTU 126 ) is not very similar to any other OTU, although it shares its highest similarities with species of Elsholtzia section Aphanochilus sensu stricto. This is confirmed by by the nearest neighbours list ( appendix 7 ) which shows the nearest 184 neighbour to be Elsholtzia yunnanensis mihi ( OTU 102 ) ( = Dysophylla mairei, see p. 174) with the very low similarity of 81.5$. The single- linkage analysis ( fig. 27 ) also shows Eurysolen gracilis to be an OTU of rather remote affinity with other members of the tribe. The principal co-ordinates ( figs 14 - 19) and clustering to maximise WGMS analyses ( tables 6-8, 11-13) clearly and consistently show Eurysolen gracilis as a member of the same group as species of Elsholtzia section Aphanochilus sensu stricto. They are not separable even in the principal co-ordinates analysis of run 2A ( figs 17 - 19 )• These results should not perhaps be taken at face value. In the principal co-ordinates analyses the inclusion of Eurysolen gracilis in the cluster of Elsholtzia species is almost certainly due to one of the flaws in this method. The axes used to produce the two-dimensional plots are those corresponding to the largest eigenvalues, i.e. those axes in the direction of maximum variance ( see p. 52 ). Only the first of four axes are normally used and this may be insufficient to split OTU's apart. The presence of Eurysolen gracilis in the same group as species of Elsholtzia in the clustering to maximise WGMS is probably due to the " rag-bag " effect. That is, since every unit must be placed in a group and the pre-selected number of groups has already been formed at higher levels of similarity, Eurysolen gracilis has been placed with its nearest neighbour Elsholtzia yunnanensis ( OTU 102 ). That the similarity of Eurysolen gracilis to Elsholtzia yunnanensis is very low ( 81.2$ ) does not affect the inclusion of these two OTU's in the same group; Eurysolen gracilis only appears to be a part of the Elsholtzia cluster. The single-linkage analysis ( fig. 27 ) suggests that this inclusion is unlikely and the nearest neighbours list confirms that they are 185 Fig. 32 • Eurysolen gracilis. a. habit x 1 b. calyx x 5 c. corolla x 5 d. dissected flower x 5 e. stamen x 10 f. annular hairs at the base of a stamen filament x kO 743 Fig. 32 187 separate taxa. The pollen data ( table 2 ) lends further support to the separation of Eurysolen and Elsholtzia ( see p. 142 )• Clearly however, Eurysolen gracilis is phenetically most closely related to species of Elsholtzia. Eurysolen gracilis does share some characters with other members of the tribe, most notably an invagination of the corolla and hairy bosses at the base of the stamen filaments as in species of Rostrinucula; unilocular anthers as in species of Pogostemon and the Comanthosphacineae. It is recognised within the Pogostemoneae by a combination of these characters, the absence of a tumescent gland on the disc, the emarginate upper lip of the corolla and the single distinctive feature of the corolla tube gibbous towards the base ( see fig. 32 ). 6.6. SUBTRIBE POGOSTEMONINEAE. 6.6.1. Pogostemon and Dysophylla. Although usually treated as separate genera, Pogostemon and Dysophylla have been considered as congeneric taxa by several authors, first by Hasskarl ( 1842 ), later by Miquel ( 1856 ) and Kuntze ( 1891 )• Bentham ( 1870 ) retained the two as separate genera and commented that if they were to be regarded as a single genus, Pogostemon, Dysophylla would still form a distinct section. Keng's ( 1978 ) arrangement in which species of Dysophylla are confined to Pogostemon section Eusteralis agrees with this idea of Bentham. El-Gazzar & Watson ( 1967 ) produced a modified treatment for the two genera. On the basis of a comparison of six characters, leaf shape, leaf arrangement and indumentum, petiole length, calyx inclusions and presence of stem aerenchyma in twenty-two species of Pogostemon 188 ( approximately 37% of the genus ) and sixteen species of Dysophylla ( approximately k7% of the genus ) they transferred the four species of Dysophylla section Qppositifoliae to Pogostemon. Although the species sample was rather restricted, their evidence and arguments were clear and persuasive. Their emended genera were characterised as follows :- Dysophylla. Leaves verticillate, 3 - 10 in a whorl, linear, sessile and usually glabrous. Corolla subequally quadrifid. Pogostemon. Leaves opposite, ovate or narrowly ovate, petiolate, usually more.or less hairy or tomentose. Corolla usually subbilabiate, upper lip trifid, lower entire. With these differing treatments of Pogostemon in mind, the results of my study can be examined more closely. Each of the analyses shows clear similarities between species of Dysophylla and Pogostemon. In the single-linkage analysis ( fig. 27 ) most species of Pogostemon and Dysophylla link on at the rather high level of 91$ similarity. They form three groups, two containing species of Pogostemon and the other containing species of Dysophylla. The first group of Pogostemon species is more similar to the species of Dysophylla, linking at the 92% level of similarity, than it is to the other group of Pogostemon species which links at the 91% similarity level. In the simplified similarity matrix ( appendix 4 ) the limits of the groups formed by species of Dysophylla and Pogostemon are rather blurred due to a number of OTU's which share high similarities with members of more than one group. These OTU's are shown in table 24 , and include all species of Dysophylla section Oppositifoliae, several species of Dysophylla section Verticillatae 189 Table 24 . Species which share high similarities with Dysophylla and Pogostemon in two analyses. OTU simplified similarity clustering to maximise matrix run 1 WGMS run 2B 43 D. trinervis + D. glabrata + 55 D. cruciata + 56 D. quadrifolia + 57 D. auricularia + + 58 D. salicilifolia + + 59 D. rugosa + + 60 D. myosuroides + + 70 P. formosanus + 71 P. menthoides + + 74 P. micagensis + + 75 P* mutamba + + 91 P. strigosus + + 92 P. hispidus + 104 D. andersoni + 105 D. koehneana + 106 D. gracilis + • 113 P. hirsutus + + 114 P. rupestris + + 120 P. nilagiricus + + 135 P« brevicorollus + 190 Fig. 33 • Dysophylla glabrata. a. habit x 1 b. calyx x 20 c. corolla x 20 d. dissected flower x 20 e. stamen x 40 f. nutlet, inner face and profile x 20 Fig. 33 . 192 and several species of Pogostemon. In the clustering to maximise WGMS analysis of ( tables 6, 7 ) the groups of species of Pogostemon and Dysophylla have relatively high WGMS values suggesting some degree of integrity for the groups. However, they also share high BGMS values with each other, suggesting that the groups are not dissimilar. The clustering to maximise WGMS analysis of run 2B is rather more revealing, particularly in the k group scheme ( table 15)- Again the WGMS and BGMS values are conflicting, indicating discrete but not easily separable groups. In .this scheme the focus of interest is group which contains all the species of Dysophylla section Oppositifoliae, several species of Dysophylla section Verticillatae and several species of Pogostemon ( appendix 10 ) and which shares high and roughly equal BGMS values with each of the other groups. As might be expected the majority of species in group k are ones whose similarities cross group boundaries in the shaded similarity matrix ( see appendix k and table 2k ). In the principal co-ordinates analysis ( figs 17 — 19 ) species of Pogostemon and Dysophylla form a single cluster but with most species of Pogostemon concentrated at one end of the cluster and most species of Dysophylla concentrated at the other end. This distribution of species is more clearly seen in the principal co-ordinates analysis of run 2B ( figs 23-25 ) which shows that species of Dysophylla at the end of the cluster are only from section Verticillatae. The middle of the cluster is occupied by the OTU's listed in table 2k , i.e. species of Dysophylla section Oppositifoliae, some species of Dysophylla section Verticillatae and some species of Pogostemon ( see figs 3k - 6). 193 Figs 34, 351 36 • Two dimensional plots of the principal co-ordinates analyses run 2B showing the positions of Dysophylla linearis, D. falcata, D. szemacensis and those species of Pogostemon and Dysophylla which are listed in table 2k . Key. species listed in table 2k and D. linearis, D. falcata, D. szemacensis. • other species of Dysophylla and Pogostemon. O 194 Fig. 34 . vector 1 cr "bo*J- k M "O^O" 1 O" u o o" O" o o*- "bo'*3 cr •* ~o8$cT d'o* dV o oW •5W * 00* o* on o* o* >* O71 On o« o* Omon vector 2 195 Fig.36 . vector 1 d* o* •s'H CP O1* 0®t 0r 5 CPq* O ' O O" b •b bo" o" o1" o1* v* 0" 0*Vei o** cT Q^O* w n 1 o qao* bo" o* QUfc O Cn Ow vector 3 196 Fig. 36 . vector 1 O® o<* 6 o* O"1 Oo«i O' kS o41 o o" 4 o* k'lS o,c* o11 O" Q.M •V u O"1 o o owt O-s O" vector 3 197 El-Gazzar & Watson's sample of species of Dysophylla section Verticillatae included only two species with hairy leaves, a character which the authors dismissed as non-significant. I have examined a number of similarly hairy species unavailable to El-Gazzar & Watson. With the exception of Dysophylla linearis ( OTU 3k ), D. falcata ( OTU 137 ) and D. szemacensis ( OTU 138 ) ( see figs 3^-36 ), these hairy species may all be found in table 2k , among those species selected from the simplified similarity matrix ( appendix k ) as sharing high similarities with species of both Dysophylla and Pogostemon. If all the species listed in table 2k and indicated in figs 3k - 36 are described in terms of the morphological features which characterise Dysophylla and Pogostemon sensu El-Gazzar & Watson, they form a transition series between the extremes of the cluster. The stages in the series are as follows :- Species with leaves in whorls of three or more, sessile, linear, truncate at the base, glabrous, corolla with upper lip equalling or shorter than lower; this describes Dysophylla sensu stricto; Dysophylla stellata ( OTU 53 ) is a typical species. Species with leaves in whorls of four, sessile, linear, truncate at the base, sparsely hairy, corolla with upper lip equalling lower; Dysophylla linearis ( OTU 3k ). Species with leaves in whorls of three or more, sessile, linear, truncate at the base, hairy, corolla with upper lip equalling lower; Dysophylla cruciata ( OTU 55 )* Dysophylla koehneana < . ( OTU 105 ), Dysophylla gracilis ( OTU 106 ) and Dysophylla szemacensis ( OTU 138 )• Species with leaves in whorls of three or four, very shortly petiolate, 198 linear, attenuate at the base, densely hairy, corolla with upper lip equalling or shorter than lower; Dysophylla falcata ( OTU 137 Dysophylla quadrifolia ( OTU 36 ). Several specimens of Dysophylla quadrifolia in the herbarium at the Royal Botanic Garden, Edinburgh have paired leaves instead of the more usual whorls of three. The isotype of Dysophylla falcata ( the only specimen examined ) appears to have leaves in whorls of three, changing to pairs of leaves at the two uppermost nodes. Species with leaves in whorls of three, sessile, broadly ovate to orbicular, cuneate at the base, sparsely hairy, corolla with upper lip equalling lower; Dysophylla tirinervis ( OTU k?> ). Species with leaves in pairs, sessile, ovate, truncate at the base, densely hairy, corolla with upper lip equalling lower; Dysophylla andersoni ( OTU 104 ). Species with leaves in pairs, petiolate, linear to narrowly ovate, sparsely hairy, cuneate at the base, corolla with upper lip equalling lower; Dysophylla salicifolia ( OTU 58 ). Species with leaves in pairs, shortly petiolate, ovate, cuneate at the base, glabrous, corolla with upper lip equalling lower; Dysophylla glabrata ( OTU kk ). Species with leaves in pairs, shortly petiolate, ovate, cuneate at the base, hairy ( sometimes densely so ), corolla with upper lip equalling lower; Dysophylla rugosa ( OTU 59 )> Dysophylla myosuroides ( OTU 60 ), Dysophylla auricularia ( OTU 57 ). Species with leaves in pairs, very shortly petiolate, ovate, cuneate at the base, sparsely hairy, corolla with upper lip longer than lower; Pogostemon strigosus ( OTU 91 ). This species bears a striking 199 resemblance to Dysophylla auricularia. Species with leaves in pairs, shortly petiolate to subsessile, ovate, cuneate at the base, sparsely hairy to subglabrous, corolla with upper lip longer than lower; Pogostemon mutamba ( OTU 751 fig*37 )i Pogostemon micangensis ( OTU 7*t Species with leaves in pairs, petiolate, linear, cuneate at the base, densely hairy, corolla with upper lip equalling lower; Pogostemon nilagiricus ( OTU 120 ). Species with leaves in pairs, petiolate, ovate to orbicular, cuneate, attenuate or rounded at the base, hairy, corolla with upper lip equalling or longer than lower; Pogostemon sensu stricto, including Pogostemon menthoides ( OTU 71 )? Pogostemon hirsutus ( OTU 113 ) and Pogostemon rupestris ( OTU 114 ). From the simplified similarity matrix ( appendix k ) Pogostemon formosanus ( OTU 70 ) and Pogostemon hispidus ( OTU 92 ) should also be regarded as members of the transition series. However their positions in the principal co-ordinates plots ( figs 3^-36 ) suggest that they should be regarded as members of Pogostemon sensu stricto and using the characters given above they are undistinguishable. The simplified similarity matrix ( appendix k ) indicates that Pogostemon brevicorollus ( OTU 135 ) should also be regarded as a member of the transition series. Much of the data for this species is missing and it may be that it does not, in fact, share high similarities with many Dysophylla species. The type specimen is very similar to Pogostemon amaranthoides ( OTU 69 ) which is the only species to appear as an intermediate species in the centre of the principal co-ordinates cluster but not in group 4 of the clustering to maximise WGMS of run 2B 200 ( table 15 ) or the simplified similarity matrix ( appendix 4 ). The position of P.. amaranthoides in the principal co-ordinates analysis is rather anomalous, as it appears to be a somewhat unusual member of Pogostemon sensu stricto. The other end of the transition series, Pogostemon sensu stricto, is slightly confused, the various categories less clear cut and not matching the left to right order of species in the principal co-ordinates plots as well as the categories containing species of Dysophylla. Nevertheless a logical progression is discernable and, taken as a whole, the transition series appears acceptable. The results of the analyses clearly show that Dysophylla and Pogostemon cannot be retained as separate genera. The consequences of this are as follows. Hasskarl's ( 1842 ) proposal to unite Pogostemon and Dysophylla must be accepted and El-Gazzar's & Watson's ( 1967 ) finding in favour of maintaining the two genera rejected. Although unconvincing in maintaining Dysophylla and Pogostemon as separate genera, El-Gazzar 8c Watson's thesis that Dysophylla section Oppositifoliae is much closer to Pogostemon sensu stricto than to Dysophylla sensu stricto is correct. Keng ( 1978 ) has produced a classification which unites Pogostemon and Dysophylla while incorporating the views of El-Gazzar 8e Watson. On' the separation of Pogostemon and Dysophylla he stated that " the phyllotaxis is insufficient taxonomically for a ( division at the ) generic level " and accepted Pogostemon in the widest sense, including Dysophylla. However he made use of phyllotaxis to define two new sections. He accepted the joining of the opposite-leaved species of Dysophylla with Pogostemon sensu stricto to form section Pogostemon and placed 201 the verticillate-leaved species of Dysophylla in section Eusteralis. Keng's conclusions were based only on studies of Malesian plants ( ten species ) but his eminently sensible arrangement fits the data from my wider sample too, and is followed here ( see pp. 225 - 2^0 ). With the exception of Dysophylla falcata and D. quadrifolia the species are easily divided between the sections. However to accomodate these exceptional species slight modification of Keng-'s distinguishing characters of the sections is required. Pogostemon section Pogostemon. All leaves in opposite pairs, usually petiolate, the corolla with the upper lip equalling or longer than the lower. Pogostemon section Eusteralis. At least some of the leaves in whorls of three or more, usually sessile, the corolla with the upper lip equalling or shorter than the lower. Supporting evidence for this treatment of Pogostemon and Dysophylla comes from work by Cook ( 1978 ) who noted the occurence of a complex of characters including erect, unbranched stems with simple, elongate leaves borne in whorls which he called the " Hippuris syndrome This complex is found in a number of unrelated flowering plant groups and Cook pointed out that it is only ever found in aquatic or amphibious plants and appears to be developed purely as a response to habitat. Among the plants named by Cook as exhibiting the Hippuris syndrome were species of Pogostemon section Eusteralis. These species do indeed show all the required characters, including absence of hairs from most parts of the plants, stems which are creeping below, erect above and small flowers with very short pedicels. The characters used by earlier authors ( e.g. El-Gazzar & Watson 19&7 ) the maintainance of Dysophylla as a genus 202 Fig.38 . Distribution of Pogostemonsens ulato. a. habit x 1 b. calyx x 10 c. corolla x 5 d. dissected flower x 5 e. stamen x 10 f. nutlet, inner face and profile x 10 203 Fig. 37 . 204 distinct from Pogostemon are precisely those of the Hippuris syndrome. All species of Pogostemon section Eusteralis are plants of wet habitats while many (though by no means all ) species of Pogostemon section Pogostemon are plants of stony, dry grounds or forest margins. Some species of section Eusteralis e.g. P. falcatus and P. cfuadrifolius inhabit the dryer parts of wet habitats or areas prone to drying out. It is noticeable that these species sometimes possess characters which are typical of species of section Pogostemon e.g. toothed, hairy, petiolate leaves borne in opposite pairs. Similarly two species of section Pogostemon, P. mutamba ( OTU 75 ) and P. micangengis ( CfTU 74 ), which grow along watercourses in southern tropical Africa resemble species of section Eusteralis in their general habit, branching and very shortly petiolate leaves. Both species are normally hairy but the type specimen of P.mican.gensis, which according to the field notes was " an aquatic herb in the marshes of the river Micango n is virtually glabrous. This evidence supports the union of Pogostemon and Dysophylla, many of the characters formerly separating the species being ecological adaptations. 6.6.2. Subgeneric divisions. Bentham ( 1832 - 36 ) divided species of Pogostemon sensu stricto into § Paniculatae and § Racemosae on the basis of inflorescence structure. Briquet ( 1897 ) adopted Bentham's divisions, expanded the characters on which they were based and further subdivided each group. His characters for these groups ( table 25 ) do not contrast*, for example § Paniculatae has linear-subulate bracts while § Racemosae 205 Table 25. Names and diagnoses of Briquet's subdivisions of Pogostemon. §1. Racemosae Benth. Spicastra einfach, unterbrochen. Bracteen lineal-pfriemlich, kiirzer als der Kelch. A. Glabriuscula Briq. Stb. mit nachten oder fast kahlen Stf. B# Barbata Briq. Stb. mit deutlich dichtbart'igen Stf. §2. Paniculatae Benth. Spicastra rispig verzweigt. Stb. uberall mit bartig behaarten Stf. A. Scheinwirtel meistens entfernt, in unterbrochenen Spicastris. B. Scheinwirtel meistens in dicken oder Spicastris ( vergl. oben P. suave ). 206 Table 26. Names and diagnoses of Briquet's subdivisions of Dysophylla. 1. Sect. 1. Rhabdocalicinae Briq. Kelchrohre cylindrisch und stielrund oder sehr undeutlich, stumpf 5eckig.BB. gegen-oder quirl- standig, gezahnt oder ganzrandig. § 1. Oppositifoliae Benth. B. gegenstandig A. Ausdauernde Arten, mit meistens holziger entwickelter Wurzel oder unteriridiseheh Stengel. B. Einjahrige Art, abstehend behaart, mit sitzenden oder kurz gestielten, eilanglichen, gesagten B. behaarten Spicastris und 3©ckigen, zur Fruchtzeit nach innen gebogenen Kelchzahnen. § 2. Verticillatae Benth. B. in Quirlen zu 3 oder k ( selten bis 10 ). Sect. 2. Goniocalicinae Briq. Kelchrdre 5kantig, mit vorspringenden Kanten. Ijahrige Arten mit quirlstandigen, sitzenden, ganz- randigen B. 207 has stamens which are hairy all over. If the characters defining each group are scored for the species of the other group only one character remains valid for defining them, i.e. spikes paniculately branched versus spikes simple. In my view this character is more accurately scored as verticels not secund versus verticels sub-secund ( see character 62 in table k ) and a division of the genus based on this character closely matches that given by Bentham ( 1832 - 36 ) and Briquet ( 1897 )• Both the simplified similarity matrix run 2B ( appendix 6 ) and the clustering to maximise WGMS run 2B ( tables 14, 15 ) indicate the presence of two groups which roughly correspond to Bentham's and Briquet's groups. However, as stated on p. 91 there is sufficient overlap to prevent clear definition of these groups, nor do they appear in either of the other analyses. It is impossible to justify the maintainance of such vague sub- generic groups on the basis of this single character. Briquet's characters for defining the subdivisions A and B in each group ( table 25 ) do contrast but are misapplied. His Glabriuscula and Barbata in § Racemosae are distinguished by the stamens naked or hairy respectively. All species of Pogostemon have hairy stamens, although some species, e.g. P. travancoricus ( OTU 81 ) ( see fig. 9 )i bear hairs on the lower portions of the filaments which are not clearly visible from examination of an undissected flower. However, even if Briquet's character is substituted by the one ' filaments hairy in the lower half versus filaments hairy in the upper half ' no useful division of the species can be made since, for example, P. rotundatus ( OTU 90 ) with filaments hairy in the lower half would be placed in a separate group from P. mollis ( OTU 80 ) with filaments hairy in the 208 upper half eventhough they share a very high overall similarity and are each other's nearest neighbour ( see appendix 7 )• Similarly the inflorescence character used by Briquet to divide A and B in § Paniculatae ( table 25 ) does not provide a clear or useful division of the species and I have discarded his groups- Briquet also divided Dysophylla into two sections, section Rhabdocalicinae and section Goniocalicinae ( table 26). Here too, there is only one contrasting character. Members of both sections have verticillate, sessile and entire leaves and may be annual, leaving only the character calyx-tube terete versus calyx-tube strongly five-angled. The opposite-leaved species of his section Rhabdocalicinae are those which form Bentham's section Oppositifoliae and are removed to Pogostemon sensu stricto. As with Pogostemon sensu stricto the analyses show no evidence of groupings which match those of Briquet and a single character seems insufficient to support a division at sectional or even subsectional level. I have discarded these sections also. 6.6.3. Relocated OTU's. No species have been removed from Pogostemon and excepting . species of Dysophylla ( for D. mairei see pp. 173 - 174 ) the analyses indicate only one species which should be included here. 6.6.4. Elsholtzia aquatica ( OTU 22 ). All the analyses group this OTU with species of Pogostemon sensu lato and examination of the characters readily shows the inclusion of Elsholtzia aquatica in Elsholtzia to be erroneous. It has linear, 209 Fig. 38 . Distribution of Pogostemon sensu lato. 210 sessile, verticillate leaves, an inflorescence of numerous small flowers in crowded whorls, a weakly bilabiate corolla with an entire upper lip which is shorter than the lower lip, stamens with unilocular anthers and hairy filaments and no tumescent gland on the disc. None of these characters is shared by any species of Elsholtzia but they all distinguish species of Pogostemon sensu lato and in particular species of P. section. Euster&Iis. Elsholtzia aquatica must, therefore, be transferred to this section ( see p. 229). 6.6.5- Distribution. The genus Pogostemon has a disjunct distribution ( fig. 38 ); most species are confined to the Indo-Chinese and Malesian regions and Japan with P.stellatus extending as far as northern Australia but three species, P. mutamba ( OTU 73 )i P.micangensis ( OTU 74 ) and P. aquaticus ( OTU 22 ) are confined to southern tropical Africa. There is no evidence for any species of Pogostemon occuring in the intervening areas and the nearest members of the tribe are the three species of Tetradenia which are found in Madegascar. The analyses do not separate the African species from the Asian and Australian species in any way and there is no morphological evidence to help explain the distribution. 6.7. SUBTRIBE COLEBROOKINEAE. 6.7.1. Colebrookea. This is the most isolated of all the genera within the Pogostemoneae and has no strong affinities with any of the other taxa. This can be seen from the simplified similarity matrix ( appendix 4 ) and 211 particularly from the single-linkage analysis ( fig. 27 ). In the principal co-ordinates analysis ( figs 14-16 ) the two species of Colebrookea occupy a central position in each of the plots and are equally distant from all other groups. In the clustering to maximise WGMS analysis ( table 8 ) Colebrookea shares very low BGMS values with all other groups but is somewhat more similar to Elsholtzia section Aphanochilus sensu stricto and Pogostemon pro parte than to other taxa. This is confirmed by the nearest neighbours list ( appendix 7 ) in which only species of Pogostemon and Elsholtzia section Aphanochilus sensu stricto are recorded as nearest neighbours of Colebrookea oppositifolia ( OTU 9 ) and C. ternifolia ( OTU 8 ). Previous authors have neither discussed nor commented on the affinities and relationships of Colebrookea beyond placing it next to Pogostemon ( see Bentham 1832 - 36, Bentham & Hooker 1876, Hooker 1885, Briquet 1897, Kudo 1929, Wu 1977 ) with which it shares the characters of unilocular anthers and an equal disc. In so placing Colebrookea these authors have given undue emphasis to the rather tenuous relationship between it and Pogostemon. While my analyses confirm this they also show that Colebrookea is not significantly more similar to species of Pogostemon than it is to species of Elsholtzia. Grouping Colebrookea with Pogostemon has also served to obscure the separation of Colebrookea which j.s so apparent in my results. Bentham ( 1832 - 36 ) believed Tetradenia to be an intermediate genus between Colebrookea and Elsholtzia but I have found no supporting evidence for this. As a genus Colebrookea is easily recognised by the distinctive features of the fruiting calyx, the calyx teeth becoming greatly elongated and plumose ( fig. 7). The single, hairy nutlet ( cf. fig. 11 ) does 212 not fall from the calyx when ripe, but remains firmly attached to the disc, fruit and calyx acting as a diaspore. The plumose calyx teeth give the infructescence a characteristic fluffy appearance. Colebrookea may also be recognised by a combination of characters which are found in other genera within the Pogostemoneae. Like Pogostemon sensu lato it has an equal disc and unilocular anthers but the filaments are naked ( cf. fig. 9 ) and the upper lip of the corolla ( cf. fig. 8 ) is emarginate. 213 7. GENERAL DISCUSSION. In retrospect the execution of this study raises a number of points. One obvious shortcoming is the incompleteness of the coded data matrix, due to lack of sufficient material of some species or, when sufficient gatherings are available, lack of certain required parts, such as ripe fruits. Furthermore, almost all of the study was carried out on herbarium material and live material would have been useful in helping to obtain and interpret data. When scoring the data it was unsatisfactory to have to sample each species and use averaged data for each OTU. It does seem that a number of apparent discrepancies in the results of the analyses may be directly attributable to this problem as shown perhaps in the relative positions of Elsholtzia fruticosa and Leucosceptrum plectranthoideum. Ideally one would like to score each gathering as separate OTU's and thus be able to account for intra- as well as inter-specific variation but the number of OTU's which would be involved in such a study obviates this approach. Minima and maxima for each of the quantitative characters might have proved useful and, in retrospect, a more definite scaling for the characters would have helped to reduce distortions of the kind found with, for example, Elsholtzia densa and E. eriostachya. Although the sample of species is large ( some 70% of the tribe ) some taxa such as the Chinese species, in particular, are rather neglected. Many of these have only been described quite recently and, while a number may prove to be conspecific with earlier species described by western authors, the study would have been greatly improved by their inclusion. Some of the problems outlined in the introduction such as the 214 affinities and position of Leucosceptrum canum ( see pp. 144-148 ) cannot be fully resolved without reference to taxa from other tribes. In this respect my sample of species was too restricted to enable solutions to be formulated. To have expanded the study to include all the appropriate taxa would have entailed too great a volume of work for such a project. Similarly the range of analyses used was not sufficient to resolve all of the problems, for example, the exact affinities of Elsholtzia concinna ( pp.172-173) and the most suitable choice of subtribes ( see pp. 134 - 144). As outlined by Sneath te Sokal ( 1975 ) numerical phenetic methods use overall similarity as the basis for establishing relationships. There have been many objections to the philosophy of phenetics ( e.g. Farris 1979 ) and there are faults in the choice of characters, choice of similarity matrices, clustering methods and so on- Particular ly relevant to this study is the choice of characters. As pointed out earlier ( see p> 33 )* overall similarity cannot be achieved since any list of characters to be used in analysis is restricted and must by definition be a selection. A cynical comment would be to say that a study of the Pogostemoneae could have easily been carried out using traditional, non-numerical methods. The principal goals of numerical phenetics are objectivity, repeatability and stability ( Sneath & Sokal 1973 )• In dealing with objectivity these methods have subjective elements at all stages of the procedure and do not produce objective results and therefore they do not give " better " taxonomy for the group than would other methods. However, the large amount of data involved in this study would have been much more difficult to handle without the use of a computer. Numerical 215 methods have allowed many more comparisons to be made than by traditional methods and have therefore helped to highlight various features, such as the relationships of the Eurysolenineae to the other subtribes,which might otherwise not have been elucidated. Despite their limitations ( see pp.^9 - 5*0, the analyses used here are flexible and statistically robust and well suited for taxonomic investigations. In general, the ways in which they function, the data required and the manipulation and interpretation of the results obtained are relatively simple and easy to understand. With the range of techniques and analyses now available ( see Sneath & Sokal 1973 ), criticism of this thesis could perhaps be levelled at the lack of sophistication in the analyses adopted here. Certainly other methods for optimising taxonomic resolution such as a weighting of characters in the initial character selection and more complex measures of best fit offer interesting and tempting alternatives. Furthermore such techniques may provide further insights to the problems of this group, but the analyses I have used do give good resolutions and I believe that my results provide a useful contribution to the taxonomy of the Pogostemoneae. Solutions have been offered to a number of the relationship problems within this tribe and where the analyses proved inadequate for this they have at least enabled the remaining problems to be adequately defined. 216 8. TAXONOMIC CONSPECTUS 8.1. KEY TO GENERA AND SECTIONS. 1a. Anthers bilocular; disc with one or more tumescent lobes 2 1b. Anthers unilocular; disc without tumescent lobes ....• •••••• 4 2a. Disc with four large, bright-red, tumescent lobes; upper tooth of calyx broad and over- lapping the lateral teeth .••••••••• Tetradenia 2b. Disc with one tumescent lobe; upper tooth of calyx not overlapping the lateral teeth ... 3 3a. Calyx with a well-defined annulus of white / hairs at the mouth of the tube; mature nutlets one Keiskea 3b. Calyx without an annulus; mature nutlets usually four Elsholtzia ia. Bracts linear or narrowly to broadly ovate, not membranous, usually green; bracteoles present section Aphanochilus ib. Bracts orbicular, usually membranous, often brown, purplish or bi-coloured; bracteoles absent ..••...•••••• iia iia. Fruiting calyx inflated, much broader than at anthesis; nutlets verrucose; style tips clavate section Platyelasmeae 217 iib. Fruiting calyx sometimes enlarged but never inflated, not much broader than at anthesis; nutlets not verrucose; style tips subulate section Elsholtzia 4a. Bracts orbicular, membranous, caducous; the whole plant covered in a thick indumentum of branched, stalked hairs 5 4b. Bracts sometimes broadly ovate but never orbicular or membranous, persistent; the indumentum variable but never of dense, branched, stalked hairs 7 5a. Ripe nutlets with a pronounced rostrate tip; corolla with an entire upper lip, hairy disc-like excrescences at the base of the filaments and a crescent-shaped invagination at the base of the lower lip Rostrinucula 5b. Ripe nutlets without rostrate tips; corolla with an emarginate upper lip and lacking excrescences at the bases of the filaments and an invagination at the base of the lower lip 6 6a Filaments shortly hairy towards the base; corolla without an annulus 218 6b. Filaments completely glabrous; corolla with an annulus of hairs Comanthosphace 7a« Anther filaments glabrous; corolla with rather broad, spreading lobes, the upper one emarginate 8 7b. Anther filaments with long, often purplish hairs; corolla usually with rather narrow and erect lobes, the upper one always entire Pogostemon iiia. All leaves opposite, usually petiolate; upper lip of corolla equalling or longer than the lower lip section Pogostemon iiib. At least some leaves in whorls of three or more, usually sessile; upper lip of corolla equalling or shorter than the lower lip section Eusteralis 8a. Calyx teeth plumose and elongated in fruit; corolla tube straight Colebrookea 8b. Calyx teeth not plumose or elongated in fruit; corolla tube gibbous towards the base Eurysolen 219 8.2. CONSPECTUS. In the following conspectus diagnoses are given for supra- specific taxa only. Species are listed in the order which best shows their relationships to each other. Species which are indented are possibly conspecific with the species which immediately precedes them. Comanthosphacineae Press subtr. nov. Bracteae caducae corolla bilabiata; labio superiori integro vel emarginato; antherae uniloculares; discus non lobatus; nuculae maturae e calyce secedentes. Pollinis granae binucleatae tricolporatae. Comanthosphace S. Moore in J. Bot., Lond. 1f?:293 ( 1877 ). Leaves ovate, petiolate; indumentum composed of branched and unbranched eglandular hairs; bracts at least as broad as long, membranous; bracteoles narrow, caducous; calyx sub-equally five-toothed; corolla with an emarginate upper lip and a complete annulus of hairs in the throat ( interrupted in C. nanchuanensis ); stamens with glabrous filaments; nutlets four at maturity, hairy ( except in C. nanchuanensis ), C. stellipila ( Miq. ) S. Moore C. japonica ( Miq. ) S. Moore C. sublanceolata ( Miq. ) S. Moore C. barbinervis ( Miq. ) S. Moore C. ningpoensis ( Hemsley ) Hand. - Mazz. C. formosana Ohwi C. nanchuanensis Y. C. Wu & Li 220 Leucosceptrum Smith, Exot. bot., 2:113, t.116 ( 1805 )• Leaves ovate, petiolate; indumentum composed of branched and unbranched eglandular hairs; bracts at least as broad as long, membranous; bracteoles narrow, caducous; calyx equally five-toothed; corolla with an emarginate upper lip, the throat without an annulus of hairs; stamens with filaments shortly hairy towards the base; nutlets four at maturity, glabrous. L. canum Smith Rostrinucula Kudo in Mem, Fac. Sci. Agric. Taihoku imp. Univ. 2,( 2 ) :30k ( 1929 )• Leaves ovate, petiolate; indumentum composed of branched and unbranched eglandular hairs; bracts at least as broad as long, membranous; bracteoles narrow, caducous; calyx sub-equally five-toothed; corolla with an entire upper lip, the throat with an interrupted annulus of hairs borne on disc-like excrescences at the base of the stamen filaments and on a crescent-shaped invagination at the base of the lower lip; stamens with naked filaments; nutlets four at maturity, hairy, the apices extended to form a strongly-hooked beak. R. dependens ( Rehder in Sargent ) Kudo R. sinensis ( Hemsley ) Y. C. Wu Subtribe Elsholtzineae Benth. ex Endl., Gen. pi.:612 ( 1838 ). Bracts persistent; corolla bilabiate, the upper lip emarginate; anthers bilocular; disc with one or more tumescent lobes; ripe nutlets separating from calyx. Pollen grains tri-nucleate / hexa-colpate. 221 Elsholtzia Willd. in Bot. Mag. 4:3 ( 1790 ). Leaves narrowly ovate to orbicular, petiolate; indumentum composed of unbranched eglandular hairs or a mixture of unbranched glandular and eglandular hairs, sometimes with branched stalked hairs; inflorescence cylindrical or secund, verticils one - many flowered; bracts linear to broader than long, sometimes membranous; bracteoles, when present, narrower than the bracts, persistent; calyx with five equal teeth or the three upper teeth shorter than the two lower; the corolla sometimes with an interrupted annulus in the throat; stamens with glabrous filaments, the anther locules confluent through partial fusion at the apex ( except in E. hunanensis where the locules are free ); disc with a single tumescent lobe; nutlets usually four at maturity, rarely one, glabrous. Section Elsholtzia. Synonyms. Cyclostegia Benth. in Bot. Reg. 13:sub t.1282 ( 1829 ). Elsholtzia section Cyclostegia ( Benth.. ) Benth., Lab. gen. sp.,:l63 ( 1833 ). Spikes usually secund, sometimes cylindrical; bracts at least as broad as long, usually membranous, veined, imbricate, the members of a pair sometimes fused to form a cyathium; bracteoles absent; fruiting calyx sometimes accresant but never inflated; style lobes subulate; nutlets rugulose or nearly smooth. E. heterophylla Diels E. bodinieri Vaniot E. strobilifera ( Benth. in Wallich ) Benth. E. luteola Diels 222 E. oldhami Hemsley E. argyi Leveille E. pseudocristata Leveille & Vaniot E. soulei Leveille E. nipponica Ohwi E. pygmaea W. Smith E. ciliata ( Thunb. ) Hylander E. feddei Leveille E. elegans Franch. E. hunanensis Hand. - Mazz. E. kachinensis Prain E. concinna Vaut. Section Aphanochilus ( Benth. ) Benth., Lab. gen. sp.,:161 ( 1833 ). Spikes usually cylindrical, sometimes secund; bracts linear to ovate, not membranous or imbricate, always free; bracteoles similar to bracts but usually narrower; fruiting calyx sometimes accrescent but never inflated; style lobes subulate; nutlets rugulose. E. myosurus Dunn E. pubescens Benth. E. pilosa ( Benth. in Wallich ) Benth. E. communis ( Collett & Hemsley ) Diels E. alopecuroides Leveille & Vaniot E. griffithii Hook.f. E. glanduligera C. B. Clarke E. elata Zoll. & Mor. E. winitiana Craib 223 E. blanda Keng E. ochroleuca Dunn E. stachyodea ( Link ) Raiz. 8c Saxena E. beddomei C. B. Clarke ex Hook.f. E. rugulosa Hemsley E. yunnanensis Press nom. nov. Synonym: Dysophylla mairei Leveill& in Bull. Geogr. bot. 22:236 ( 1912 ), E. mairei ( L6veill£ ) Press would be a later homonym for E. mairei Leveille ( 1915 ) ( * E. rugulosa ). E. capituligera ( Dunn ) Y. C. Wu E. stauntoni Benth. E. flava ( Benth. in Wallich ) Benth. E. fruticosa ( D. Don ) Rehder Leucosceptrum plectranthoideum ( L&veille ) Marquand E. penduliflora W. Smith Section Platyelasmeae ( Briq. ) Press stat. nov. Basionym: Series Platyelasmeae Briq. in Engl. 8e Prantl, Nat. Pflantzenfam. 3a: 32? ( 1897 ). Spikes cylindrical; bracts at least as broad as long, not membranous or imbricate, free; bracteoles absent; fruiting calyx inflated in fruit; style lobes with clavate tips; nutlets verrucose. E. eriostachya ( Benth. in Wallich ) Benth. E. densa Benth. E. manshurica ( Kitagawa ) Kitagawa 781 Keiskea Miq. in Annls Mus. bot. Lud.-Bat. 2:105 ( 1865 ). Leaves ovate, petiolate; indumentum composed of unbranched eglandular hairs only; inflorescence secund, verticils two-flowered; bracts ovate, not membranous; bracteoles absent; calyx with the three upper teeth shorter than the two lower, the throat with an annulus of white hairs; corolla with a complete annulus of hairs in the throat; stamens with glabrous filaments, anther locules free; disc with a single tumescent lobe; nutlets one ? at maturity, glabrous and with a somewhat reticulate pattern of ridges. K. .japonica Miq. K. sinensis Diels K. elsholtzioides Merrill K. szechuanensis Y. C. Wu K. glandulosa Y. G. Wu Tetradenia Benth. in Bot. Reg. 15:sub t-1300 ( 1829 ). Leaves ovate, petiolate; indumentum composed of unbranched eglandular hairs or a mixture of these and branched stalked hairs; inflorescence cylindrical, verticils many-flowered; bracts broader than long, somewhat membranous; bracteoles absent; calyx five-toothed, the upper tooth much broader than, and overlapping, the two lateral teeth; corolla with a complete annulus of hairs in the throat; stamens with glabrous filaments, the anther locules confluent through partial fusion at the apex; disc with four bright-red tumescent lobes equally spaced around the edge; nutlets four at maturity, glabrous. T. fruticosa Benth. e 225 T. goudotii Briq. Eurysolenineae Press subtr. nov. Bracteae persistentes; corolla bilabiata labio superiori emarginato; antherae uniloculares; discus non lobatus; nuculae maturae e calyce secedentes. Pollinis granae binucleatae tricolporatae. Eurysolen Prain in Scient. Mem. med. Offrs Army India 11:43 ( 1898 ). Leaves ovate, petiolate; indumentum composed of unbranched eglandular hairs only; bracts ovate; bracteoles similar but narrower; calyx subequally five-toothed; corolla with a gibbous tube and an emarginate upper lip, the throat with an interrupted annulus of hairs borne on disc-like excrescences at the base of the stamen filaments and on a papilla-like invagination on the ventral surface of the tube just below the gibbous curve; stamen filaments glabrous, the anthers unilocular; disc equal; nutlets four at maturity, glabrous. E. gracilis Prain. Subtribe Pogostemonineae Benth. ex Endl., Gen. pi.:612 ( 1838 ). Bracts persistent; corolla weakly bilabiate, the upper lip entire; anthers unilocular; disc equal; ripe nutlets separating from the calyx. Pollen grains bi-nucleate / tri-colpate. Pogostemon Desf. in Mem. Mus. Hist. nat. Paris 2:154 ( 1815 ). Leaves linear to orbicular, sessile or petiolate; indumentum composed of unbranched eglandular hairs, unbranched eglandular and glandular hairs or branched sessile hairs; bracts linear to ovate, not membranous; 226 bracteoles similar but narrower; calyx sub-equally five-toothed or with the three upper teeth shorter than the two lower; corolla with an entire upper lip; stamens with filaments hairy in the middle or towards the base, the anthers unilocular; nutlets four at maturity, rarely one, glabrous. Section Pogostemon . Synonym: Dysophylla section Oppositifoliae Benth., Lab. gen. sp.,:157 ( 1833 ). Leaves in opposite pairs, usually petiolate; the corolla with the upper lip equalling or longer than the lower lip. P. nilagiricus Gamble P. amaranthoides Benth. in DC. P. brevicorollus Y. Z. Sun P. formosanus Oliver in Hook.f. P. hispidus Prain P. wattii C. B. Clarke P. battakianus Ridley P. championii Prain P. gardneri Hook.f. P. heyneanus Benth. in Wallich P. cablin ( Blanco ) Benth. in DC. P. griffithii Prain P. paniculatus ( Willd. ) Benth. in Wallich P. villosus Benth. P. pubescens Benth. in DC. P. purpurascens Dalz. in Hook.f. 227 P. benghalensis ( Burm.f. ) Kuntze P. parviflorus ( Benth. in Wallich ) Benth. P. nelsoni Doan P. tuberculosus Benth. in Wallich P. dielsianus Dunn P. glaber Benth. in Wallich P. elsholtzioides Benth. in DC. P. rotundatus Benth. in Wallich P. mollis Benth. P. mutamba ( Hiern ) G. Taylor P. micafigensis G. Taylor P. reflexus Benth. in DC. P. atropurpureus Benth. in DC. P. speciosus Benth. in Wallich P. travancoricus Beddome P. litigiosus Doan in Humbert P. velatus Benth. in DC. P. williamsii Elmer P. fraternus Miq. P. nigrescens Dunn P. menthoides Blume P. brachystachyus Benth. in DC. P. paludosus Benth. in DC. P. reticulatus Merr. P. phillipensis S. Moore P. wightii Benth. P. macgregori W. Smith 228 P. rupestris Benth. P. hirsutus Benth. P. strigosus Benth. in DC. P. auricularius ( L. ) Hassk. P. rugosus ( Hook.f. ) El-Gazzar & Watson P. myosuroides ( Benth. in Wallich ) Kuntze P. salicifolius ( Dalz. ex Hook.f. ) Kuntze P. glabratus Chermserivathana ex Press sp. nov. Based on an unpublished description given by Chermserivathana ( thesis, 1963 ) and to be published at a later date. P. andersoni ( Prain ) Press comb, nov. Basionym: Dysophylla andersoni Prain in J. Asiat. Soc. Beng. 59:298 ( 1891 ). Section Eusteralis ( Kafin. ) Keng in Fl. Males. 8(1):352 ( 1978 ). Synonyms: Eusteralis Rafin., Fl. Tellur. 2(4): 95 ( 1837 ). Dysophylla section Verticillatae Benth., Lab. gen. sp.,: 15$ ( 1893 )• At least some leaves in whorls of three or more, usually sessile; the corolla with the upper lip equalling or shorter than the lower lip. P. falcatus ( Y. C. Wu ) Press comb, nov. Basionym: Dysophylla falcata Wu in Acta phytotax. sin. 10:237 ( 1965 ) P. quadrifolius ( Benth. in Wallich ) Kuntze P. cruciatus ( Benth. in Wallich ) Kuntze P.-ssemaseasis ( Y. C. .Wu 8c Hsuan ) Press comb, nov. Basionym: Dysophylla szemacensis Wu 8e Hsuan in Acta phytotax. sin. 10:238 ( 1967 ) 229 P. peguanus ( Prain ) Press comb, nov. Basionym: Dysophylla peruana Prain in J. Asiat. Soc. Beng. 59:298 ( 1891 ) P. yatabeanus ( Makino ) Press comb, nov. Basionym: Dysophylla yatabeana Makino in Bot. Mag., Tokyo 1:55 C 1898 ) P. pumilus ( Graham ) Press comb, nov. Basionym: Dysophylla pumila Graham in Edinb. New phil.J. 4:393 ( 1828 ), non Host ( 1831 ) P. lythroides ( Diels ) Press comb, nov. Basionym: Dysophylla lythroides Diels in Notizbl. bot. Gart. Mus. Berl. 9:1031 ( 1926 ) P. linearis ( Benth. in DC. ) Kuntze P. crassicaulis ( Benth. in Wallich ) Press comb, nov. Basionym: Dysophylla crassicaulis Benth. in Wall., PI. As. Ear. 1:30 ( 1830 ) P. faurei ( Leveill6 ) Press comb, nov. Basionym: Dysophylla faurei Leveille in Fedde, Reprium nov. Spec. Regni veg. 10:476 ( 1912 ) P. helferi ( Hook.f. ) Press comb, nov. Basionym: Dysophylla helferi Hook.f., Fl. Brit. India 4:640 ( 1885 ) P. tsiangii ( Y. Z. Sun ) Press comb, nov. Basionym: Dysophylla tsiangii Sun in Acta phytotax. sin. 11:50 ( 1966 ) P. aquaticus ( C. H. Wright in Dyer ) Press comb, nov. Basionym: Elsholtzia aquatica C. H. Wright in Dyer, Fl. Trop. Afr. 5:451 ( 1900 ) P. stellatus ( Lour. ) Kuntze P.,sampsoni . ( Hance ) Press comb, nov. 230 Basionym: Dysophylla sampsoni Hance in Annls Sci. nat. ser. 5,5:284 ( 1866 ) P. griffithii ( Hook.f. ) Press comb, nov. Basionym: Dysophylla griffithii Hook.f., Fl. Brit. India 5:641 ( 1885 ) P. pentagonus ( C. B. Clarke ex Hook.f. ) Kuntze P. stocksii ( Hook.f. ) Press comb, nov. Basionym: Dysophylla stocksii Hook.f., Fl. Brit. India 4:642 ( 1885 ) P. koehneanus ( Muschler in Fedde ) Press comb, nov. Basionym: Dysophylla koehneana Muschler in Fedde, Reprium nov. Spec. Regni. veg. 4:269 ( 1907 ) P. gracilis Hassk. P. deccanensis ( Panigr. ) Press nom. nov. . v &4St'ariigr\: deccaneosis fksu^r. in Pkijhol^^ 32.: Synonym: Dysophylla tomentosa Dalz. P. tomentosa ( Dalz. ) Press would be a later homonym for P. tomentosa Hassk. ( 1844 \ P. trinervis Chermserivathana ex Press sp. nov. Based on an unpublished description given by Chermserivathana ( thesis, 1963 ) and to be published at a later date. Colebrookineae Press subtr. nov. Bracteae persistentes; corolla bilabiata, labio superiori emarginato; antherae uniloculares; discus non lobatus; nuculae maturaee calyce non secedentaS. Pollinis granae binucleatae tricolporatae. Colebrookea Smith, Exot. bot. 2:111, t.115 ( 1805 ). Leaves ovate, petiolate; indumentum composed of unbranched eglandular hairs only; bracts linear, not membranous; bracteoles similar but 231 smaller; calyx with five equal, long, slender, plumose teeth; corolla with an emarginate upper lip; stamens with glabrous filaments; nutlets one at maturity, hairy. G. oppositifolia Smith C. ternifolia Roxb. Species excludendae Elsholtzia integrifolia Benth. = Schizonepeta tenuifolia ( Benth. ) Briq. Elsholtzia chinense = Agastache sp. ( ? rugosus Fisch. & Mey. ) 232 ACKNOWLEDGEMENTS. This study has been carried out at the Department of Botany, British Museum ( Natural History ). I would like to thank the Keeper of Botany for providing facilities for the work and the trustees of the museum for providing financial assistance. My sincere thanks go to my supervisors, Drs C. J. Humphries, D. Dalby and T. C. Whitmore for their patience and guidance. I would also like to thank Mr A. 0. Chater for many useful discussions and comments and Miss Ann Lum for translations of Chinese literature. On the technical side I am grateful to Kay Shaw for explaining the computer programs and to Roger White for running them. 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Y. 1965* Hostrinucula sinense. In C. Y. Wu, H. W. Li, S. J. Hsuan and Y. C. Huang, Materiae ad floram Labiatarum sinensium (2). Acta phytotax. sin. 10(3):215 - 24-2. & HUANG, Y. C. 1974. Materiae ad floram Labiatarum sinensium Acta phytotax. sin. 12(3):337 - 3^6. 1977- Labiatae. In Flora Reipublicae Popularis Sinicae 66. Peking, Science Press, pp. 64-7. WUNDERLICH, R. 1963. The Pogostemoneae - a debatable group of Labiatae ( some remarks on their taxonomic position with regard to their pollen grains ). J. Indian bot. Soc. 42A:321 - 330. APPENDICES. 241 Appendix 1. Specimens used for scoring and producing averaged data. For the names of species to which the acronyms refer see table 3. KJAP 1; Bot. gard. Tokyo s.n. 1880 ( TI ), Herb. Terasaki s.n. 1906 ( K )., Yano s.n. 1890 ( TI ) - CSTE 2; Murata 19058 ( TI ), 19171 ( TI ), 36037 ( TI ) - CBAR 3; Humsawa s.n. 1947 ( TI ), Kanai 6012 ( TI ), Yamozaki 6 ( TI ) - CSUB 4; Bisset 102285 ( BM ), Hara s.n. 1953 ( TI ), Togasi & Matsuoka 180 ( BM ) - CJAP 5; Maximovicz s.n. 1862 ( BM ), in Herb. Hance 13214 ( BM ), Tschonoski s.n. 1864 ( BM ) - CNIN 6; Law 1012 ( TAI ), Liou 1378 ( E ), Handel - Mazzetti 2602 ( E ) - CTER 8; Roxburgh s.n. Mysore ( BM ), s.n. Mysore Bot. gard. Calc. ( BM ) Isotype?, s.n. Hort. Calc. ( BM ) - COPP 9; Clarke 34640B ( BM ), Hooker s.n. Sikkim ( BM ), Thomson s.n. Mont. Nilghiri 8c Kurg. ( BM ) - ECON 10; Ludlow 8c Sherriff 9057 ( BM ), Polunin 1020 ( BM ), 1612 ( BM ) - ECIL 11; Togasi 787 ( BM ), Wilson 5717 ( BM ), Yu 10728 ( BM ) - EKAC 12; Forrest 6729 ( E ), Kingdom - Ward 20365 ( BM ), Put in Herb. Kerr 3478 ( BM ) - EPYG 13; Forrest 17128 ( E ) Holotype - ESOU 14; Maire 321 ( E ), 599 ( E ), Soulie 226 ( E ) Holotype - EARG 15; Cavalerie 7993 ( E ), Leveille s.n. ( E ) - ELUT 16; Forrest 11145 ( E ), 15184 ( E ), Yu 13952 ( BM ) - EBOD 17; Bodinier 2(E) Holotype, Forrest 2997 ( BM ), 7371 ( BM ) - EHET 18; Forrest 934 ( E ) Holotype, Maire 1203 ( E ), Collector unknown 096035 ( TAI ) - ESTR 19; Ludlow Sherriff 8c Taylor 7139 ( BM ), Nicolson 2652 ( BM ), Stainton Sykes 8c Williams 8526 ( BM ) - EERI 20; Dhwoj 0180 ( BM ), Ludlow Sherriff 8c Hicks 16682 ( BM ), Stainton Sykes 8c Williams 8102 ( BM ) - EDEN 21; Ludlow Sherriff 8c Elliot 14130 ( BM ), Stainton 24 2 Sykes & Williams 2169 ( BM ), 8102 ( BM ) - EAQU 22; Johnson 15 ( K ) Holotype, Fanshawe 8503 ( K ), 8512 ( K ) - EINT 23 Staunton s.n. ( BM ) Holotype - EHUN 24; Handel - Mazzetti 2702 ( E ) Isotype - EPEN 25; Forrest 11686 ( E ) Holotype, Yu 17600 ( E ) - ERUG 26; Forrest 6527 C BM ), Maire 633 ( E ), Yu 16748 ( E ) - ESTA 27; Jackson s.n. 1929 ( BM ), Licent 2996 ( BM ), Staunton s.n. ( BM ) Holotype - EFLA 28; Forrest 6283 ( BM ), Stainton 5090 ( BM ), Wallich 1553 ( BM ) Isotype - EFRU 29; McLaren's native collectors 326d ( BM ), Polunin Sykes 8c Williams 3126 ( BM ), Young s.n. 1880 ( BM ) - EPUB 30; Horsefield 337 ( BM ) - ECOM 315 Maire 561 ( BM ), Yu 14498 ( BM ), 14712 ( BM ) - EALO 32; Cavalerie 1426 ( E ) Holotype - EGRI 33; Haines s.n. 1914 ( E ), Lace 4391 ( E ) - EGLA 34; Clarke 37393 ( K ) Holotype - EELA 35; Koorders 37621B ( K ), Herb. Kuntze 5613 ( K ), Ridley s.n. 1915 ( K ) - EWIN 36; Kerr 1607 ( BM ), s.n. ( BM ), Put in Herb. Kerr 4412 ( BM ) - ESTC 37; Buchanan s.n. 1802 ( BM ) Holotype of Elsholtzia lept-ostachya, Clarke 23765 ( BM ), Polunin Sykes 8c Williams 5827 ( BM ) - EBLA 38; Ludlow Sherriff 8c Taylor 7075 ( BM ), Norkett 8631 C BM ), Stainton Sykes 8c Williams 8323 ( BM ) - EMYO 39; Forrest 7220 ( E ) Holotype, Wilson 3533a ( K ) - EOCH 40; Maire s.n. ( E ) Holotype of Elsholtzia lampradena - EPIL 41; Forrest 28965 ( BM ), Maire 1204 ( BM ), Stainton Sykes 8c Williams 4452 ( BM ) - ECAP 42; Forrest 1680 ( BM ), 20697 ( BM ), 22956 ( BM ) - DTRI 43; Kerr 81150 ( BM ) Holotype, Put in Herb. Kerr 2223 ( BM ) - DGLA 44; Kerr 10274 ( BM ), Put in Herb. Kerr 132 ( BM ) Holotype - DTOM 45; Buchanan s.n. ( BM ), Hohenacker 371 ( BM ), Young s.n. 1879 ( BM ) - DPEN 46; Clarke 20438 ( BM ) Isotype, Garrett in Herb. Kerr 59 ( BM ), Kerr 1465 ( BM ) - DSTO 47; Stocks s.n. in Herb. Hooker ( K ) 243 Holotype - DSAM 48; Sampson in Herb. Hance 10946 ( BM ) Holotype - DPEG 49; Kurz 2401 ( K ), 2405 C K ), Put in Herb. Kerr 1971 ( BM ) - DGRI 50; Chattaya 5224 ( K ), Gamble 13748 ( K ), Griffith 3968 ( K ) - DMT 51; Science College imp. Univ. Japan s.n. 1883 ( K ) Isotype, Tanaka 7338 ( TAI ) - DCEA 52; Clarke 23691 ( BM ), Griffiths 1024 ( BM ), Wallich 1545 ( BM ) Isotype - DSTE 53; Clarke 8073 ( BM ), Loureiro s.n. ( BM ) Holotype, Simpson 9189 ( BM ) - DLIN 54; Clarke 45731 ( BM ), Kingdom - Ward 14260 ( BM ), Ludlow Sherriff 8c Hicks 21019 - DCRU 55; Beddome s.n. ( BM ), Clarke 18059 ( BM ), Polunin Sykes 8c Williams 5875 - DQUA 56; Roxburgh s.n. ( BM ) Isotype, Rugel s.n. ( BM ), Wallich 1538 ( BM ) - DAUR 57; Clarke 26464 ( BM ), Forbes 89 ( BM ), Stainton Sykes 8c Williams 6481 ( BM ) - DSAL 58; Dalzell s.n. ( K ) Holotype, Lain s.n. in Herb. Hooker ( K ), Young s.n. 1882 ( BM ) - DRUG 59; Beddome s.n. ( BM ), Herb. Bottler s.n.1828 ( K ), Wallich 1547 ( K - W ) Holotype.- DMYO 60; Fricker 4724 ( K ), Koenig s.n. ( BM ), Wallich 1547 ( K - W ) Holotype - PBEN 61; Roxburgh s.n. India ( BM ), Stainton 13 ( BM ), 5220 ( BM ) - PPAR 62; Andrews 348 ( BM ), Beddome s.n. ( BM ), Metz 1393 ( BM ) - PPAN 63; Stocks 8c Law s.n. Malabar 8c Concan ( BM ), Thomson s.n. Mont. Nilghiri 8c Kurg. ( BM ), Wallich 1561 ( BM ) - PTUB 64; Clarke 26368 ( BM ), Ludlow Sherriff 8c Taylor 6759 ( BM ), Sarail 23 ( BM ) - PGLA 65; Flatt 161 ( BM ), Hooker s.n. Sikkim ( BM ), Nicolson 2941 ( BM ) - PHEY 66; MaCrae 737 ( BM ), Robinson 8c Kloss 88 ( BM ), Thomson s.n. Maisor ( BM ) - PCAB 67; Clemens s.n. 1924 ( BM ), Horsefield s.n. Java ( EM ), Ramos 22432 ( BM ) - PELS; Griffiths 1018 ( BM ), Kingdom - Ward 14234 ( BM ), Ludlow Sherriff 8c Taylor 7211 ( BM ) - PAMA 69; Hooker s.n. Sikkim ( BM ), Murata 06306529 ( BM ), Stainton Sykes 8c Williams 9276 244 ( BM ) - PFOR 70; Kao 4883 ( TAI ), Kudo & Mori 2401 ( TAI ), Simada 5419B ( TAI ) - PMEN 71; J. & M.S. Clemens 32575 ( BM ), 40267 ( BM ), P^telot 5110 ( BM ) - PFRA 72; Clarke 44065 ( BM ), Horsefield s.n. Java ( BM ), Kerr 9658 ( BM ) - PBRA 73; Clarke 40303 ( BM ), Griffiths 222 ( BM ), Hooker & Thomson s.n. Khasia ( BM ) - PMIC 74; Grossweiler 2545 ( BM ) Isotype, 9668 ( BM ), Raynal 12161 ( EM ) - PMUT 75; Grossweiler 12134 ( BM ), Welwitsch 5496 ( BM ), 5990 C BM ) Holotype - PNIG 76; Henry 9082 ( K ), 11174 ( K ) Holotype, 12563 ( K ) - PPHI 77; Ramos 33320 ( BM ), Ramos & Edano 45012 ( BM ), Whitehead s.n. 1896 ( BM ) Holotype - PREF 78; Herb. Hooker 80 ( K ), Thwaites 154 ( BM ), Walker s.n. Ceylon ( K ) Lectotype - PRET 79; Ahern's collector Forestry Bureau no. 3395 ( BM ) Holotype, Edano 48842 ( BM ) - PMOL 80; Clarke 10673 ( BM ), Gardner s.n. in Herb. Miers ( BM ), Vine 216 ( BM ) - PTRA 81; Beddome 109 ( BM ) Holotype - PATR 82; Beddome s.n. Anamallays ( BM ), s.n. W. slopes of Nilgherries ( BM ), Herb. Wight 2127 ( K ) Holotype - PSPE 83; Metz 1225 ( BM ), Schmidt s.n. Nilghiries ( BM ), Vine 215 ( BM ) - PVEL 84; Cuming 1097 ( BM ) Isotype, McGregor 11339 ( BM ), Mendoza 40923 ( BM ) - PWIL 85; Elmer 22225 ( BM ) Isotype - PPUR 86; Herb. Dalzell s.n. ( K ) Holotype, Stocks s.n. Concan ( BM ), Vasnoli s.n. 1881 ( BM ) - PPAL 87; Gamble 17853 ( BM ) - PVIL 88; Masters s.n. in Herb. Hooker ( K ), Roxburgh s.n. India ( BM ), Wallich s.n. 1831 ( K ) - PWIG 89; Gamble 18389 ( BM ), Schmidt 74 ( BM ), Stocks & Law s.n. Malabar, Concan ( BM ) - PROT 90; Lawson s.n. 1884 ( BM ), Wallich 1535 ( K - W ) Holotype - PSTR 91; Clarke 5501 ( BM ), 15617 ( BM ), Kingdom - Ward 18763 ( BM ) - PHIS 92; Herb. Hooker s.n. ( K ), Jenkins 346 in Herb. Hooker ( K ) Paratype, Kerr 6631 ( BM ) - PPUB 93; Kerr 2384 ( BM ), 3113 ( BM ), 245 Petelot 5297 ( BM ) - CFOR 94; Huang 7040 ( TAI ), Huang 8c Hsieh 7297 ( TAI ), Kao 8611 ( TAI ) - RDEP 95; Cavalerie s.n. in Herb. Leveille ( E ), Wilson 5534 ( E ) Holotype, 4313 ( BM ) - RSIN 96; Bodinier 2709 ( E ) Holotype of Leucosceptrum bodinieri, Henry 7765 ( E ) Holotype - LPLE 97; Maire s.n. ( E ) Holotype - ENIP 98; Ikeo 9567 ( TI ) Isotype, Kurata s.n. 1964 ( BM ), Mayebara 4125 ( TI ) - EOLD 99; Oldham s.n. 1864 ( K ) Holotype, Tagawa 4162 ( TI ), Yamazaki 3059 ( TI ) - ECHI; d'Argy s.n. Kiangsu ( E ) - EPSE 101; Taquet 1223 ( K ) Paratype, 1224 ( K ) Paratype - DMAI; Maire s.n. 1911 ( E ) Holotype - DFAU 103; Faurie 760 ( E ) Holotype - DAND 104; Anderson s.n. 1867 ( K ) Holotype - DKOE 105; Hosseus 704 ( EM ) Isotype - DGRA 106; Dalzell s.n. ( K ) Holotype, Stocks & Law s.n. Malabar & Concan ( BM ) - DLYT 107; Fan & Li 563 ( EM ) - DHEL 108; Heifer 194 ( BM ), 3968 ( K ) Holotype - DPUM 109; Wallich 1546 ( K ) - PNEL 110; Cook's 3rd voyage s.n. ( BM ) - PBAT 111; Ridley s.n. 1921 ( K ) Holotype - PWAT 112; Clarke 41719 ( K ) Holotype - PHIR 113; Beddome s.n. ( BM ), Simpson 9049 ( BM ), Thwaites 283 ( BM ) - PRUP 11-4; Cramer 4277 ( K ), MaCrae 396 ( K ) Holotype, Thwaites 343 ( BM ) - PMAC 115; Hansen & Smitinand 12661 ( K ), Iwatsuki Fukuoka 8c Chintayungkun 9659 ( K ) - PGAR 116; Gardner 1847 ( K ) Holotype, Herb. Wight s.n. ( K ), V. Row 3229 ( K ) - PCHA 117; Champion 339 ( K ) Holotype, Shiu Ying Hu 12421 ( K ) - PDIE 118; Forrest 875 ( K ) Isotype - PGRI 119; Griffith 3962 ( K ) Holotype - PNIL 120; Bourne 5094 ( K ), Herb. Hooker s.n. ( K ) - PLIT 121; Evrard 1834 ( K ), Poilane 24241 ( K ), s.n. ( K ) - TG0U 122; Goudot s.n. ( G ), Hildebrandt 3471 ( G ) Holotype - THZL123; Hildebraat 3971 ( G ) Holotype - TFRU 124; Forsythe - Major s.n. 1895 ( G ), Lyle 278 ( K ) Holotype - EMAN 125; Nakai Honda 8c Kitamura s.n. 1933 ( TI ) 246 Holotype, Togashi 828 ( TI ), 1578 ( TI ) - EGRA 126; Put in Herb. Kerr 4404 ( BM ), Robinson & Kloss 130 ( BM ), s.n, 1914 ( EM ) - EFED 127; Forrest 22386 ( E ), Soulie 227 ( E ) Holotype - EELE 128; Bau Hwa Shan 1490 ( E ) - EBED 129; Beddome 147 ( BM ) Holotype, MacGregor 69 ( E ), 608 ( E ) - KELS 130; Jiangsu group 2667 ( KUN ) - KGLA 131; Y. Ling 728 ( KUN ) Isotype - KSIN 132; Chen Quan 999 ( KUN ) - KSZE 133; T. P. Tsung 39354 ( KUN ) Isotype - CNAN 134; Z. H. Hsiung & Z. L. Chow 93127 ( KUN ) Isotype - PBRE 135; Chen 3206 ( KUN ) Isotype - DTSI 136; Tsiang 9449 ( KUN ) Isotype - DFAL 137; Wang 79441 ( KUN ) Isotype - DSZE 138; Tsiang 12713 ( KUN ) Isotype. 2 47 Appendix 2. 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S 76,4 68.1 72.8 74.2 73.7 69.8 64 65.0 70.7 70.5 70.7 70.3 73.3 69.6 73,5 71 ,0 67.9 76.1 70.3 73.7 76.3 74.1 70.9 65 69.8 70.7 73.8 70.7 70.7 70.6 70.0 72.7 74.1 72.8 70.9 69,5 66.7 70.7 68.1 69.1 66 64.4 69.1 68.0 67.6 63.5 70.7 67.2 72.0 69.* 71 ,0 80.4 70.5 76.5 76.5 76.0 70 P 5 67 63.6 68.1 66.9 66.6 67,4 69.7 66.2 70.3 69.4 74,6 76.9 67.2 71.7 72.2 75.0 65.9 68 69.4 73.1 73.2 73.4 73.6 7^.5 72.5 77.3 75.1 70.5 76.9 72.4 77.2 77.9 76.5 75.1 ^ 69 68.2 68.9 71 ,7 68.8 68,5 70.2 71.0 79.4 77.2 71 ,3 73.2 71 .8 69.3 70.6 69.8 70.3 ^ 70 64.8 70.4 69.9 70.2 70.1 73.1 70,7 73.1 70.6 70.6 80.2 71.0 76.9 76.6 78.5 OTU 1 2 3 4 3 6 7 8 9 10 11 12 13 14 15 % Appendix 3 ( continued). 812 S 255 o ro 1— o co 0 in p*- ccr o cc r\j I— O o ^r in r j co f\I vO T— Pv. o C- tv- T— O CM COr j © O O O rj o vO rofM N-CM r- in vt <1 s o o vl- ro o ro vT •O O VR CM0 0 ooo CMc c o (M CO o ^ ro o c ro IM ro CM oc e- I - • • • • • • a a •> • a a S• • • • a a a a » • a • a a • B a a ^ B a a B l^v •4S " OT — O ROs o O C R— O IN. •oI -m p-r- rc ro o rv. 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MstOC1 Is- sf r- C1 n M fM in ^KICO vJ^CCONOM^r-NOvr^ ro m O J>3 " <>3r" fFMM • • a • • • a • -J- 0 0 0 FM ro rj t— CO cc 00 OC Is- O IN.I s- s s s 00 r\j co C tN o Is- OC vj D- in- ,-NC Oc T—< 0 r-fNC MfM O xC* c co O CJ O T—C C —1 x— 0 0 r- O CN* — G C co a a » a » a a a a a a •a • a a a •- a * a a a a a » a a • a a a e c; c:- s <3 r- COC NO t- CMr o LA NO I C-O 0 0 T~ r j ro m <3 r- 0OO NO T—r o to 10 NO| N CO 0 0 C-' G v- t- T • T- r- t " TV-" *— ro fVj CMrM C j ro r j rvj o-i ro ro ro ro ro fO ro ro Ki t- x— r- t- V \ - , - \ - —t r- K— T" V" T— V X —x— x- 257 OlfltOCNNOinT-NKMOOCvl-vOxtK'OKlS OCMOOKlK'JDOCCNMO^CinrMtMOxJ O O-O O cc 00 CO OC oc CC r^vO NOO NN vO N Appendix 8 ( continued )« OWMON-inr-r-W <5 (M O O0C O fM fO O N O (\J • •• • •• • a > » « t • •• O 'J> n N N N o C t- ir> o CO O N vT -J CO N CC-C C CCCOCCC^COM^KNNKVOVCNNON OONCOOfOfO• •O• •OOMMNOOKI^COO^(M•• ••• » • •>• o o ro t— i— o pjo t—rvj o o -O rMrv_room*— r\JON~) O O O O O O C> CO O C- CO 00 OC N N N ^o N CO'Or-NvJxrK(\JsOr-CCLn>f CC'^OCCl^T-CvfMCMVO' K: fO O C N n v} CO O (V vf vf O • •••••••••••••a aaaaaaaaaaa VO Ot^OMCMT.OlACCT-NMSir, tO O M O >3 T- RE U'I N ^ N O ro -3 ^3- o:> co co co sj- UNt o O CO'r o o o o cc *— c ro -r 00o CN < ' CM • • m a a a a « a a a a * a a a c t a a CM v. cc o *— to r • to ro c- co irc c to P- r^ NCO CO CMLT . 00in ^o UN OO P-. o p- P- r - r- -o r-- r- r^- r- p- r- r- r- -0 P- C 00 p- COC O p-oot o 0C CO •LO ro ro c CO >TR J O N LTIT,. 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S 70.3 67.1 69.3 73,9 77,2 73,0 75.4 72.5 78.2 66 76.1 i c.. n 68.1 67.2 69.3 69.6 66.7 69.6 71.3 70.3 78.0 78.8 79,5 72.6 73,1 77.5 7 *? 7 67 74.3 f r. . r 70.3 67.6 65.6 65.3 67.2 65,3 69.0 73.6 73.5 76.9 76,3 69.4 67,8 77.5 OTU 33 34 35 36 37 38 4o - 39 41 42 43 44 46 hi 4S Appendix 3 ( continued ). 262 CNJ Ov C o •vj-r\j (\l m oc >C ro in CM 00 COo >o ro » • • » • • • • • • * m • • k • • •m • • OO r— r- CNlo o rs- O N- fV-m o o e- in o o p- p- r*. CO 00 co r^-<3 O o >G o ND r*. r^ o oc O in r- Sf -O M OO N- r- N M fOr O vOvi - mP -m o <1- M -r o o O LTlI O CO SB »••»>•• » 1 » • • m • • • » • • • •m • o • • • • • m vo N in «0 N r- K Ln —\ o O r- o CODC oc 30 in i/i «- N. m r- T~ CO ft • p LOv • • • • m * • • • •• •• • • V• • -Ct- o P- CNJ p-r- o c- h- o o ro Csl o V-CO »o Kl «— p- r^-N- r- CO o- oc r^-sC CM tn m m m o POo rv. tn c-r- in T— N-oc 00 o <\i • • • • • • • m • • • • • • • • p• • m cc c o o rv in o ru o co oo o <\J r-ru CO o o in T—in NO <0o o N COr - r- tv- >o o O o O •Q 00 lOO r- *— ro in N- O to o o 00 o sO o in LTL -J- ro CNJ CVJ Pwi n PO o cr Ps. O G O r- to N CNJ CNJ o • • • • • • • • • • • • • • • • » • • • a C\J O CO < O 00 fl rr TsiO 1/ CNJ M T— cv 00 o in -J- o rv- ro C cc •c o -o o CM tv. o o P- PW Pv. r^. r- nC CC p~ p- CO cc o r— o •o ro c in C rvj O vO o RS.o o o • • • • • • • • • • • • • • • • » •• < a • e o O tO1 G O O Lf\ O rO m oo 00 T-CO •sj- o <3 O o fVi CNJ c o o CO ir. LO • • • i • • • » • • • • • • • > • • • • a N > , S vjinOt-0"0 000 KmC (\ rx.' in vO m o (V ro m 0 T— r- CO ro o rj CO o N N N N N -O <0 V N O >C C V <> o >W <> p- o P»- -O N- o cc cc cc p- co cO iT' ro T- o c o R^-C rj r\. co O O in o O • > • • • • • » • • • • • • • * • • r • • ON c in O^MWirn-^lAr-KlC ro e- O ir> cr o T— o rv. CNJ o o P- ro CO in r.i oo oo _ a •• •*••••< • » • • • • • • m > • • • • • • • • k a a ro ro T- CM R- ro O ro ro >0 c o CO *— L*. c. r\j T- r- m ru 00 ro ru rj r- o Ki r- CC e O o Cv in -o T— C in o CO O- • • • • • • • • • • • • • * • • • • • • a • < n O Mn K, c o Cs. Tn. & C' e to o r- r. o CO sT T— cr o rj o CN n N f» N n N s s r^ r- r^ r- NO N0 P-p^- O o r- fv. R- p- p^- vC cc cc CC cc CO tC F^ R- UI C> N ^ C T- R- R- cc P^.t ~ v— U". x— Cr bn vO CNJJ V P-*o vO in ro rvj * • • • • • • » r • • » • » > » • • • • «» vo t^, < n ^ c c- o to o o o LT c: io T- t— m ro o P- T" O o rj r-j CN o 00 c 00 JV FVL N N N F- S N P- P- C O F- o c- P-r- P-P- r-. o p- 00 p» p- oo c C- n ' ^ O r Tvlr- C N c; r , IT. »T vT re o 00 00 o m PO P- ro o VD t— co VQ.C s o ro o o CN 00 o C rj r j • • • • • • • • t • « • • • • • • • i • < <3 OO COo o r- to CN sC P-m CN eo t- fv- o r - r-- p- N- r-- CO r- OO oo p- r«- t^ vD P^ 00 oo oo 00 oo jo0 Os O r J roV F in o r^ oo o o T— ru to^ r m vO Pw Os o CVJ 00 oo 00 oo 00 00 00 oo o cr> O ChC N o CNO CN o o o o O Appendix 3 ( continued ). 263 o vrOomoNt-oco-oo xvooooococrN.*—cc OrOlAfCOOOC O W r Ot oin N MO L O LT N oto N r «0 O O0 fO fx -OO -N C' O M N O OKI N •> • • a a a a a a'a i a a a CN. 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O © ro © O ro O to LPv a • a a a a a a a a a a a a a a a • a a a • a a a a a a a a a a r • O X— © O CM P- Cxi px- xO (N IN vO >G ro m © CJ ro in xO xO IN © 0 T— T— LO O xO UN xf N) fO OC ro — © O © CO © tN CO OD CO 00 co CO 00 IN 00 CO fx. xO -O xG IN IN IN IN fN xG xG xG xO xG CO cc © O- © iC N- r\j © O xf rvj CJ px- CM UN Px. © xf © © co fx- LPi CO K: IM T- CM © xC LP, ro © K) © -4- a a a a a a a a a a a a a a a a a a a a a a a a a a a a a a r a » a 1—1 © in xC tN T— CO xf ro xf P- CC ro © ro cj ro xG U*. r— NO Nf v- Nf ro r- © LPi CO xG fx T— xC' p- © O s CO CO |N CO cc CO CC CO IN IN tN IN N P-. P- fx— p- \©— oc 00 CO CO co iC c c lO cc I — xO vO xC xG CO © © - Nf cc OC px UN tN ro T ro 0 © Xf xC Xf Nf CM ->f © © fN © O LO © cc CJ CC xO LC .•N © © ro UN ro a a a a a a a a a a a a a a a a a a a » a a • a • a a a aaa a a a a O tr- O © UN O ro vO © © cj ro O < © O © T— O NO in. © O P- 00 r- r j C. LP 1 xG UN CJ *— —. cc oz © © CO © P- P~ CO to cc bo IN P- CO 00 CO P- O xO •O IN Px NC •G N- •c •G •<3 •O xG CO © © © © 0 ro rO 00 IN O © xO © © © CJ xt x~ IT. co N' 0 LP Nf IPI © ro © ro xO u'^ -f xG 03 « ro • a a a a a a a a a a a a a a a a a a a a a < a a a a a a a a a a 1 • O oa CM 10 CJ © f\J in. IO. C, CO P- © ro 0 to -Q L-' LP. Nf © © CO © in U". P- N". x— UN CM in NO S R- px- FN P- N xG |N I - 00 P- fN P- p- IN fN |N r- IN IS- P—' fx- CO IS- fN P- r- fx. |N- fx. FN r~ fN r- TN co ro to T- in fN xf © © fM C © © xf in Nf O fM INJ © Cv! Cxi xC CJ c 00 CO IN •G CO fN v- aaa a a a a a a a a a a a a • a a a a r a a # a a a a a a a • • • O xf CM N» ©X xf f\J |N IN O0 L- © P- O f O OO 00 © O xf r- 00 r. © c © CJ .f 1.0 V s NN N fx. IN fN I - CO IN fN fN fN px p- fN IN IN r- r- cc- N © © N co CC cc cc F- r- xO r- i — » f ^ © ro T- x-0 ro © r-N RN xG L-N xO OO |N r- CJ 00 r- CJ tN xO © fx- M IN vO © CM ro UN -f Nf * a a a a a a • a a a a a a a a a a a c a a a a a a a a a a a a a a a 0 - 0 O UN XC O UN 00 IN OO 10. rv ro xf © UN © OO TO in ro ro K~ © © r- xO Nf in fN 00 T CJ CJ UN N- S s IN r- p- !N tN IN >G I -fx- 00 so 00 IN |N IN CC P- P- r- P- f- 00 fN I — 00 P- IN fN |N IN IN.C O r- |N 'N xG X? • ir. ro LO UN (M r— fN vf UN IN 00 © © r— CJ ro xf UX) xO tN 00 © O t— CJ ro OTU 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 -nru3 ro 129 100.0 130 74.1 100.0 131 72.7 93,3 100.0 132 76.7 93.7 87.7 100.0 133 75.6 96.0 89.5 92.7 100.0 134 73.0 70.3 73.6 66.1 67.6 100.0 135 75.9 67.6 66.7 70.9 71.2 66.8 100.0 136 67.5 61.3 60.5 60.7 63.0 62.7 81.5 100.0 13 7 76.7 63.9 67.3 63.1 62.2 70.5 84.2 84.1 100.0 138 75.2 63.5 64.7 64.3 64.0 66.4. 83.6 90.3 94.6 100.0 OTU 129 ^30 131 132 133 134 133 136 137 138 8 • 08- £Zl IZl 9Z V • g*- S<5 »+- 96 Hi o- •44XXX+*-* -•-••-- i c\j *XXXX8—•• *t6 • 8008 • — 2 • 000 * "' • 08 ' s £ 9 • -•^•-••••••••--•••••••••••••X'tX**''*'*' S2 21 02 • X" 12 S21 121 006711 •XXX— 69 ,444+X+4-4 4 -x«xx*xx»-*x*xxxxxxxxxxxxxxxxx & »-• +-*».XXXXXXXXXX« •XX-»»-- — • — * • zz »+«+44-+4 X* •••••«-« -»+•••- •---•xXX *XX4X-XXXXXX- ••»«••••«••».»* — Oil • ••XXXX+ ••••X-« • +-+.-«. x«»» 4«-«-»x»-»x-x * xx xx--" * • - C8 • 9AXXX* -»*XX-»*4*4•»4+4x»-4»•• X—X4»*>r**•-•-»• XX — £^ 11 ' u 21 > 21 44 + - + + * • • • • • •• X «•• «••» •><•«•»«•»•»• »*4K4X»»+XXXXX XX* X x • - 91 # - x»*---XX«•*-•»•» + *X*4 4 KXX + «~XX-»>XXX t* XXXXXXX-+ X4 X + i + 44 x • x* — is, *•*-*-' 98 «X8**X-*—• 4 XXX + XXX* *X+4 XX* »XX*^XXX »•••» XXYX-- ozi *8*XX4-*-4 •XXYXyXXXXXVXX*-»XXxXX4-*» XX^XXX******"*******— *ri «-XX+*+-»*-» + t*'»4x+xx«-+^xxxxx*x-»xvx+*xx**xxxxxx+xy*vxxxxxx— it •••4xXXXX*XXXXXX4*44XXXXX-4t -444*444 +4+4+-_ 9CI *84*-+^<*4XX*tXXXXX4XXX*-X*^***t+*+XX-»XXXX*XX*Xf»^-»*XXXX-4 * + + -+4->+ • 6095 •+x+m«4xxx*xx/«xxx*8"'-xx*x+»+*x»+8X4XX-tXXyX4*444.xxXX • '*xxxxxx**x+x**— 4.+—-4-*—4 4^xyxy*-»—•-«•—•—t+++-44—4 - • ty •xxxxxxe®xy-*x+xxxxxx—• t 901 •flXX8XXXX8*X«+*XXX*+* , 4- ----——> 4 — 05 »XX8XXXX8*XXXXXXXt--+-4—— ——-i——— 1 • »XX8X«88XXXXXXXXX-4«-4-44-*4»»--»«-4«-H-4444fX+-«-4«.4X4X £8S •*8iB888XyXXXXXX<«-*44.—V—4-4-4.+4+V 808XXXX8XXXXXXX4-4-X—• ••^-••••••••••4-4-t»-X4-4*44>tXl-X— — to V Z s * 888XXX 88X8 8 *••••«• 4 •••*••• •4-»X*-X4 4-X4X--**44VXX-- /Ol * 80XX XX8888S4-4--M 4-4 + 4-44 f-H-f.*)^**** 4 4 4 4+VXX 6ol *8X XXXXXXXX**X4-4-4 4444+)(444 ••**•* X444** + XX X" vs *XXXXXXgaX*-*-4-*-«-4f 4 4+4 4 4 + 4 4 4-4 4 4*X4f •••••4 -f ' • 6 ) • 8yxxxxxxxxx-x+444 4 4.x+-**4 4+-»yx4X+4 4-»4txyy---+-+4-4-44 - 444444 tot • 88aXXXX^^4"*^'444+ — 4y4^*44'*4X44X44 + 444X'44* — 4.+"* ss 4€XXXXy>XX-4 4 44+44.X4.44-44»fXX4X4«-44*4+X+ -- + 4 4 95 »8xyfla«-xx-4 4*4»4+4 4 4f^4 4-xx+«-44 44 4i.yxx-4 LZi • X0XXXX* 44-4 44 44 444 4 4 444*44444+-4444XXX-4 9£t *88X*«-*r—4«—-4-++4**X*'**+4-4'4 + 4*4TX4-X • - SI • 88 44 4~4 4-4 4 44t4-44»-4-VX4-XX*X-t-4 4 4*1'XXX• •— 4-' . 90 I • 84-4 4 + 44 44 44444 44XXX-»XX*X4 4-44-»-) Appendix 5 . Simplified similarity matrix run 2A. •ax- ZZl iZv * - 92 V tct i it, JZ ci PQ 9 S2 »-+"'--•• 4 H--4-44 + -4 • *• S? , «• 4--* + f4-4-44444"* 4 oov vo• ?v rA oz ON 12 • 44 - • 4-4-• 4 444 52 V V »-44 4-4-44 4--4-44444.4 44* 4 4-- 621 X 4 ' " -4 --* 44-44 X4 44+4444X-- *Z HCO • 4 4 44 4 X-4 + --4 +-' •• 16 O V • 4 44-+444-X4 + 44-4- '** 6Z »-V—4-44X-444 444 + 4 + 4444 — 4 82 oj 01 fi CN »-44+ + 444 + 444-XXX"-- + 4+444 44- • LZ CO OO • 4-44-44XXXXXX4-XX** 201, bO V ZY »4-44.4X4X*-XXX-"--+44-44 4 44 • +X4+XXXX44* " ' • • • 9Z o V "•XX + 4-X4XXX+* '4 * — — 4 6! H 444XXXXX XX '-4- 4 4444 ' 01 H oo • X XXX4- X44 * 4444—* * * IS— - O • • XXX 4-44-X* ' oy I *•-•.+•+•-+4-4 •• • • • t-t•--'- ••--.»•+ 4 4 .4. ••+—-_ t • • - • •-* ' 4 - - * --4 + -t1. + +4 + • + • 69 »--*44--x-xy-44-• •• 4 + + -+-- > + x•+xx++ - a oil + • -•»«• +4-4-+44xXx+—+• 58 £1 • X*44—4 4VX 44--444-4+44 44XXXXX4+4+444- u 91 • +X + 44 * -4 44.---4 + --4-4 4 4-4 * 4 4»XX4 + 44X -4- 21 --* 4-. -+--+-+4V--+4 *+ + 4X44y4.4-44i.4_- • 28 • 44-' -44 4-4 4- 44-X+4 —4X XXX 44 4 44 8 L • + -,._--4 + -4+XX+++4 44+4 <44 4+ X 44 4 +4 4XX-4 4-4 4 444—44X-44--. 4 IS4-4 - + X4-44XXX4-4+XXX XX+4-+ + +X -- 99 »-4----44-»XX-XX4 4- + 4- 4+--•+-- 4-4.4 4XX4.4t4 4+-44 + *-4-f+—4+4-44XXX4+44-4-X-4.4'-4--4 4*4 * XX 44 + X+-4-4 +' •4X XXXXX4on X ' X4 tl • • 4X4 444 44-»44 XX 4+X X X 4 4 4+4 4+4 4 4 X X XX X X+44 4 4I X4 4 * 44 • +-++- • "** 4-4X4 4XXXX4-X+"4XXX) Appendix 7 List showing the five nearest neighbours for each OTU with the similarities expressed as percentages. OTU 1 2 4 5 OTU % OTU % OTU % OTU * OTU $ 1 132 94.0 • 130 89.7 133 88.7 131 83.0 25 80.1 2 5 97.7 if 97.7 3 95.7 6 92.3 94 88.4 3 4 96.1 5 95.9 2 95.7 6 93.7 94 89.4 4 5 99.2 2 97.7 3 96.1 6 93.4 94 85.2 5 4 99.2 2 97.9 3 95.9 6 93.2 94 89.4 6 3 93.7 94 93-5 93. ^ 5. 93.2 2 92.3 7 5 85.7 4 85.2 3 85.0 2 84.3 94 84.2 8 9 95.2 135 79.8 69 79.4 33 78.7 36 77.3 9 8 95.2 135 80.5 60 77.9 59 77.8 129 77.7 10 102 88.5 35 88.3 38 85.9 36 85.5 93 85.5 11 99 95.6 101 95-5 98 94.8 15 94.3 13 93.3 12 18 85.7 14 8if.8 .17 83.9 98 83.8 99 82.6 13 101 96.3 98 95.4 99 95.3 14 93.7 15 93.7 14 98 97.2 101 96.9 99 96.7 15 94.7 127 94.0 15 99 97.0 98 96.5 101 96.0 14 94.7 11 94.3 16 98 '90.7 101 90.7 99 90.6 14 90.0 19 89.6 17 18 96.7 19 88.1 99 84.2 101 84.0 12 83.9 18 17 96.7 19 90.1 98 86.5 99 86.0 12 «5.7 19 101 92.4 98 92.1 13 91.8 99 91.3 15 91 .c 20 21 90.7 34 83.9 125 83.4 39 83.3 102 £2.6 21 20 90.7 125 88.4 34 85.6 128 85.2 127 34.0 22 52 90.7 5T 83.6 103 88.6 107 83.4 109 88.4 23 33 84.1 38 82.3 30 82.1 3': 81.5 25 30.5 24 19 87.3 101 87.3 99 86.5 9? 86.2 07 84.9 25 135 85.7 35 3 4.7 38 8*4.4 81 S3.8 69 83.6 26 35 91.3 42 91.0 30 89.8 34 88.4 36 87.9 27 31 89.9 32 83.2 33 88.1 23 87.5 30 86.4 28 30 87.9 27 87.5 26 36.4 16 35.0 31 35.0 Appendix 7 ( continued ). OTU 1 2 3 4 5 CPU % OTU * OTU % OTU % OTU % 29 40 87.8 42 86.4 36 85.8 38 85.6 30 84.6 30 32 92.2 36 92.2 39 92.0 33 91.8 31 91.6 31 32 95.2 33 93.3 34 92.6 30 91.6 41 91.2 32 31 95.2 33 94.5 34 92.8 30 92.2 36 90.7 33 32 94.5 31 93.3 30 91.8 39 90.5 36 89.8 34 32 92.8 31 92.6 35 92.4 30 91.6 38 90.1 35 36 95.0 38 94.8 34 92.4 26 91.3 102 90.5 38 96.0 35 95.0 30 92.2 32 90.7 37 90.5 37 38 92.8 40 90.8 36 90.5 102 89.8 35 89.2 38 36 96.0 35 94.8 4o 93.4 37 92.8 30 91.3 39 30 92.0 33 90.5 34 88.9 32 88.5 36 87.3 40 38 93.4 37 90.8 35 90.3 36 90.1 29 37.8 41 31 91.2 33 88.6 32 88.4 35 87.3 On -> 42 26 91.0 w «/ 97 87.7 34 87.5 32 87.2 43 45 90.6 54 90.5 74 90.4 105 90.2 58 90.1 44 108 86.0 1"C 84.' 59 83.7 60 83.5 57 83.1 45 . 106 95 105 93.7 109 92.9 53 91.8 49 91.5 45 50 95- ' 47 93.0 49 92.4 103 92.3 107 90.5 47 46 9} 0 50 90.1 136 39.9 45 89.6 106 89.5 48 51 94.0 •>09 93.0 49 92.9 52 92.3 107 91.6 49 51 97.3 1C9 95-4 107 95.2 52 95.0 103 94.1 50 46 95.1 49 93.6 103 92.8 52 91.7 107 91.1 51 49 97.3 109 97.1 107 96.7 52 96.4 103 94.9 52 109 96.8 51 96.4 107 95.4 49 95-0 53 94.2 53 52 94.2 103 93.0 55 92.6 107 92.5 45 91.8 54 55 91.7 51 91.4 52 90.3 43 90.5 53 90.4 55 56 95.4 137 <54.1 104 93.9 138 93.2 53 92.6 56 137 96.9 55 95.4 138 92.0 51 91.1 105 91.1 57 112 91.3 135 51.1 60 89.2 61 89.0 b? 88.8 58 105 94.6 59 92.5 137 92.5 106 92.4 109 92.4 59 CO 93.3 137 92.9 58 92.5 104 91.5 75 91.2 279 Appendix 8 ( continued )« OTU 1 2 3 4 5 OTU * OTU * OTU % OTU % OTU % 60 59 98.3 58 91.8 137 91.4 104 90.5 56 90.1 61 62 98.8 112 93.5 116 93.4 117 93.1 66 92.7 62 61 98.8 112 93.7 116 93.1 117 92.9 65 92.7 63 66 94.5 116 94.1 61 92.4 62 91.9 64 91.8 64 118 93.4 66 93-1 68 92.6 119 92.2 62 92.0 65 62 92.7 61 92.4 93 91.6 112 90.9 119 90.9 66 116 95.2 63 94.5 117 94.4 112 94.1 64 93.1 67 66 92.9 88 92.8 116 91.7 117 91.3 112 90.7 68 64 92.6 118 92.5 117 92.2 119 92.2 66 91.8 69 135 89.9 112 86.8 37 86.7 118 86.3 92 85.8 70 92 93-^ 66 92.1 114 91.2 118 91.0 67 90.5 71 73 94.7 76 93.2 91 90.9 59 90.2 113 90.2 72 76 92.8 79 91.2 1J> 91.1 78 90.7 77 89.6 73 71 94.7 ?l- 3 72 91.1 78 89.4 85 87.0 74 75 95.^ 4"* 90.'; 90 89.6 91 89.3 71 88.4 75 ' 74 95.4 "SO 92.4 111 91.8 119 91.5 59 91.2 T> 76 71 93.2 ( - 92.1 73 92.2 90 90.6 75 88.6 77 79 94.'i 89 93.9 81 9 2.7 78 91.3 84 91.3 73 82 92..^ 77 91.3 81 35 90.9 72 90.7 79 77 94,0 84 93.7 89 93.6 35 92.6 72 91.2 80 90 94.8 120 91.1 75 87.9 76 87.2 74 86.7 81 77 9 2.7 73 91-3 115 90.9 79 90.1 135 89.7 8 82 ? 92.8 77 91.2 83 88.4 85 88.3 81 87.4 83 79 90.9 77 90.2 78 90.1 89 39.5 . 72 89.1 84 85 96.2 79 93.7 90 92.2 77 91.3 73 90.7 85 84 9 6.2 79 9 2.6 78 90.9 77 90.7 90 89.1 86 93 91.6 116 91.5 112 91.3 62 90.6 61 90.5 87 135 93.3 89 92.2 115 91.-u 92 89.9 112 89. k 88 119 95.1 117 93.6 67 92.8 61 92.1 62 91.9 89 77 93.9 79 93-6 115 93.0 87 92.2 135 90.8 90 80 94.8 84 92.2 75 91.1 76 90.6 74 89.6 280 Appendix 7 ( continued ). OTU 1 2 3 4 5 OTU % OTU % OTU % OTU % OTU * 91 113 96 .'1 114 93.8 115 93.1 58 92.1 117 91.0 92 70 93.4 135 93.1 115 92.3 66 91.6 112 90.6 93 112 94.1 61 92.3 62 92.3 66 92.3 117 91.8 94 6 93.5 3 89.4 5 89.4 4 89.2 2 88.4 95 96 94.8 134 83.6 6 83.4 5 82.1 3 81.8 96 95 94.8 6 83.9 94 82.7 134 81.6 5 81.4 97 42 87.7 29 84.1 129 84.1 26 83.5 39 83.5 98 99 98.5 101 97.4 14 97.2 15 96.5 13 95.4 99 98 98.5 101 97.3 15 97.0 14 96.7 11 95.6 100 91 87.1 72 85.6 113 85.6 77 85.3 33 84.9 101 98 97.4 99 97.3 14 96.9 13 96.3 15 96.0 102 38 90.8 35 90.5 37 89.8 36 89.2 10 88.5 103 107 95.7 51 94.9 49 94.1 52 93.8 ICO 93.7 104 55 93.9 59 91.5 60 90.5 137 90.3 '•3 5C.-; 105 106 96.3 58 94.6 '45 93.7 137 93.0 49 92.8 106 105 96.3 45 95.4 138 93-2 49 93.0 109 92.6 107 109 99.1 51 9 6.7 103 95.7 52 95.4 95.2 108 107 93.5 109 93.5 51 91.3 4o 90.5 103 89.5 109 107 99.1 51 97.1 52 96.8 49 95.- 103 93.7 110 111 90.9 116 89.5 117 88.9 66 88,5 112 88.4 111 112 97.6 117 94.0 116 93-3 66 92.4 75 91.8 112 111 ' 97.6 117 95.5 116 95.2 66 94.1 . 93 94.1 113 91 96.1 114 93.8 58 91.6 115 91.2 71 •30.2 114 91 93.8 113 93.8 70 91.2 58 90.5 117 90.5 115 • 91 93.1 89 93.0 135 93.0 112 92.6 92 92.3 116 66 95.2 112 95.2 117 94.9 63 94.1 61 93.4 117 112 95.5 119 95.1 116 94.9 66 94.4 111 94.0 118 64 93.4 68 92.5 66 92.3 62 91.1 63 91.O 119 88 95-1 117 95.1 64 92.2 68 92.2 66 92.0 120 75 92.4 80 91.1 59 90.6 91 89.5 90 89.4 121 72 86.3 71 86.1 113 86.0 91 84.4 73 84.0 281 Appendix 8 ( continued )« 7 OTU 1 2 3 4 5 OTU % OTU * OTU * OTU % OTUl % 122 123 94.-1 124 92.2 90 79.6 15 79.4 14 79.3 123 124 97.7 122 94.1 129 79.9 15 79.1 40 79.1 124 123 97.7 122 92.2 15 81.0 14 80.9 98 80.0 125 21 88.4 98 86.3 101 86.2 14 85.4 33 85.2 126 102 81.5 34 81.1 32 79.8 38 79.6 91 79.5 127 128 97.3 14 94.0 99 91.8 101 91.6 98 91.3 128 127 97.3 14 92.0 99 91.4 101 91.2 98 90.9 129 34 86.7 26 86.1 35 85.8 32 85.2 127 84.7 130 133 96.0 132 93.7 131 93.3 1 89.7 99 83.0 131 130 93-3 133 89.5 132 87.7 101 86.4 99 85.7 13? 1 94.0 130 93.7 133 92.7 131 8 7.7 10 82.1 133 130 96.0 132 92.7 131 89.5 1 88.7 16 84.5 134 6 85.1 85.1 95 83.6 96 81.6 99 80.5 r 135 87 93.3 i2 93. • -.15 93.0 112 92.1 57 91.1 136 103 92.0 hu 91.6 107 90.7 51 90.6 109 90.6 137 56 96.9 133 94.6 - 55 94.1 105 93.0 59 92.9 138 137 94.6 93.2 106 93.2 107 92.7 109 92.7 282 Appendix S . Distribution of OTU's in the six and nine group schemes for clustering to maximise WGMS analysis. Six group scheme Nine group scheme Group 1. Group 1. 8 : 9 : 59 : 60 : 71 : 73 : 2 : 3 : if : 5 : 6 : 7 7b : 75 : 76 : 80 : 87 : 90 : 9b : 95 : 96 91 : 92 :10*f :113 :11*f :115 : 120 :135. Group 2. Group 2. 57 : 61 : 62 : 63 : 6b : 65 : 1 :130 : 131 :132 :133. 66 : 67 : 68 : 69 : 70 : 86 : 88 : 93 :110 :111 :112 :116 : 117 :118 :119. Group 3- Group 3» 1 : 10 : 11 : 12 : 13 : 1b : 122 :123 15 : 16 : 17 : 18 :' 19 -: 20 : 21 : 23 : 2b : 25 : 26 : 27 : 28 : 29 : 30 : 31 : 32 : 33 : 3b : 35 : 36 : 37 : 38 : 39 : bo : bl : b2 : 97 : 98 : 99 : 100 :101 :102 :122 :123 :12*f : Group b. 125 :126 :127 :128 : 129 :130 : 22 : b3 : b5 : b6 : K7 : bS 131 :132 :133. b9 : 50 : 51 1 52 : 53 : 5b Group km 55 : 56 : 58 :103 :10*f :105 2 : 3 : b : 5 : 6 : 7 : 106 :107 :108 :109 :136 :137 9b : 95 : 96 138. 283 Appendix 8 ( continued )« Six group scheme. Nine group scheme. Group 5* Group 5* 72 : 77 : 78 : 79 : 81 : 82 : 59 : 60 : 71 : 72 : 73 74 - 83 : 84 : 85 : 89 :121 • 75 : 76 : 77 : 78 : 79 : 80 : 81 : 82 : 83 : 84 : 85 : 87 : 89 : 90 : 91 : 92 :113 : 115 :120 :121 :135- Group 6. Group 6. 22 : 43 : 44 : 45 : 46 : 47 : 10 : 20 : 23 : 25 : 26 : 27 : 48 : 49 : 50 : 51 : 52 : 53 : 28 : 29 : 30 : 31 : 32 : 33 : 54 : 55 : 56 : 58 :103 :105 : 34 : 35 s 36 : 37 : 38 : 39 s 106 :107 :108 :109 :136 :137 : 40 : 41 : 42 : 97 :100 :102 : 138, 126 :129. Group 7» 57 : 61 : 62 : 63 : 64 : 65 : 66 : 67 s 68 : 70 : 86 : 88 : 93 :110 :111 :112 :116 :117 : 118 :119- Group 8. 8 : 9 : 44 : 69. Group 9- 11 : 12 : 13 : 14 : 15 : 16 : 17 : 18 : 19 : 21 : 24 : 98 : 99 :101 :125 :127 :128. / 284 Appendix • Distribution of OTU's in the four and five group schemes for clustering to maximise WGMS analysis run 2A. Four group scheme. Five group scheme. Group 1 . Group 1. 1 : 11 : 12 : 13 : 14 : 15 : 16 : 10 : 23 : 25 : 26 : 27 : 28 : 17 : 18 : 19 ::24 : 98 : 99 :101 : 29 : 30 : 31 : 32 : 33 : 34 : 127 :128 :130 :131 :132 :133. 35 : 36 : 37 : 38 : 39 : 40 : 41 : 42 : 97 :100 :102 :126 : 129. Group 2. Group 2. 2:3: 4 : 5 : 6 : 7 20 : 21 :122 :123 :124 :125. 94 :134. Group 3« Group 3- 10 : 23 : 25 : 26 : 27 : 28 : 29 : 1 :130 :131 :132 :133. 30 : 31 : 32 : 33 : 34 : 35 : 36 : 37 s 38 : 39 : 40 : 41 : 42 : 97 : 100 :102 :126 :129. Group 4. Group 4. 20 : 21 :122 :123 :124 :125. 11 : 12 : 13 : 14 : 15 : 16 : 17 : 18 : 19 : 24 : 98 : 99 : 101 :127 :128. Group 5* 2:3: 4 : 5 : 6 : 7 : 94 : 134. 285 Appendix 10 • Distribution of OTU's in the four and five group schemes for clustering to maximise WGMS analysis run 2B. Four group scheme. Five group scheme. Group 1. Group 1. 61 : 62 : 63 : 64 : 65 : 66 : 63 : 64 : 66 : 67 : 68 : 70 67 : 68 : 69 : 70 : 86 : 88 : 88 :110 :111 :116 : 117 :118 93 :110 :111 :112 :116 :117 : 119. 118 :119. Group 2. Group 2. 72 : 73 : 76 : 77 : 78 : 79 : 71 : 72 : 73 : 74 : 75 : 76 80 : 81 : 82 : 83 : 84 : 85 : 77 : 78 : 79 : 80 : 81 : 82 87 : 89 : 90 : 92 :115 :121 : 83 : 84 : 85 : 89 : 90 : 91 135. 113 :114 :121 • Group 3- Group 3. 22 : 45 : 46 : 47 : 48 : 49 : 43 : 55 : 56 : 58 : 59 : 60 50 : 51 : 52 : 53 : 54 : 55 : 104 :105 :120 :137 :138. 56 :103 :105 :106 :107 :108 : 109 :136 :137 :138. Group 4. Group 4. 43 : 44 : 57 : 58 : 59 : 60 : 22 : 44 : 45 : 46 : 47 : 48 71 : 74 : 75 : 91 :104 :113 : 49 : 50 : 51 : 52 : 53 : 54 114 :120. 103 :106 :107 :108 :109 :136 Group 5* 57 : 61 : 62 : 65 : 69 : 86 87 : 92 : 93 :112. :115 :135 f I co © © FN. in CM to to 0 W'. r~ to N?