Tentang Taksonomi

MELALEUCAS
THEIR BOTANY, ESSENTIAL OILS AND USES
Joseph J. Brophy, Lyndley A. Craven

and John C. Doran
MELALEUCAS
THEIR BOTANY, ESSENTIAL OILS AND USES
Joseph J. Brophy
School of Chemistry, University of New South Wales
Lyndley A. Craven
Australian National Herbarium, CSIRO Plant Industry
John C. Doran
Australian Tree Seed Centre, CSIRO Plant Industry
2013
The Australian Centre for International Agricultural Research (ACIAR) was

established in June 1982 by an Act of the Australian Parliament. ACIAR operates
as part of Australia's international development cooperation program, with a
mission to achieve more productive and sustainable agricultural systems, for
the benefit of developing countries and Australia. It commissions collaborative
research between Australian and developing-country researchers in areas
where Australia has special research competence. It also administers Australia's
contribution to the International Agricultural Research Centres.
Where trade names are used this constitutes neither endorsement of nor
discrimination against any product by ACIAR.
ACIAR MONOGRAPH SERIES

This series contains the results of original research supported by
ACIAR, or material deemed relevant to ACIARs research and
development objectives. The series is distributed internationally, with
an emphasis on developing countries.
Australian Centre for International Agricultural Research (ACIAR) 2013

This work is copyright. Apart from any use as permitted under the
Copyright Act 1968, no part may be reproduced by any process without
prior written permission from ACIAR, GPO Box 1571, Canberra ACT 2601,
Australia, aciar@aciar.gov.au
Brophy J.J., Craven L.A. and Doran J.C. 2013. Melaleucas: their botany, essential
oils and uses. ACIAR Monograph No. 156. Australian Centre for International
Agricultural Research: Canberra. 415pp.
ACIAR Monograph No. 156
ACIAR Monographs ISSN 1031-8194 (print), ISSN 1447-090X (online)
ISBN 978 1 922137 51 7 (print)
ISBN 978 1 922137 52 4 (online)
Technical editing by Mary Webb, Canberra
Design and typesetting by Sam Highley, Canberra
Printing by CanPrint Communications, Canberra
Cover: The crimson bottlebrush, Melaleuca citrina, a common species in coastal
and subcoastal eastern Australia, and widely cultivated as an ornamental shrub
(Photo: Sam Highley)
Foreword
The genus Melaleuca currently comprises nearly 300 species distributed in Australia
and South-East Asia. Melaleucas have been used for many purposes in Australia,
including brushwood fencing and as ornamental trees and shrubs for gardens and
street planting. They have also been used in farm shelterbelts and for rehabilitating
salt-affected lands. In South-East Asia, melaleuca fuelwood is harvested from natural
forests, melaleucas are planted for poles and posts on potentially acid-sulfate soils, and
research is being undertaken into the suitability of their wood for fibre. Extraction of
essential oils from the foliage of three species (Melaleuca alternifolia, M. cajuputi subsp.
cajuputi and M. quinquenervia) is the basis of industries in Australia and elsewhere,
and is a key contributor to several local economies.
The Australian Centre for International Agricultural Research (ACIAR) has previously
supported collaborative research between scientists in Australia, Indonesia and
Papua New Guinea to provide the basis for local essential oil industries. It is therefore
appropriate that it should take the initiative to publish a comprehensive account of
melaleucas to further assist development of economic uses of these species. I anticipate
that this book will be very beneficial to those planning and funding research on
Melaleuca species, especially in Australia and the AsiaPacific region. It should also
be useful to people involved in reforestation and agroforestry who require information
on species that can produce economic products and are capable of reasonable growth
under poor environmental conditions.
The authors of this book are leading scientists in their respective fields and ACIAR
appreciates their commitment to preparing such a scholarly and comprehensive
account of melaleucas. This book is the first serious attempt to compile a consolidated
account of the taxonomy, essential oils, silvicultural characteristics and utilisation of
melaleucas. Detailed descriptions and natural distribution maps of all the species are
included in this volume, many of which are being published for the first time. The
authors made extensive field collections of melaleucas and analysed their essential
oils to fill large gaps in published information. As a result, several species worthy
of further study of their potential to produce economically important essential oils
were identified. A searchable database of the melaleuca oil profiles is provided on the
ACIAR website.
Foreword
Many people and organisations helped with the creation of this book, and are
mentioned in the Acknowledgments section. The Commonwealth Scientific and
Industrial Research Organisation (CSIRO) and the University of New South Wales
provided institutional support for this activity over many years. Special thanks go to
the Essential Oils and Plant Extracts Program of the Rural Industries Research and
Development Corporation (RIRDC) for funding a collecting expedition to Western
Australia. This allowed leaves to be collected from nearly 40 species not previously
sampled for analysis of their essential oils.
Nick Austin
Chief Executive Officer
ACIAR
Contents
Foreword.........................................................................................................................................................................................................3
Melaleuca species...................................................................................................................................................................................8
Melaleuca synonyms...........................................................................................................................................................................11
Preface.............................................................................................................................................................................................................13
1. Taxonomic history and systematics................................................................................................................................15
Historical context...............................................................................................................................................................................15
Studies based on morphological evidence..........................................................................................................................16
Incorporating DNA evidence in classification..................................................................................................................16
Current and future classification challenges..................................................................................................................... 17
2. Introduction to the genus Melaleuca............................................................................................................................19

General information.........................................................................................................................................................................19
Family and tribe..............................................................................................................................................................................19
Botanical name..............................................................................................................................................................................19
Common names............................................................................................................................................................................19
Ploidy....................................................................................................................................................................................................19
Number of species......................................................................................................................................................................20
Botanical features.............................................................................................................................................................................20
Habit and size................................................................................................................................................................................20
Bark......................................................................................................................................................................................................20
Foliage.................................................................................................................................................................................................20
Flowers................................................................................................................................................................................................21
Reproductive biology..................................................................................................................................................................21
Timing of flowering..................................................................................................................................................................... 23
Pollination and pollen biology.............................................................................................................................................. 23
Hybridisation.................................................................................................................................................................................. 25
Fruits.................................................................................................................................................................................................... 25
Seeds...................................................................................................................................................................................................26
Cotyledons.......................................................................................................................................................................................26
Contents
Geographical distribution and ecology................................................................................................................................26

Natural occurrence and ecology.........................................................................................................................................26
Locations of planted forests.................................................................................................................................................. 27
Tolerance of difficult conditions..............................................................................................................................................29
3. Uses............................................................................................................................................................................................................. 33
Ethnobotanical.................................................................................................................................................................................... 33
Ornamental, landcare, honey, bark and wood................................................................................................................. 34
Ornamental and amenityhorticultural use................................................................................................................ 34
Land rehabilitation...................................................................................................................................................................... 36
Brushwood fencing and related products..................................................................................................................... 37
Honey................................................................................................................................................................................................. 37
Bark...................................................................................................................................................................................................... 38
Wood................................................................................................................................................................................................... 38
Fuelwood................................................................................................................................................................................... 38
Posts, poles, stakes and sticks......................................................................................................................................... 39
Sawn wood............................................................................................................................................................................... 39
Woodchips...............................................................................................................................................................................40
Extractives..............................................................................................................................................................................................40
Non-volatile extractives..........................................................................................................................................................40
Foliar essential oils.......................................................................................................................................................................40
Commercially important oils..........................................................................................................................................40
Inter- and intra-specific variation................................................................................................................................42
Species by oil type................................................................................................................................................................42
Groundbreaking work on gene control of terpene biosynthesis in melaleucas................................. 45
4. Propagation, silviculture and management........................................................................................................... 47

Propagation........................................................................................................................................................................................... 47
Propagation by seed................................................................................................................................................................... 47
Vegetative propagation............................................................................................................................................................48
Silviculture and management....................................................................................................................................................49
Plantations for wood production........................................................................................................................................49
Plantations for production of essential oils..................................................................................................................50
Australian tea tree oil..........................................................................................................................................................50
Indonesian cajuput oil........................................................................................................................................................ 52
5. Pests, diseases and other limitations............................................................................................................................ 55

Pests and diseases............................................................................................................................................................................. 55
Other limitations................................................................................................................................................................................ 56
Weediness/biological control............................................................................................................................................... 56
Source of allergens...................................................................................................................................................................... 58
Conservation status......................................................................................................................................................................... 59
Prospects................................................................................................................................................................................................60
Contents
6. Conservation and prospects............................................................................................................................................... 59
Opportunities for wider use..................................................................................................................................................60

Caution..............................................................................................................................................................................................60
Advice is at hand..........................................................................................................................................................................62
7. Species accounts............................................................................................................................................................................. 63
Acknowledgments............................................................................................................................................................................395
Photograph credits........................................................................................................................................................................ 396
Appendix 1Oil types by species.........................................................................................................................................397

Appendix 2Species with potential for planting for a variety of uses in different
climatic zones....................................................................................................................................................................................... 403
References............................................................................................................................................................................................... 407
Melaleuca species
Species name
Page no.
Species name
Page no.
Species name
Page no.
Melaleuca acacioides
65
Melaleuca bracteosa
96
Melaleuca comboynensis
125
Melaleuca acuminata
66
Melaleuca brevifolia
97
Melaleuca concinna
126
Melaleuca acutifolia
68
Melaleuca brevisepala
98
Melaleuca concreta
127
Melaleuca adenostyla
69
Melaleuca bromelioides
99
Melaleuca condylosa
129
Melaleuca adnata
70
Melaleuca brongniartii
100
130
Melaleuca agathosmoides
71
Melaleuca brophyi
101
Melaleuca
conothamnoides
Melaleuca alsophila
72
Melaleuca buseana
102
Melaleuca cordata
131
Melaleuca alternifolia
73
Melaleuca caeca
103
Melaleuca cornucopiae
132
Melaleuca amydra
74
Melaleuca cajuputi
104
Melaleuca croxfordiae
133
Melaleuca apodocephala
75
Melaleuca calcicola
106
Melaleuca ctenoides
134
Melaleuca apostiba
76
107
Melaleuca cucullata
135
Melaleuca araucarioides
77
Melaleuca
calothamnoides
Melaleuca cuticularis
136
Melaleuca arcana
78
Melaleuca calycina
108
Melaleuca dawsonii
137
Melaleuca argentea
79
Melaleuca calyptroides
109
Melaleuca dealbata
138
Melaleuca armillaris
80
Melaleuca campanae
110
Melaleuca deanei
139
Melaleuca aspalathoides
82
Melaleuca camptoclada
111
Melaleuca decora
140
Melaleuca atroviridis
83
Melaleuca capitata
112
Melaleuca decussata
141
Melaleuca barlowii
85
Melaleuca cardiophylla
113
Melaleuca delta
142
Melaleuca basicephala
86
Melaleuca carrii
114
Melaleuca dempta
143
Melaleuca beardii
87
Melaleuca cheelii
116
Melaleuca densa
144
Melaleuca biconvexa
88
Melaleuca chisholmii
117
Melaleuca densispicata
145
Melaleuca bisulcata
89
Melaleuca ciliosa
118
Melaleuca depauperata
146
Melaleuca blaeriifolia
90
Melaleuca citrina
119
Melaleuca depressa
147
Melaleuca boeophylla
91
Melaleuca citrolens
120
Melaleuca dichroma
148
Melaleuca borealis
92
Melaleuca clarksonii
121
Melaleuca diosmatifolia
149
Melaleuca brachyandra
93
Melaleuca clavifolia
122
Melaleuca diosmifolia
150
Melaleuca bracteata
94
Melaleuca cliffortioides
123
Melaleuca dissitiflora
151
Melaleuca coccinea
124
Melaleuca eleuterostachya
152
Page no.
Species name
Page no.
Species name
Page no.
Melaleuca elliptica
153
Melaleuca holosericea
196
Melaleuca manglesii
238
Melaleuca ericifolia
154
Melaleuca howeana
197
Melaleuca marginata
239
Melaleuca eulobata
156
Melaleuca huegelii
198
Melaleuca megacephala
240
Melaleuca eurystoma
157
Melaleuca huttensis
200
Melaleuca megalongensis
241
Melaleuca eximia
158
Melaleuca hypericifolia
201
Melaleuca micromera
242
Melaleuca exuvia
159
Melaleuca idana
202
Melaleuca microphylla
243
Melaleuca fabri
161
Melaleuca incana
203
Melaleuca minutifolia
244
Melaleuca faucicola
162
Melaleuca interioris
205
Melaleuca monantha
245
Melaleuca ferruginea
163
Melaleuca irbyana
206
Melaleuca montana
246
Melaleuca filifolia
165
Melaleuca johnsonii
207
Melaleuca montis-zamia
247
Melaleuca fissurata
166
Melaleuca keigheryi
208
Melaleuca nanophylla
248
Melaleuca flammea
167
Melaleuca kunzeoides
209
Melaleuca nematophylla
249
Melaleuca flavovirens
168
Melaleuca laetifica
210
Melaleuca nervosa
251
Melaleuca fluviatilis
169
Melaleuca lanceolata
211
Melaleuca nesophila
252
Melaleuca foliolosa
170
Melaleuca lara
213
Melaleuca nodosa
253
Melaleuca formosa
171
Melaleuca lasiandra
214
Melaleuca ochroma
255
Melaleuca fulgens
172
Melaleuca lateralis
215
Melaleuca oldfieldii
256
Melaleuca genialis
174
Melaleuca lateriflora
216
Melaleuca orbicularis
257
Melaleuca gibbosa
175
Melaleuca lateritia
217
Melaleuca ordinifolia
259
Melaleuca glaberrima
176
Melaleuca laxiflora
218
Melaleuca orophila
260
Melaleuca glauca
177
Melaleuca lazaridis
219
Melaleuca osullivanii
261
Melaleuca glena
179
Melaleuca lecanantha
220
Melaleuca oxyphylla
263
Melaleuca globifera
180
Melaleuca leiocarpa
221
Melaleuca pachyphylla
264
Melaleuca glomerata
181
Melaleuca leiopyxis
222
Melaleuca pallescens
265
Melaleuca gnidioides
183
223
Melaleuca pallida
266
Melaleuca grieveana
184
Melaleuca
leptospermoides
267
185
224
Melaleuca paludicola
Melaleuca groveana
Melaleuca leucadendra
268
186
226
Melaleuca pancheri
Melaleuca
halmaturorum
Melaleuca leuropoma
Melaleuca linariifolia
227
Melaleuca papillosa
269
Melaleuca halophila
187
Melaleuca linearifolia
229
Melaleuca parviceps
270
Melaleuca hamata
188
Melaleuca linearis
230
Melaleuca parvistaminea
272
Melaleuca hamulosa
190
Melaleuca linguiformis
232
Melaleuca pauciflora
273
Melaleuca haplantha
191
Melaleuca linophylla
233
Melaleuca pauperiflora
274
Melaleuca hemisticta
192
Melaleuca longistaminea
234
Melaleuca pearsonii
276
Melaleuca hnatiukii
194
Melaleuca lutea
235
Melaleuca penicula
277
Melaleuca hollidayi
195
Melaleuca macronychia
236
Melaleuca pentagona
278
Melaleuca phoenicea
280
Melaleuca species
Species name
Melaleuca
Melaleuca species
10
Species name
Page no.
Species name
Page no.
Species name
Page no.
Melaleuca phoidophylla
281
Melaleuca sclerophylla
319
Melaleuca teuthidoides
356
Melaleuca phratra
282
Melaleuca sculponeata
320
Melaleuca thapsina
357
Melaleuca pityoides
283
Melaleuca seriata
321
Melaleuca thymifolia
358
Melaleuca platycalyx
284
Melaleuca sericea
322
Melaleuca thymoides
359
Melaleuca plumea
285
Melaleuca serpentina
323
Melaleuca thyoides
360
Melaleuca podiocarpa
286
Melaleuca sheathiana
324
Melaleuca tinkeri
361
Melaleuca polandii
287
Melaleuca shiressii
325
Melaleuca torquata
363
Melaleuca polycephala
288
Melaleuca sieberi
326
Melaleuca tortifolia
364
Melaleuca pomphostoma
289
Melaleuca similis
327
Melaleuca trichophylla
365
Melaleuca preissiana
290
Melaleuca societatis
328
Melaleuca trichostachya
366
Melaleuca pritzelii
291
Melaleuca sophisma
329
Melaleuca triumphalis
367
Melaleuca procera
292
Melaleuca sparsiflora
330
Melaleuca tuberculata
368
Melaleuca protrusa
293
Melaleuca spathulata
331
Melaleuca ulicoides
370
Melaleuca psammophila
294
Melaleuca spectabilis
332
Melaleuca uncinata
371
Melaleuca pulchella
295
333
Melaleuca undulata
373
Melaleuca pungens
296
Melaleuca
sphaerodendra
374
297
334
Melaleuca urceolaris
Melaleuca punicea
Melaleuca spicigera
375
298
335
Melaleuca uxorum
Melaleuca pustulata
Melaleuca squamea
376
299
337
Melaleuca venusta
Melaleuca pyramidalis
Melaleuca squamophloia
377
300
338
Melaleuca villosisepala
Melaleuca quadrifaria
Melaleuca squarrosa
378
301
339
Melaleuca viminalis
Melaleuca quercina
Melaleuca stenostachya
380
302
340
Melaleuca viminea
Melaleuca quinquenervia
Melaleuca stereophloia
382
304
341
Melaleuca vinnula
Melaleuca radula
Melaleuca stipitata
383
305
342
Melaleuca violacea
Melaleuca recurva
Melaleuca stramentosa
384
306
343
Melaleuca virens
Melaleuca rhaphiophylla
Melaleuca striata
385
307
344
Melaleuca viridiflora
Melaleuca rigidifolia
Melaleuca strobophylla
387
308
345
Melaleuca williamsii
Melaleuca ringens
Melaleuca styphelioides
389
309
347
Melaleuca wilsonii
Melaleuca rugulosa
Melaleuca subalaris
390
310
348
Melaleuca wimmerensis
Melaleuca ryeae
Melaleuca suberosa
391
311
349
Melaleuca wonganensis
Melaleuca sabrina
Melaleuca subfalcata
392
312
350
Melaleuca xerophila
Melaleuca salicina
Melaleuca subtrigona
393
313
351
Melaleuca zeteticorum
Melaleuca saligna
Melaleuca subulata
394
314
352
Melaleuca zonalis
Melaleuca sapientes
Melaleuca sylvana
Melaleuca scabra
315
Melaleuca systena
353
Melaleuca scalena
316
Melaleuca tamariscina
354
Melaleuca sciotostyla
318
Melaleuca teretifolia
355
Melaleuca synonyms
Synonym
Accepted name
Synonym
Accepted name
Astroloma marginatum
Melaleuca marginata
Callistemon pancheri
M. pancheri
Callistemon acuminatus
M. flammea
Callistemon pauciflorus
M. faucicola
Callistemon brachyandrus
M. brachyandra
Callistemon pearsonii
M. pearsonii
Callistemon brevisepalus
M. brevisepala
Callistemon phratra
M. phratra
Callistemon buseanus
M. buseana
Callistemon phoeniceus
M. phoenicea
Callistemon chisholmii
M. chisholmii
Callistemon pinifolius
M. linearis var. pinifolia
Callistemon citrinus
M. citrina (Curtis)
Dum.-Cours.
Callistemon pityoides
M. pityoides
Callistemon comboynensis
M. comboynensis
Callistemon polandii
M. polandii
Callistemon flavovirens
M. flavovirens
Callistemon pungens
M. williamsii subsp.
williamsii
Callistemon formosus
M. formosa
Callistemon glaucus
M. glauca
Callistemon pungens subsp.

fletcheri
fletcheri
Callistemon gnidioides
M. sphaerodendra
Callistemon hemistictus
M. hemisticta
Callistemon pungens subsp.

synoriensis
synoriensis
Callistemon lanceolatus
M. citrina (Curtis)
Dum.-Cours.
Callistemon pyramidalis
M. pyramidalis
Callistemon quercinus
M. quercina
Callistemon lazaridis
M. lazaridis
Callistemon recurvus
M. recurva
Callistemon linearifolius
M. linearifolia
Callistemon rigidus
M. linearis var. linearis
Callistemon linearis
M. linearis var. linearis
Callistemon rugulosus
M. rugulosa
Callistemon rugulosus var.

flavovirens
M. flavovirens
Callistemon sabrina
M. sabrina
Callistemon salignus
M. salicina
Callistemon serpentinus
M. serpentina
Callistemon shiressii
M. shiressii
Callistemon sieberi
M. paludicola
Callistemon speciosus
M. glauca
Callistemon suberosum
M. dawsonii
Callistemon subulatus
M. subulata
Callistemon macropunctatus M. rugulosa

Callistemon macropunctatus M. rugulosa
var. laevifolius
Callistemon megalongensis
M. megalongensis
Callistemon montanus
M. montana
Callistemon montis-zamia
M. montis-zamia
Callistemon nervosus
M. nervosa
Callistemon pachyphyllus
M. pachyphylla
Callistemon pallidus
M. pallida
Callistemon paludosus
M. paludicola
11
Melaleuca synonyms
12
Synonym
Accepted name
Synonym
Accepted name
Callistemon teretifolius
M. orophila
M. spectabilis
Callistemon viminalis
M. viminalis subsp.
viminalis
Melaleuca longistaminea
subsp. spectabilis
Melaleuca maidenii
M. quinquenervia
Callistemon viminalis subsp. M. viminalis subsp.

rhododendron
rhododendron
Melaleuca minutifolia subsp. M. monantha

monantha
Callistemon viminalis var.

minor
M. viminalis subsp.
viminalis
Melaleuca nervosa f. latifolia M. nervosa
Callistemon viridiflorus
M. virens
Melaleuca nervosa subsp.

crosslandiana
M. nervosa
Callistemon wimmerensis
M. wimmerensis
Melaleuca paludosa
M. glauca
Calothamnus suberosus
M. suberosa
Melaleuca smithii
M. quinquenervia
Melaleuca acacioides subsp.

alsophila
M. alsophila
Melaleuca styphelioides var.

squamophloia
M. squamophloia
Melaleuca acerosa
M. systena
M. irbyana
Melaleuca apodocephala
subsp. calcicola
M. calcicola
subsp. irbyana
M. pallescens
Melaleuca arenaria
M. tuberculata var.
arenaria
subsp. pallescens
M. dichroma
Melaleuca calycina subsp.

dempta
M. dempta
Melaleuca urceolaris var.

virgata
Melaleuca virgata
M. dichroma
Melaleuca cardiophylla var.

longistaminea
M. longistaminea
Melaleuca viridiflora var.

angustifolia
M. quinquenervia
Melaleuca citrina Turcz.
M. lutea
M. quinquenervia
Melaleuca coccinea subsp.

eximia
M. eximia
Melaleuca viridiflora var.

rubriflora
Metrosideros citrina
Melaleuca coccinea subsp.

penicula
M. penicula
M. citrina (Curtis)
Dum.-Cours.
Metrosideros decora
M. decora
Melaleuca coronicarpa
M. marginata
Metrosideros glauca
M. glauca
Melaleuca crosslandiana
M. nervosa
Metrosideros linearifolia
M. linearifolia
Melaleuca cymbifolia
M. halmaturorum
Metrosideros nodosa
M. nodosa
Melaleuca densa var. pritzelii M. pritzelii
Metrosideros pinifolia
M. linearis var. pinifolia
Melaleuca halmaturorum
subsp. cymbifolia
M. halmaturorum
Metrosideros quinquenervia M. quinquenervia
Melaleuca lanceolata subsp.

occidentalis
M. lanceolata

planifolia
M. lanceolata

thaeroides
M. lanceolata
Melaleuca lateriflora subsp.

acutifolia
M. acutifolia
Melaleuca lateriflora var.

acutifolia
M. acutifolia
Metrosideros rugulosa
M. rugulosa
Metrosideros saligna
M. salicina
Metrosideros viminalis
M. viminalis subsp.
viminalis
Metrosideros viridiflora
M. virens
Myrtus leucadendra
M. leucadendra
Petraeomyrtus punicea
M. punicea
Regelia punicea
M. punicea
Preface
Melaleuca is the basis of several industries in Australia and elsewhere. Predominantly
these industries are based on the extraction of essential oils from the foliage of three
species, Melaleuca alternifolia (but sometimes including minor use of M. linariifolia
and M. dissitiflora), M. cajuputi subsp. cajuputi and M. quinquenervia. An emerging industry is underway in South-East Asia, especially on potentially acid-sulfate
soils, where trees are being grown primarily for roundwood, and research is being
undertaken into the suitability of their wood for fibre. In view of the challenge to find
novel sources of commercially significant oils, we have been collecting samples over
the past three decades, and have been extracting and analysing the oils. In spite of this
activity, when this book was mooted, information on the oils of about 100 species was
still lacking. Dedicated fieldwork was undertaken in south-western Western Australia,
where most of the unsampled species occurred, and requests were made of individuals
and organisations that might have been in a position to assist. These efforts were successful and there are only one species and four infraspecific taxa for which there are
still no data on their essential oils. The majority of the oils reported in this volume have
not previously been described in the scientific literature. As a result of the fieldwork,
several species (e.g. M. halophila, M. hamata, M. ochroma) have been identified as
potentially valuable sources of essential oils and may warrant further investigation of
their oil content and yield.
As it is presently circumscribed, Melaleuca consists of
290 species with 37 infraspecific taxa. Work on the systematics of the genus in recent years has indicated that
Melaleuca may be made larger due to the inclusion of
several genera presently regarded as distinct. As this work
is at present incomplete, Melaleuca is treated in a conventional concept in this volume, although with the addition
of the species previously known as Callistemon. Many of
the species included in this volume have previously been
treated only skeletally and, for these, detailed descriptions
are being published for the first time. Distribution maps
are included for all species, and colour photographs of the
flowers are included for those we have been able to source.
Apart from essential oils, melaleucas have been used
for a wide range of purposes, from making brushwood
fencing, as ornamental shrubs, as shelterbelt species for
farmland, for rehabilitating salt-affected lands, for street

and park trees, and so on. One interesting potential use
for M. bracteata is as a source of water-soluble betaines
compounds that act as osmoprotectants against stress in
plants. These compounds may increase stress tolerance in
agricultural crops which in turn would increase national
income from agriculture.
A threat to the future health and genetic diversity of a
substantial number of Melaleuca species in eastern Australia is from Puccinia psidii sensu lato (synonym Uredo
rangelii). This exotic pathogen has the common name of
myrtle rust in Australia but it is known as guava or eucalyptus rust elsewhere, with origins in Brazil. Myrtle rust
targets species of the family Myrtaceae, including Melaleuca. First observed in Australia on the central coast of
New South Wales in 2010, it has now spread from Victoria
13
Preface
to northern Queensland. In susceptible plants, young

spore-covered leaves and shoots become curled and distorted and severe infection can cause shoots to die, causing
these plants to become stunted after repeated infections.
In the worst cases, death of the whole plant can occur after
repeated destruction of new growth. As this book goes to
press, this disease is of concern to all with an interest in
the conservation and sustainable use of Australian plants
of the family Myrtaceae.
Joe Brophy, Lyn Craven, John Doran
September 2013
14
Taxonomic history
and systematics
Historical context
Melaleuca is almost entirely Australian in its distribution yet the first of its species to
be formally described, Melaleuca leucadendra, was based on material from Ambon, in
present-day Indonesia. Georgius Everhardus Rumphius, a merchant with the Dutch
East Indies Company, compiled a detailed account of many of the plants growing in
the Malesian biogeographical region but this was not published until 1741; this important work has recently been translated into English and published with annotations
(Rumphius 2011). The plant we now know as Melaleuca leucadendra was called Arbor
alba by Rumphius. Rumphius 1741 publication predated the accepted starting point for
the scientific botanical nomenclature of flowering plants and the formal description of
the species occurred in 1754 when the Swedish botanist Carolus Linnaeus gave it the
name Myrtus leucadendra, taking his descriptive data from Rumphius work (Linnaeus
1754). Subsequently, Linnaeus realised that his Myrtus leucadendra did not have very
much in common with the other species of Myrtus and in 1767 he described the genus
Melaleuca to accommodate this plant (Linnaeus 1767).
The nomenclature applied to the first endemic Australian
melaleucas to be described was inconsistent due to a lack
of appreciation of the relationships of the species. Doubtless this was due to the limited numbers of specimens that
had been collected and consequent uncertainty as to how
the genera of Myrtaceae should be circumscribed. Several
species, such as M. armillaris and M. decora, described in
1788 and 1796, respectively, initially were placed in Metrosideros. In other cases, the author recognised a relationship
with Linnaeus Melaleuca and the species was placed in
that genus, e.g. M. ericifolia and M. gibbosa, described
in 1797 and 1806, respectively. From the perspective of
having a foundation on which to build new knowledge
of Melaleuca species, George Benthams treatment of the

genus in his Australian flora (Bentham 1867) provided the
first comprehensive summary of the species known, and
those species that had been described in Metrosideros were
then brought into Melaleuca. Benthams important account
enabled later workers to identify their materials and thus
determine if there were additional species that should be
described. During the next 100 years, many species were
added to the genus, including two from New Caledonia
(M. brongniartii and M. gnidioides) and one from Lord
Howe Island (M. howeana).
Melaleuca was circumscribed by Bentham (1867) in part
as having stamens united in bundles opposite the petals.
15
1. Taxonomic history and systematics
The related genus Callistemon was distinguished from

Melaleuca by Bentham as having free stamens, although
he noted that the stamens of Callistemon speciosus (now
M. glauca) were often in bundles. Most of the species
placed in Callistemon are Australian but five species were
also described from New Caledonia, two of which have
stamens in bundles; one wonders why these were not therefore placed in Melaleuca. The recognition of Melaleuca and
Callistemon as separate genera had been regarded as being
artificial from the time of the description of the latter.
When describing Callistemon, Robert Brown (1814, p. 547)
wrote: The maximum of Melaleuca exists in the principal
parallel, but it declines less towards the south than within
the tropic, where its species are chiefly of that section
which gradually passes into Callistemon, a genus formed of
those species of Metrosideros that have inflorescences similar to that of Melaleuca, and distinct elongated filaments.
Ferdinand Mueller, who was well acquainted with both
genera in the field and in the herbarium, also regarded Callistemon as being artificial (Mueller 1864). Bentham may
in fact have had his own doubts about the distinctness of
Callistemon, for he commented in Flora Australiensis that
Callistemon passes gradually into Melaleuca, with which
F. Mueller proposes to reunite it (Bentham 1867, p. 118).
The majority of authors have accepted the two genera as
being distinct but the nineteenth century botanist Henri
Ernest Baillon included Callistemon and two other genera
(Conothamnus and Lamarchea) in Melaleuca, recognising them at sectional level (Baillon 1876). In 1998, Lyn
Craven and John Dawson transferred the New Caledonian
Callistemon species to Melaleuca, as they considered the
endemic New Caledonian species of the complex should be
placed in the same genus (Craven and Dawson 1998). Craven (2006) then discussed morphological evidence relevant
to the separation of the two genera and transferred all the
accepted Australian species of Callistemon to Melaleuca.
Studies based on
morphological evidence
Surprisingly for a genus of nearly 300 species, few revisionary-level treatments of species groups or of prescribed
geographical regions within Australian Melaleuca have
been published. Perhaps fittingly, in view of the ecological and/or economic significance of its several species, the
first was an account of the broad-leaved paperbarks, the
M. leucadendra species group, by Stan Blake (Blake 1968);
these species are a common component of savannah and
woodland communities in northern and north-eastern
Australia, south-eastern Malesia and New Caledonia.
John Carrick and Kosmyn Chorney (Czornij) published a
revisionary-level account of the South Australian species
16
of Melaleuca in 1979 which gave an insight into arid-zone

species of the genus (Carrick and Chorney 1979). Norm
Byrnes in 1984 published the first part of a concise revision
of the Melaleuca species of northern and eastern Australia
(Byrnes 1984, 1985, 1986). Several of these species were
removed from Melaleuca in 1989 as a result of the resurrection of Asteromyrtus (Craven 1989). Asteromyrtus is related
to Agonis and Leptospermum and presumably had been
included in Melaleuca only because its stamens are in bundles. Byrnes M. punicea was also removed from the genus
because of its anomalous androecium, gynoecium and seed.
Initially the species was placed in Regelia (Barlow 1987a)
but it was subsequently placed in the new genus Petraeomyrtus (Craven 1999), as the species was as anomalous in
Regelia as it had been in Melaleuca. However, following
recent investigations using molecular data, M. punicea is
treated in the present volume under Melaleuca (see below).
Bryan Barlow initiated revisionary studies of Melaleuca
in Canberra in the early 1980s. In 1986, Barlow published
the results of his studies of three species complexes: the
M. acacioides, M. tamariscina and M. minutifolia complexes (Barlow 1987b). Subsequently, the M. cuticularis
and M. lanceolata species groups were revised by Barlow
and Kirsten Cowley (Barlow and Cowley 1988) and the
M. fulgens and M. laxiflora species groups were revised
by Cowley and collaborators (Cowley et al. 1990). An
enumeration of the Australian species of Melaleuca sensu
stricto was published by Craven and Brendan Lepschi in
1999; this paper included an identification keythe first
key including all these species of the genus since 1867
(Craven and Lepschi 1999). Four species belonging to
the M. thymoides species groupM. lutea, M. pungens,
M. striata and M. thymoideswere not included in the
1999 enumeration, as Craven, at that time, considered the
species of this group warranted separate generic status;
however, in the present volume they are again included
as melaleucas.
Incorporating DNA
evidence in classification
Plant classification until recently has been based largely
upon morphological evidence, utilising data from anatomy,
cytology, chemistry (secondary metabolites) and any other
sources where these have been available. The overarching objective of classification has been to group plants
according to their presumed natural relationships, with
putatively closely related species classified together. The
technological advances that have permitted sequencing
of DNA (deoxyribonucleic acid), together with development of computer programs for detecting related species,
have enabled biologists to study the genetic relationships
agreement with the nuclear DNA phylogenies of Ladiges et

al. (1999) and Brown et al. (2001); these were based upon
data derived from a smaller number of species than in the
study of Edwards et al. (2010) but the sampling was drawn
from most of the various genera of the complex.
Current and future

classification challenges
The taxonomic implications of the DNA studies are that
one either includes all the related genera within Melaleuca
or one retains the existing segregate genera and recognises
many new ones, perhaps 10 or more. Given the lack of
distinctive morphological features available to differentiate many of them, such new genera would not be readily
recognisable. Consequently, Melaleuca will be enlarged to
include Beaufortia, Calothamnus, Conothamnus, Eremaea,
Lamarchea, Phymatocarpus and Regelia (L.A. Craven, R.D.
Edwards and K.J. Cowley, in preparation). The species
treated in the present volume are those that accord with
the conventional concept of Melaleuca and also include
M. punicea, the M. thymoides group and those species
previously placed in Callistemon.
Morphological evidence also has been a primary source
of data for developing species concepts in Melaleuca. It
is unlikely that this situation will change dramatically in
the foreseeable future but the few studies that have been
made using DNA data have shown that this will be a very
powerful tool for taxonomists. Linda Broadhurst et al.
(2004), utilising nuclear restriction fragment length polymorphism (RFLP) data derived from a range of Western
Australian populations of the broombush (M. uncinata)
complex (and including one South Australian population from the type locality of M. uncinata), found that
phylogenetic analysis showed the sampled populations
represented seven species of the broombush complex. The
results of the phylogenetic analysis were congruent with
those of a parallel morphological study that encompassed
the whole of the broombush complex in Western Australia
(Craven et al. 2004).
The broad-leaved paperbark group (M. leucadendra
group) is an ecologically important component of vegetation in northern and eastern Australia. Conventionally it
has been regarded as a taxonomically difficult group and,
in the mid nineteenth century, it was treated by Bentham
(1867) as a single species with numerous varieties. Blake,
with an insight gained from studying the complex in the
field, published a detailed and workable account of the
group that, despite several new species being described
since, remains the most useful guide to the group (Blake
1968). One of the broad-leaved paperbarks, M. quinquenervia, is a common wetlands tree in eastern Australia and
of plants in detail for the first time. The consequences of

such studies for taxon delimitation, and thus classification,
are varied. In some cases, generic circumscriptions are supported by analysis of DNA data and there are no changes
to the classification, and hence nomenclature, of a genus.
In other cases, the current classification is not supported,
with part or all of the species sampled falling (or nesting)
within one or more other genera. There is no automatic
procedure to be adopted in such cases. Typically, if all the
species of a genus or group of genera nest within a single
genus, a decision has to be made to either combine all the
genera into one or to break the original single genus into
several. This clearly has implications for nomenclature. In
other cases, some of the species of a genus are found to nest
within another, with the remaining species still comprising
a distinct group, and in those cases the nested species are
transferred to the other genus. This still has implications
for nomenclature, especially if the name of the genus is
transferred along with the nested genus, as the remaining
species will then require a new generic name.
Species of Melaleuca and the morphologically closely
related genera Beaufortia, Calothamnus, Conothamnus,
Eremaea, Lamarchea, Phymatocarpus and Regelia, and
including M. punicea (Petraeomyrtus), the four species of
the M. thymoides group, and several Australian and New
Caledonian species of Melaleuca that formerly had been
placed in Callistemon, were included by Robert Edwards
and his collaborators in an analysis of chloroplast DNA
data (Edwards et al. 2010). Chloroplast DNA is maternally
inherited, and phylogenies derived from chloroplast DNA
data give a good estimate of the maternal family tree.
The results of Edwards et al. (2010) showed there were
three major groupings or clades, each of which contained
species of Melaleuca sensu stricto. The whole Melaleuca
group itself formed a well-supported clade relative to the
outgroup taxa included in the analyses. In one of the three
clades are species of the M. leucadendra, M. acacioides,
M. scabra and M. thymoides groups, two New Caledonian
species previously placed in Callistemon, M. punicea,
and representative species of Beaufortia, Calothamnus,
Conothamnus, Eremaea, Lamarchea, Phymatocarpus
and Regelia. In a second clade are representatives of the
M. cuticularis, M. fulgens and M. laxiflora groups, together
with Australian species previously placed in Callistemon
and several species not placed within a particular grouping. The third clade contains members of the M. bracteata,
M. cuticularis, M. lanceolata and M. minutifolia groups,
together with other species not allocated to a particular
grouping and the morphologically anomalous M. foliolosa.
The incorporation of nuclear DNA data in phylogenetic
studies enables the paternal contribution to be assessed;
using only the maternal chloroplast data could give a
biased result. The broad structure of the inferred chloroplast DNA phylogeny given in Edwards et al. (2010) is in
17

18
also occurs naturally in Papua province of Indonesia,

Papua New Guinea and New Caledonia, and is a major
woody weed in Florida, United States of America (USA).
This species is harvested for its essential oil, niaouli oil, in
New Caledonia and also in Madagascar, where it is cultivated for this purpose. In a study using DNA sequence
data from two chloroplast and two nuclear regions, Cook
et al. (2008) found that the genome of M. quinquenervia
contained alleles that link the species to several other
broad-leaved paperbark species and that there was regional
sharing of chloroplast haplotypes, indicating introgression
or lineage sorting. This has significance for biological control studies of M. quinquenervia as it will be necessary to
match the genetics of the weed populations with naturally
occurring populations in Australia when seeking control
agents. A significant conclusion of the Cook et al. (2008)
study was that species boundaries within the complex were
not clear. This work is being extended by Robert Edwards,
drawing upon a comprehensive sampling of the complex
across northern Australia and the results should be of
interest to systematists and others.
The extent to which hybridisation has played a role
in the evolution of Melaleuca species is not known but
it may be important and occurs quite widely across the
genus (Craven 2006). DNA studies may assist in clarifying
species circumscriptions in the bottlebrush (Callistemon)
group, which is at least as difficult taxonomically as the
broad-leaved paperbarks. The bottlebrush M. paludicola
and its putative close relatives M. phratra, M. quercina,
M. sabrina and M. wimmerensis make up one group
worthy of investigation. The red-flowered Queensland
bottlebrushes M. hemisticta, M. lazaridis, M. montis-zamia
and M. pyramidalis are another, as there are populations
in southern Queensland that presently are not definitively
identifiable to one of these species and the whole complex
requires a comprehensive population genetics study of the
type conducted by Broadhurst et al. (2004) on members of

the broombush group. Similarly, the relationship between
M. citrina and M. subulata in southern New South Wales
and eastern Victoria needs investigation. Bill Molyneux
has described Callistemon forresterae, C. genofluvialis,
C. kenmorisonii and C. nyallingensis from this region
(Molyneux 1993, 1994, 1997, 2005) but these are regarded
by one of us (LAC) as being hybrids or hybrid derivatives
between M. citrina and M. subulata and are not accepted
as species in the conventional sense; consequently, they
are not included in the present volume. Apomixis in two
bottlebrush species was studied by James (1958) and this
genetic process may also be a factor in the evolution and
relationships of the bottlebrushes in eastern Australia
(Craven 2009).
It is clear there is need for an infrageneric classification
of Melaleuca, with closely related species being grouped
together in sections and subsections etc. Already there are
nearly 300 species in the genus and with the transfer of the
species flagged above there will be nearly 400; this is too
large a number to be left in an unstructured arrangement.
A preferred classification would be one based upon morphological data alone, but in the case of Melaleuca these are
not sufficient and it will be necessary to incorporate DNA
results in the synthesis. The DNA studies that have been
published to date, while sufficient to support the merger
of the genera of tribe Melaleuceae into a single genus, i.e.
Melaleuca, do not adequately resolve the clades that were
found and contain too few species to permit a classification to be prepared. Research presently underway at the
Australian National University, Canberra, using nextgeneration sequencing methods and a very much greater
sampling of species across Melaleuca sensu lato should give
information that will guide development of a robust classification (Mike Crisp and Bo Choi, pers. comm.).
Introduction to
the genus Melaleuca

General information
Family and tribe
The genus Melaleuca is in the family Myrtaceae and tribe Melaleuceae.
Botanical name
The generic name is derived from the Greek melas, meaning black or dark, and leucon,
meaning whiteapparently alluding to the white branches and black trunk of the first
named species, M. leucadendra, the trunks of which are often blackened by fire.
Common names
Species with thick, spongy, peeling bark comprising many papery layers are commonly
referred to as paperbarks, with some qualifying adjective (e.g. silver-leaved paperbark,
M. argentea). In southern Australia, the common name honey myrtle is also well
established for many shrub-sized species (e.g. bracelet honey myrtle, M. armillaris). A
few have distinctive Aboriginal (e.g. winti, M. arcana) or locality (e.g. South Australian
swamp paperbark, M. halmaturorum) names, while some are referred to as tea tree
(e.g. black tea tree, M. bracteata), a common name shared with many species of Leptospermum. As explained above, this treatment of Melaleuca includes species previously
belonging to the genus Callistemon. They retain their common name of bottlebrush
which alludes to the resemblance of the flowers and emerging new growth to a kitchen
bottlebrush (e.g. crimson bottlebrush, M. citrina).
Ploidy
The great majority of Melaleuca species are diploid with
2n = 22 (Brighton and Ferguson 1976; Rye 1979). Polyploidy appears to be relatively infrequent in this genus,
with only a few recorded instances of aneuploidy
(2n + 2 = 24), triploidy (3n = 33), tetraploidy (4n = 44)

and hexaploidy (6n = 66) (James 1958; Brighton and
Ferguson 1976; Rye 1979) which have been linked with
hybridisation and apomixis. The apomictic species, M. linearis, may have populations that are diploid, triploid or
tetraploid (James 1958).
19
2. Introduction to the genus Melaleuca
Number of species
Bark
Melaleuca, as circumscribed in this work, comprises 290

species (327 entities inclusive of infraspecific taxa). As
such, it is the third-largest genus of Myrtaceae after Eucalyptus and Syzygium in the Australasian region.
The bark of the majority of species is of the papery type.

Here, the bark consists of thin, paper-like layers of cork
separated by thin fibrous layers, which may reach 5 cm in
thickness. The outer layers peel naturally to give a distinctive ragged, torn and unkempt appearance to the lower bole
(Figure 2A). In addition, a substantial proportion (c. 20%)
of Melaleuca species has hard, deeply furrowed, rough bark,
as exemplified by M. bracteata (Figure 2B). There is a further 20% of species where the bark is described as fibrous.
There is some overlap in bark types in some species. For
example, M. clarksonii is variously described as displaying
all three bark types in different individuals.
Botanical features
Habit and size
Melaleucas range from woody, multistemmed shrubs to
very large, single-stemmed trees of timber-producing
value. By far the majority of species are shrubs or small
trees less than 10 m tall, 40 of which do not exceed 1 m in
height. These ground-hugging types are found largely in
the south of Western Australia. Fifteen species have been
documented as being over 10 m in height, with seven of
these species exceeding 20 m. Boland et al. (2006) report
that M. cajuputi has been recorded with heights up to
46 m in the Northern Territory and is the tallest tree in
the region; similarly, M. leucadendra has been measured
to 43 m in northern Queensland (Figure 1).
Foliage
Melaleucas are evergreen and usually carry abundant
green, bluish-green, grey-green or silvery-grey foliage
unless drought or other stresses (e.g. salt) have stimulated
leaf abscission. Leaves are minute to large. In all, 60% of
recognised species have short (<30 mm long) to very short
(<10 mm) leaves, while the others have medium to long
leaves. Melaleuca leucadendra, with its narrow lanceolate
Figure 1.A substantial

Melaleuca leucadendra
on a well-watered site in
north-western Queensland
20
Figure 2.Bark types in Melaleuca: papery, as in (A) M. exuvia; and rough, as in (B) M. bracteata
leaves, produces leaves measuring 75270 mm in length,

representing some of the longest in the genus. Leaves are
entire, dorsiventral or isobilateral, usually coriaceous, flat,
concave, centric or semi-terete, sessile or petiolate. Leaf
shape is also highly variable, including elliptic, cordate,
falcate, lanceolate, linear, oblong, ovate, obovate, elliptic or
triangular. Leaves are cordate, attenuate, cuneate, truncate
or obtuse at the base. Mature leaf blades may be glabrous,
pubescent or woolly. They are carried in either alternate,
opposite, spiral, decussate or occasionally ternate (whorls
of three leaves) arrangement. Mature leaves are without
stipules and without a persistent basal meristem. Melaleuca leaves are usually obviously oil-gland dotted and
are aromatic or without marked odour. Leaf venation
is pinnate, longitudinal or longitudinal-pinnate and is
frequently obscure.
Flowers
Flowers are in spikes or clusters, or sometimes solitary;
the basic floral unit is a monad, dyad or triad; the calyx
lobes are five or rarely may be fused into a ring of tissue;
the petals are five; the hypanthium is fused to the ovary in
the proximal region only, or for up to three-quarters the
length of the ovary or, rarely, for almost all the length of the
ovary; the stamens are few to numerous, the filaments are
fused for part of their length into five bundles and inserted
on a staminal ring or free and not in bundles wherein the
filaments are inserted on the hypanthium apex with the
staminal ring obsolete, the anthers are dorsifixed (or rarely
basifixed) and versatile, with two parallel cells that open via
longitudinal slits; the ovary has three locules, the placentae
are peltate and axile-median to axile-basal, the ovules are
few to numerous (Figures 3 and 4).
Species vary widely in flower colour, with about half

of all species having filaments of white through cream to
yellow or green, while the others have pink, red or mauve
filaments. In many species, the contrasting yellow anthers
at the ends of these brightly coloured filaments result in a
spectacular floral display.
Reproductive biology
Melaleucas generally produce only morphologically
bisexual (hermaphroditic) flowers but this is not universal. About 160 species are always hermaphroditic, i.e.
all inflorescences have only bisexual flowers. Examples
are M. quinquenervia and M. viminalis. About 90 species are always andromonoecious, i.e. there are male and
hermaphroditic inflorescences on the one plant. Typically,
the male inflorescences are on the outside of the plant and
possibly serve as advertisements to potential pollinators,
while the hermaphroditic inflorescences are within the
canopy, where they may be somewhat protected from
predation but are still close enough to the outside of the
plant to attract a pollinator. Examples of andromonoecious species are M. gibbosa and M. uncinata. About 30
species include some plants that are hermaphroditic and
others that are andromonoecious. Examples of this type are
M. hamulosa and M. incana. One species, M. cornucopiae,
is particularly interesting as some plants are monoecious
(with both male and female inflorescences on the same
plant) and others are gynoecious (with female inflorescences only). The above data are based on the study of
herbarium specimens supplemented with observations
on living plants (L.A. Craven, unpublished data) and need
verification with additional field studies. However, some
general statements can be made about specific groups of
21
Figure 3.Flower and fruit of Melaleuca leucadendra, a species with its stamens in bundles (drawings by M. Fagg)
Figure 4.Flower and fruit of Melaleuca linearis, a species with free stamens (drawings by M. Fagg)
22
Timing of flowering
Flowering starts early in many species. For example,
M. alternifolia planted in breeding populations in northern
New South Wales set the first flower buds as early as 2 years
after planting. However, the first reasonable flowering
(defined as 45% of trees) did not occur until almost 4 years
from plantings within the species natural range (Doran
et al. 2002). In M. alternifolia, a cold winter (minimum
temperatures below 5 C) appears to stimulate floral bud
formation while good spring rains are needed to support
a good flowering and retention of the developing fruit
(Baskorowati et al. 2010a, c).
Flowering periodicity and intensity in Melaleuca species
are highly variable between species and sites. In general,
most species flower in the late winter spring summer
period, some peak in autumn and winter, and others
may flower all year round. The natural flowering pattern
can be disrupted when a species is planted outside its
normal range. For example, flowering of M. alternifolia
in its region of natural occurrence is at its peak in spring
(OctoberNovember) but the species flowers in winter
when planted in Western Australia. The period for the
morphological development of buds, flowers and fruit
leading to the development of mature seed also varies
between Melaleuca species. In M. alternifolia, a period
of 1618 months is required (Baskorowati et al. 2010a, c)
but in summer-flowering tropical species (e.g. M. leucadendra), geared to shed their seed in response to the next
summers monsoonal rainfall, a shorter period of about 12
months is typical.
experiments. Pollination is probably also effected by birds,

notably lorikeets and honeyeaters, which are often seen
visiting the flowers of bottlebrushes and broad-leaved
paperbarks. Fruit bats (family Pteropodidae) also feed on
flowering broad-leaved paperbarks and may be pollinators
for these species.
Thornhill et al. (2012) studied the pollen morphology
of 21 species of Melaleuca (including six species of the
former genus Callistemon) using scanning electron and
light microscopy. A general description from this work
follows, combining methods and descriptions of the two
relevant genera. Pollen grains were tricolporate (Figure 7),
except for some pollen of M. citrina (as C. citrinus) which
was tetracolporate. Pollen had a rugulate exine, except for
M. nesophila grains that had a granulate/scabrate exine.
Pollen sides were straight, or less commonly convex or
concave, and the colpal morphology was consistently parasyncolpate with angular colpi, except for some grains of
M. virens (as C. viridiflorus) which had arcuate colpi. Pollen
ambs were round or pointed, or less commonly notched or
flat. Colpal edges were smooth or occasionally rough and
interest. It appears that all the species of the broad-leaved

paperbark group are hermaphroditic, all of the broombush
group are andromonoecious, and most of the species allied
to M. alternifolia are hermaphroditic (M. linophylla may
be hermaphroditic or andromonoecious but this needs
verification in the field).
Baskorowati et al. (2010a, b, c) describe the reproductive biology of M. alternifolia (summarised in Figure 5)
which is an example of floral structure and development
in a hermaphroditic species.
Pollination and pollen biology

Melaleucas are mostly insect-pollinated. Hawkeswood
(1980), for example, showed that jewel beetles (Diadoxus
spp.) were the main pollinators of M. pauperiflora in
Western Australia and South Australia. Baskorowati et al.
(2010a, b) observed a wide variety of insects visiting the
flowers of M. alternifolia in New South Wales, including
large flying insects like honey bees (Apis mellifera), flies
and wasps (Figure 6). These authors also found that small
insects like thrips (Thrips imaginis and T. tabacci) were
the dominant visitors to the flowers of M. alternifolia
and are important pollinators, as confirmed by exclusion
Figure 5.Monthly progress in flowering and fruit

development in Melaleuca alternifolia (derived
from Baskorowati 2006)
23
Figure 6.Some of the potential insect pollinators of Melaleuca alternifolia: (A) honey bee (Apis mellifera);
(B) butterfly of family Lycaenidae; (C) butterfly of family Nymphalidae; (D, E) wasps of family Sphecidae; (F) wasp
of family Vespidae; (G) beetle of family Lycidae; and (H) fly of family Calliphoridae (Source: Baskorowati 2006)
24
the apocolpial field was not visible or psilate. Some pollen

of the former genus Callistemon lacked an apocolpial island
or had a small irregular polar island, but all other observed
species possessed closely fitting apocolpial islands. Overall,
pollen length range was 11.029.5 m and the colpus to
length ratio range was 26.456.8%. Two different pollen
types were observed in Melaleuca by these authors: those
with large apocolpial islands, such as in many species of the
M. leucadendra complex and former Callistemon species;
and those with medium-size, syncolpate pollen with faint
granulate patterning. The structure of the Melaleuca pollen grain, with its lack of sculpturing and its propensity for
stickiness and clumping, supports the view that it is mainly
dispersed by insects or flying vertebrates.
There is limited published information about pollen
viability in Melaleuca and its longevity under different
storage conditions. A report on viability of M. cajuputi
subsp. cajuputi pollen using an agar medium showed
that pollen was highly viable (66%) soon after collection,
had 35% viability after 3 months of storage in an airtight
bottle in the refrigerator (35 C), but this had decreased
to 4% viability after 4.5 months (Hendrati et al. 2002).
Baskorowati et al. (2010a, b) found that pollen viability in
M. alternifolia varied significantly with time (tested at 1,
14, 26, 39 and 52 weeks) and temperature (2124 C, 5 C
and 18 C) of storage. Pollen of this species was still viable
after 52 weeks of storage at all temperatures, with storage
at 18 C giving the best results (22%).
Most is known of the sequence of pollen release and
stigma receptivity in M. alternifolia following the work of
Baskorowati et al. (2010a, b, c). Consistent with reports
on other Melaleuca species (e.g. Barlow and Forrester
1984), there is only slight dichogamy in individual flowers in M. alternifolia. The male phase, when pollen is first
shed, occurs 1.5 days post anthesis, while the female phase,
defined as the period of stigmatic exudate formation, occurs
Figure 7. Scanning electron micrograph of Mela

leuca alternifolia pollen (Source: Baskorowati 2006)
34 days post anthesis. The synchrony of flowering within

and between inflorescences on the one tree, the duration of
flowering and abundant pollen provide ample opportunity
for geitonogamy in the species. Despite this, M. alternifolia
displays a breeding system that is preferentially outcrossing: Butcher et al. (1992) reported an outcrossing rate of
93% and Rosseto et al. (1999) 86% for M. alternifolia. Selfpollination, however, can occur, at least when manipulated.
J.C. Doran and G.F. Moran (unpublished report, 2002)
reported a selfing rate of up to 28% among progeny of some
trees when their flowers were bagged without emasculation. A similar result was reported by Kartikawati (2005)
for M. cajuputi subsp. cajuputi in a seed orchard in Yogyakarta, Indonesia, where a few individual trees in the orchard
proved to be self-compatible, although most were found to
be self-incompatible. Baskorowati et al. (2010a, b), reported
that a self-incompatibility system operates in the style and is
complemented by late-acting, self-incompatibility mechanisms discriminating against self-pollen tubes when they
descend to the ovary, based on microscopic observation of
pollen-tube development in M. alternifolia (Figure 8). Barlow and Forrester (1984) also studied self-incompatibility
in various Melaleuca species, although not in M. alternifolia,
and found that self-pollen tubes do not penetrate past the
base of the style.
Hybridisation
Natural hybridisation in Melaleuca appears to be restricted
to within groups of closely related species, although there
has been anecdotal mention of wider crosses occurring
spontaneously in cultivated melaleucas. Hybridisation
occurs very widely across the genus and examples noted
in both the field and the herbarium have been listed by
Craven (2006). In all, over 20 examples are known. It is
expected that, as comprehensive DNA studies are undertaken on species complexes within the genus, more will
become known as to the extent of past and (relatively)
recent hybridisation events.
Natural hybridisation between M. alternifolia and
M. linariifolia has long been suspected in tea tree populations near Port Macquarie, New South Wales. This
suspicion has arisen due to the intermediate leaf morphology, the occurrence of transgressive oil components in
leaves of the Port Macquarie population of M. alternifolia,
similarities in oil composition with M. linariifolia, and
sympatry with M. linariifolia (Butcher 1994). Butcher et
al. (1995) were able to confirm the hybrid status of the Port
Macquarie populations in a study of relationships using
chloroplast DNA.
Fruits
The fruit consists of a three-celled capsule within a usually
woody to subwoody fruiting hypanthium, which is often
cup-shaped but also frequently is described as globular,
25
urceolate, spherical, cylindrical, barrel-shaped, or ovoid;

calyx lobes persist in the fruits of some species and are a
useful aid to identification. Individual capsules are usually small and not greater than 0.5 cm long 0.5 cm in
diameter, although some are larger. Fruits are subsessile
to sessile (e.g. see Figure 5), persistent (partial immersion
in the stem is common in southern Australian species) or
shedding (fruits of some tropical species ripen and shed
in rhythm with the wet season). They are dehiscent and
usually many seeded. Melaleuca alternifolia, for example,
gave from 26 to 57 viable seeds per capsule in a study
of variation in this character between individual trees
(Baskorowati et al. 2010a, c).
Seeds
The seed has a membranaceous or rarely coriaceous testa
containing an embryo but no endosperm. It is unwinged
and smallseeds of 46 largely Western Australian species
had a mean length of 1 mm with a range of 0.52.0 mm
(Sweedman 2006). Seeds are highly variable in shape and
in the sculpturing and colour of the seed surface (see scanning electron micrographs in Sweedman 2006).
Typically in melaleucas, as in eucalypts and several
other genera of the family Myrtaceae, the fine particles
that dehisce from the fruit are a mixture of viable seed
and unfertilised ovules/ovulodes commonly referred to
as chaff . Because of their similarity in shape, colour and
small size in many Melaleuca species, it is almost impossible to separate the two by the naked eye and even when
employing a microscope and other mechanical aids like
winnowing. For this reason, Melaleuca seed is usually handled as this mixture where the percentage of viable seeds
to chaff may be less than 10% (Rayamajhi et al. 2002). Germination rates for most species are given as viable seed per
unit weight of seed and chaff mixture. For example, 27 Melaleuca species that have each received multiple (233 seed
tests per species) four-replicate seed tests over time at
the Commonwealth Scientific and Industrial Research
Organisations (CSIROs) Australian Tree Seed Centre gave
an overall average of 3,700 germinants/g of seed and chaff
mix with a range of 1,6006,000 germinants/g. The highest
germination rate on record at the Centre was for a seedlot
of M. bracteata that gave an average of 17,000 germinants/g
of seed and chaff mix.
Cotyledons
Cotyledons are planoconvex to obvolute and are not or
scarcely foliaceous.
Geographical distribution
and ecology
Natural occurrence and ecology
Figure 8.Fluorescence micrograph of pollen

tube growth (stained with decolourised aniline
blue) in the pistil of Melaleuca alternifolia, 4 days
after cross-pollination (Source: Baskorowati
et al. 2010a)
26
Melaleuca principally is an Australian genus. Eight species,

two of which are endemic (M. pustulata and M. virens),
occur in Tasmania; one endemic species, M. howeana,
occurs on Lord Howe Island in the Tasman Sea; and
there are seven endemic species in New Caledonia
(M. brevisepala, M. brongniartii, M. buseana, M. dawsonii,
M. gnidioides, M. pancheri and M. sphaerodendra). Several
species of the Australian monsoon-tropical, broad-leaved
paperbarks (the M. leucadendra group) also occur in
adjacent areas of Papua New Guinea; i.e. M. dealbata,
M. leucadendra, M. nervosa, M. stenostachya, M. viridiflora, with M. dealbata and M. viridiflora also in Papua
where t is a taxon in the set of taxa T, and Rt is the global

range of taxon t across the dataset (i.e. the number of cells
in which it is found)
The number of Melaleuca species in south-eastern and
eastern Australia is not high, although the phylogenetic
diversity may not be greatly lower than that occurring in
the south-west of the continent. Of particular note are the
bottlebrush species (formerly Callistemon) of which about
30 occur in the south-east and east and only four in the
remainder of Australia. A few species occur in the arid
zone but, in general, Melaleuca has not adapted well to very
dry regions. Within the monsoonal tropics, members of
the broad-leaved paperbark group are often dominant and

may be a characteristic feature of the landscape. Several of
the larger tree species of this group (M. argentea, M. leucadendra, M. nervosa and M. viridiflora) occur across, or
nearly across, the whole northern zone.
Ecological notes are provided for each species in the
Species accounts (Chapter 7) but a brief overview for
the genus is given here. Melaleuca species occur in a wide
range of habitats (Figure 11). Many species are found in
low to tall heathlands and shrublands, and some occur
as low shrubs or trees in open eucalypt forests. A few
species are found in wetlands, sometimes in open water,
and some commonly occur on saline soils. No species are
rainforest plants, although some apparently are able to
survive encroachment by tropical rainforest, notably the
broad-leaved paperbark M. leucadendra. The south-eastern
and eastern M. pallida can occur in wet sclerophyll forest
that may be considered temperate rainforest. Because of
the range of climates, landscape positions and soil types
to which Melaleuca species have become adapted, there is
good scope for selecting species for trial in many applications, e.g. saline land reclamation, mining rehabilitation
and ornamental and amenity use.
province in Indonesia and M. leucadendra extending as

far as the Maluku Islands of Indonesia. Melaleuca acacioides also occurs in southern Papua New Guinea. Two
species of the M. leucadendra group have a much broader
distribution: M. quinquenervia extends to Papua province,
Papua New Guinea and New Caledonia and M. cajuputi
occurs from northern Australia through Malesia to SouthEast Asia. The indigenous western Malesian South-East
Asian populations of the latter species are referable to
M. cajuputi subsp. cumingiana and represent an interesting example of dispersal across Wallacea (Barlow 1988;
Lum 1994).
Within Australia, the majority of the species and the
greatest phylogenetic diversity occur in the south-western
region of Western Australia, especially on the leached
north-western and central to southern sand plains, and in
the clay soils in the drier eastern region of the south-west.
The species distribution maps in the Species accounts
(Chapter 7) show some species are widespread within
the south-west; presumably these are tolerant of variations in soils, landscapes and climates. One such species
is M. concreta which, although restricted to the coastal
plain country north of Perth, can occur on a wide range of
substrates and in different parts of the landscape. Examples
of other widespread species in the south-west are M. carrii
and M. hamulosa. On the other hand, species such as
M. agathosmoides and M. venusta are local endemics and
probably have highly specific requirements as to soil type
and landscape position.
Species richness is shown in Figure 9, and endemism
is shown in Figure 10. The Biodiverse software version
0.17 was used to generate the maps (Laffan et al. 2010)
and the calculations were based on a 100 km grid. The
final maps were then generated using ESRI ArcGIS version 10.0. A continuous gradient from 1 to 72 species is
shown in Figure 9 and this map depicts the lowest and
highest richness. The endemism map (Figure 10) shows the
unweighted endemism score for each grid cell calculated
using equation (1).
1
=
Unweighted endemism
(1)
Rt
tT
Locations of planted forests

Reliable information on plantation areas worldwide by
species is largely unavailable. Of the three main commercial species, M. quinquenervia is arguably the most
widely planted. This species has been planted in numerous
countries, including southern Africa, Central America
(Costa Rica and Honduras), North America (southern
California, Florida, Hawaii, Louisiana and in the extreme
south of Texas), India, Fiji, Madagascar, Mexico, northern Nigeria, the Philippines, Puerto Rico and the West
Indies (Streets 1962; NAS 1983; Turnbull 1986; Geary
1988; von Carlowitz 1991; CABI 2000; Dray et al. 2006).
Two million trees have been planted on State Forest
Reserve alone in Hawaii. Melaleuca quinquenervia was
first introduced into southern Florida in the late 1800s,
where it escaped cultivation on seasonally wet sites and
has assumed weed status (NAS 1983). It occurs now on
more than 200,000 hectares (ha) of wetlands in southern
Florida (Turner et al. 1998).
Melaleuca cajuputi subsp. cajuputi has been planted
since 1926 for oil production in central Java, Indonesia,
using seed originally imported from the Maluku Islands.
The extent of government-owned plantations on Java is in
the order of 20,000 ha (Anto Rimbawanto, pers. comm.
2012). It has been widely planted also in Malaysia and
Vietnam. Melaleuca leucadendra is a relatively underutilised species. Most reports of plantings of this species in
Africa, America and Asia refer to arboretum or trial plantings rather than broad-scale plantations. The substantial
literature on the cultivation of M. leucadendron (e.g. see
27
Species richness
High : 72
Low : 1
0
740
1,480
2,220
2,960
Kilometres
Figure 9.Melaleuca species richness calculated using a continuous gradient from 1 to 72 species
Endemism
High : 9.92035
Low : 0.00518135
0
740
1,480
2,220
2,960
Kilometres
Figure 10.Melaleuca endemism. The greatest endemism occurs in the south-central coastal region
of Western Australia with some moderately strong areas of endemism in the north-western part of
the south-west. There are some lesser areas of endemism in eastern Australia.
28
There are extensive, but largely undocumented, plantings of melaleucas as ornamentals, street and public park
trees, shelterbelts on farms and for land reclamation in
Australia and elsewhere.
Tolerance of difficult
conditions
As well as being tolerant of periodic (Figure 12) or
even continuous waterlogging, many of the wetland
melaleucas will also survive and grow in moderately
to highly saline soils (e.g. M. armillaris, M. bracteata,
M. cuticularis, M. decussata, M. ericifolia, M. lanceolata,
M. lateriflora, M. leucadendra, M. linariifolia, M. quinquenervia, M. squarrosa, M styphelioides and the M. uncinata
Fenton et al. 1977), particularly in South-East Asia and

USA, can be ascribed to M. cajuputi and M. quinquenervia and not M. leucadendra. Most current interest in
the species for plantation establishment is in the Mekong
Delta of Vietnam, where M. leucadendra outperforms
the indigenous M. cajuputi subsp. cumingiana on seasonally inundated and potentially acid-sulfate sites which
are very difficult for tree establishment (Hoang Chuong
et al. 1996).
Melaleuca alternifolia is planted for the production of
essential oil in Australia3,000 ha producing 450500
tonnes (t) of essential oil/yearbut the total area planted
and the total production worldwide is potentially twice
this from plantings outside Australia in several countries,
including China. Broombush (M. uncinata complex)
plantings in Australia for brushwood fencing and related
products exceed 1,000 ha.
Figure 11. Some of the variable habitats occupied by Melaleuca species: (A) a pure stand of tree-form
M. quinquenervia in a coastal swamp in eastern Australia; (B) the shrub M. protrusa on a dry stony hillside in
south-western Western Australia (WA); (C) shrubby M. sophisma under an overstorey of mallet in southwestern WA; and (D) trees of M. cuticularis on the edge of a wet, possibly saline area in south-western WA
29
Figure 12.Melaleuca cajuputi surviving and growing under seasonal inundation in the Mekong Delta of
Vietnam: (A) a young planting as the inundation recedes; and (B) a dense plantation nearing rotation age
complex). A few melaleucas are tolerant of extremely

saline conditions (e.g. M. halmaturorum and M. thyoides).
Many of these same species are also tolerant of alkaline
soils, drought and frost (Marcar et al. 1995; Marcar and
Crawford 2004).
Melaleucas appear to employ a diverse range of
physiological strategies to adapt to difficult growing
conditions. For example, Naidu et al. (2000) found that
species with the capacity to accumulate one or, preferably, more of the methyl prolines in their leaves were
better adapted to saline and/or sodic soils than species
that accumulated only L-proline. This was the case with
M. cuticularis: Carter et al. (2006) found that the ability
of this species to tolerate saline-waterlogged conditions
was related to production of methyl proline, as well as
regulation of foliar sodium, chloride and potassium
concentrations. Ionic stress and the differential ability of
seedlings of M. leucadendra provenances to adjust for a
declining potassium concentration in leaf sap under combined salt and aluminium stress was thought to be the
main cause of variation in growth of 16 M. leucadendra
provenances in a glasshouse trial (Nguyen et al. 2009).
Mensforth and Walker (1996) found that the root dynamics of M. halmaturorum in response to fluctuating saline
groundwater contributed to the survival of this species in
saline swamps. Melaleuca halmaturorum accessed water
from deep in the profile in late summer when salt had
accumulated in the surface soils and used rainfall and
shallower groundwater after winter rains had replenished
the profile. The ability of this species to take up water from
saline substrates through maintenance of low leaf water
potentials was also a contributing factor (Mensforth and
Walker 1996).
30
Some wetland melaleucas, like the tropical M. leucadendra group, develop aerial adventitious roots on their
stems and within the papery bark to the height of the
maximum water level during flooding (Figure 13). These
are dense in aerenchyma cells which have large intracellular air spaces that improve internal root aeration and
gas exchange during inundation. The fine adventitious
roots on the stems of M. quinquenervia in a seasonally
inundated forest in northern Queensland, for example,
were considered an important part of the reason that transpiration in this species was unaffected by inundation of
up to 24 weeks (McJannet 2008). In M. cuticularis, they
appear to contribute to this species enhanced tolerance to
combined salinity and waterlogging (Carter et al. 2006).
Tanaka et al. (2011) reported that seedlings of M. cajuputi
in the tropical peat swamps of southern Thailand were
able not only to survive complete submergence for 8 weeks
but also to photosynthesise and grow during this period.
This was due to the strong development in the leaves and
stems of submerged seedlings of schizogenously formed
aerenchyma which improved uptake of gases from the
water.
Many melaleucas are highly fire-tolerant during all but
the early seedling stages before a thick protective layer
of bark has formed. Fire-ravaged individuals regenerate
through stimulation of epicormic buds under the thick
bark to sprout vigorously after fire in a process called
coppicing (Figure 14). Populations may expand through
fire-induced release of seed from serotinous capsules on
the trees and stimulation of germination of seed in soil
seedbanks.
Some melaleuca species have the ability to root sucker,
and through root extension and interconnectivity form
Figure 13.Aerial adventitious roots on the stems of: (A) Melaleuca quinquenervia growing naturally in a
seasonal swamp beside the Bensbach River, Western Province, Papua New Guinea; and (B) M. leucadendra
planted in the Mekong Delta, Vietnam
dense clumps of single clones. This is a common adaptive

characteristic of wetland plants subject to very difficult
conditions for survival, growth and sexual recruitment
(see review in Robinson et al. 2012). Using DNA markers,
Robinson et al. (2012) showed that the large, dome-shape
stands of M. ericifolia in Dowd Morass, Gippsland Lakes,
Victoria, were individual clones that did not intermingle (phalanx growth habit) with adjacent clones. They
speculate that this is also the case with other stands of
this species in southern Australia. Melaleuca viridiflora
also forms root suckers. Crowley et al. (2009) showed that
population density of this species increased dramatically
in grasslands and grassy woodlands in northern Queensland in the absence of fire by recruitment of suckers to
the sapling layer.
31
Figure 14.Western Australian broombush, Melaleuca concreta, coppicing after fire
32
Uses
Ethnobotanical
Some Melaleuca species were used extensively by the Aboriginal peoples of Australia
for a wide variety of cultural uses (refer to Williams 2011 for a description of 17 species
used by Aboriginal communities).
The papery bark of several, mainly tropical, melaleucas (e.g. M. argentea, M. dealbata,
M. cajuputi, M. leucadendra and M. viridiflora) had many domestic uses, including
water-repellent roofing material, raft-making, in food preparation, bandages, blankets,
baby slings, body wraps in burial ceremonies and for dresses denoting marriage, to
name but a few (Levitt 1981; Wrigley and Fagg 1993; Yunupingu et al. 1995; Blake
et al. 1998; Puruntatameri et al. 2001; Wiynjorrotj et al. 2005; Williams 2011; Wiersema
and Len 2013). The leaves of species such as M. acacioides, M. argentea and M. leuca
dendra were used as flavouring in cooking and M. argentea leaves were burnt to repel
mosquitoes. The trunks of some species (e.g. M. cajuputi, M. leucadendra and M. viridi
flora) were used for construction of canoes and shields.
Bee bread (produced from pollen) and honey were foods

collected from native bee hives prevalent in Melaleuca
swamp forests of M. acacioides, M. lasiandra, M. leucadendra, M. minutifolia, M. nervosa and M. viridiflora in
northern Australia (Williams 2011). Williams (2011) also
notes reports of early explorers (e.g. Leichhardt in 1847
and Mitchell in 1848) in northern Australia of the collection by Aboriginal peoples of melaleuca honey and
melaleuca flowers (e.g. from M. saligna). The latter were
soaked in water to produce a sweet-tasting drink.
Melaleucas were an important source of dry-season
water for the nomadic Aboriginal peoples (and early
European explorers), particularly in the wet/dry tropics
of northern Australia. Bulges in the trunks of individual

trees of such species as M. argentea, M. cajuputi, M. dealbata, M. nervosa and M. viridiflora when undercut could
yield about a litre of brackish but nevertheless potable
water (Yunupingu et al. 1995; Puruntatameri et al. 2001;
Wiynjorrotj et al. 2005; Williams 2011).
Melaleucas played an important role in traditional
Aboriginal medicines. The leaves and inner bark of
M. argentea and others like M. cajuputi and M. leucadendra were used medicinally (coughs and colds, aches and
pains, cuts and sores, ringworm, vomiting and diarrhoea
and other malaises), either directly following crushing or
burning (smoking medicine) and inhaling the odours or as
33
3. Uses
a liniment or drink after soaking the leaves or inner bark

in water and heating (Blake 1968; Levitt 1981; Aboriginal
Communities of the Northern Territory 1993; Wrigley and
Fagg 1993; Yunupingu et al. 1995; Puruntatameri et al.
2001; Wiynjorrotj et al. 2005; Lassak and McCarthy 2011;
Williams 2011). The bark of some melaleucas was used as a
poultice on wounds and for splinting broken bones, where
the juice of the bark was said to penetrate the skin and aid
in healing (Williams 2011). The milky extract of squashed
bee brood (bee pupae and larvae) collected from native
beehives prevalent in melaleuca swamp forests was used
as a topical antiseptic for sores, tinea and eye complaints
(Williams 2011).
Ornamental, landcare,
honey, bark and wood
Ornamental and amenityhorticultural use
Melaleuca species, especially the bottlebrushes, have long
been popular garden subjects in Australia. The first Australian melaleucas to be cultivated, however, were grown in
Europe, presumably from seed taken to England in 1771
by Joseph Banks. Melaleuca armillaris, M. decora, M. ericifolia, M. hypericifolia, M. nodosa, M. styphelioides and
M. thymifolia were in cultivation by 1793 (Elliot and Jones
1993; Wrigley and Fagg 1993). Seed and/or transplants of
the bottlebrush species M. citrina and M. linearis also were
taken to Europe in the late 1700s and these species rapidly
became popular conservatory plants. Melaleuca citrina was
in fact named and described (as Metrosideros citrina) in
1794 from material cultivated in England and M. linearis
was described in 1796 from material cultivated in Germany.
Many species (including cultivars derived from selection or from interspecific hybrids) are hardy in cultivation
in Australia (Elliot and Jones 1982, as Callistemon; Elliot
and Jones 1993; Wrigley and Fagg 1993, also as Callistemon; Holliday 2004; Stewart 2012, also as Callistemon).
Some tolerate moderate levels of frost and others grow well
in poorly drained soils. Because there are species of diverse
habit, flower form and colour, and substrate preference
in most of the major climatic zones within Australia, it
is possible to select a species for a specific purpose, e.g.
for a hedge, windbreak, specimen shrub or tree. Several
species have particularly attractive papery bark and are
worth cultivating for this feature. Several of the larger
shrub and tree species are ideal for amenity plantings as
street trees, screens for industrial sites, highway verges and
so on. Many of the tree species are ideal for use in parks
and large-scale landscape applications. The trees that have
colourful, nectariferous flowers (Figure 15) usually attract
nectar-feeding birds if these occur in the region.
There are surprisingly few melaleuca cultivars available
in the horticultural industry in Australia and most, if not
all, of these are selections of species. The bottlebrushes are
an exception. Barriers to successful hybridisation between
certain species apparently do not exist and, when grown in
a common garden, bird- or insect-mediated hybridisation
has resulted in the occurrence of hybrid plants. In many
cases, selections have been named and propagated commercially. The following species (as Callistemon) have been
recorded as being a parent, or putatively a parent, of named
cultivars: M. citrina, M. comboynensis, M. glauca, M. pachyphylla, M. phoenicea, M. polandii (doubtfully this species
and more likely to have been M. hemisticta or M. pyramidalis), M. recurva, M. salicina, M. subulata and M. viminalis
(Elliot and Jones 1982; Wrigley and Fagg 1993). Much more
Figure 15.Ornamental melaleucas: (A) Melaleuca ryeae, a shrub with profuse pink flowerheads; and (B)
Melaleuca cultivar Harkness
34
series of red- to orange-red-flowered, bushy plants to rival

the bottlebrushes, as the species of the M. fulgens group
also have bottlebrush inflorescences. Naturally occurring
hybridisation in Melaleuca has been reported from a wide
range of species (Craven 2006, p. 470) and it seems barriers
to hybridisation within a species group or species complex
are not absolute; in fact, where two or more species of the
same group occur in biotic sympatry, hybridisation may
be frequent.
An interesting application is the use of melaleucas as
bonsai (known as penjing in China and cay canh in Vietnam). Bark texture, small leaves and flowering propensity
are some of the criteria by which species are rated. Over
60 species, varieties and cultivars of Melaleuca have been
recorded by the Australian Plants as Bonsai Study Group
as being grown in bonsai within Australia (Roger Hnatiuk,
pers. comm.). Early use of melaleucas tended to mimic the
styling of classic Japanese trees but in recent years bonsai
artists are exploring the beauty of branches and crowns of
old mature trees of the species found growing in nature.
Bonsai, inspired by such forms, are beginning to be seen
in public displays (Figure 16).
Notes on species of particular, or potential, value for
ornamental and amenityhorticultural use are provided in
the Species accounts (Chapter 7). The main impediment
to their wider use within Australia and elsewhere is the
difficulty in obtaining planting material of superior colour
and growth forms of the species, or forms from particular
soil types.
3. Uses
work needs to be done as far as bottlebrush breeding is

concerned. There is considerable opportunity for breeders
to select growth and colour forms of species and cross them
with an aim of obtaining a particular combination of characters. The two broad climatic zones that should be targeted
for such hybridising endeavours are the tropicalsubtropical and subtemperate zones in Australia and elsewhere.
As to other species groups within the genus, it seems
there have been no hybridisation programs. The field is
open for such programs to be initiated. The pom-pomflowered M. scabra group is an obvious candidate as there
is much variation within this group that could be bred into
novel, improved garden plants. Although many of the species of the M. scabra group occur on leached, sandy soils,
some of them occur on clay soils and these species may
confer some degree of soil adaptability to hybrids. Melaleuca nesophila, an M. scabra group species of restricted
range and habitat in the high rainfall, coastal part of southwestern Western Australia, is remarkably hardy and grows
well on heavier soils in the dry Western Slopes region of
New South Wales. Using species such as M. carrii, M. fabri,
M. hamata, M. nematophylla, M. nesophila, M. oldfieldii,
M. sapientes and M. systenaand there are many othersit may be possible to develop hybrids of merit for
Mediterranean and dry-temperate climates.
Another group with potential as garden plants is the
M. fulgens group, and a hybridisation program involving such species as M. coccinea, M. elliptica, M. eximia,
M. fulgens, M. lateritia and M. macronychia may result in a
Figure 16.Examples of bonsai melaleucas: (A) Melaleuca lateritia, usually a 23 m tall shrub from southwestern Western Australia; and (B) M. bracteata, usually a medium-size tree up to 22 m tall from inland and
coastal northern Australia
35
3. Uses
Land rehabilitation
Melaleuca comprises many species of trees and shrubs that
are hardy and adaptable to a wide range of habitats and
soils. They regularly occupy sites that are very challenging for tree survival and growth (Figure 17), as discussed
in other sections of this volume. Their diversity in form,
adaptability and utility sees them listed prominently
among candidate species for planting for land reclamation, with natural resource benefits including mitigation
of salinity, waterlogging, and water and wind erosion.

Biodiversity improvements, carbon sequestration and
potential to increase farm income (e.g. through production of brushwood fencing, essential oils and bioenergy)
are among other commonly stated benefits of planting
melaleucas on degraded lands.
Highly topical at present is the increasing problem of
saline landscapes in Australia and elsewhere. There are
over 4 million ha of secondary or human-induced saline
soils in Australia, in addition to 29 million ha of naturally
Figure 17.Salt-tolerant melaleucas: (A) Melaleuca halmaturorum on a saline site

in southern Australia; and (B) M. atroviridis surviving on the margin of an area
severely affected by secondary salinity in south-western Western Australia
36
Brushwood fencing and related products

Ornamental brushwood fencing comprising the grey
stems, twigs and dry foliage of the M. uncinata complex
of species (broombushes), hand-packed on wires in situ
or, in more recent times, prefabricated in panels, has been
in use in Australia for more than 80 years (McKelvie et al.
1994) (Figure 18). It represents an important market for
melaleucas, despite its small size compared with alternative fencing materials (e.g. 1% of the fencing market in
Western Australia). Other uses of brushwood include
manufacture of garden furniture, gazebos, pergolas, gates,
hanging baskets and decorative bird feeders (Robinson
and Emmott, no date). McKelvie et al. (1994) indicated
that about 600,000 bundles of brushwood (each c. 25 kg,
consisting of stems with foliage of 1.41.8 m in length and
715 mm in diameter) were used in Australia each year,
with predicted annual market growth of 5.5%.
Melaleuca uncinata is the most common broombush
used for brushwood fencing. It is widespread in southern
Australia, mainly south of the Tropic of Capricorn. Melaleuca uncinata is a hardy, bushy shrub to 7 m in height with
multiple long, thin, woody erect stems topped with foliage
(broom-like). It is adapted to a wide diversity of habitats
and soils, mainly in semi-arid and arid Australia (see range
given in its species description). Other broombushes in the
M. uncinata complex having potential for brushwood fencing are the Western Australian endemics M. atroviridis,
M. concreta, M. hamata and M. osullivanii (Robinson and
Emmott, no date). Melaleuca acuminata and M. hamulosa

are also worth trialling as alternative species as they have
similar physical characteristics to the traditionally used
M. uncinata (Peter White, pers. comm. 2012).
Until recent times, all harvesting of brushwood has
been in native stands and producers are licensed by the
various state governments. Overall, 5070% of plants are
harvested manually of which 90% can be expected to coppice in a typical bush operation based on a broombush
population of about 3,000 plants/ha. This yields between
36 t product/ ha. The interval between harvests is 1014
years (Wrigley and Fagg 1993). Concerns about adverse
environmental effects have seen the gradual withdrawal
of public lands from resources available for harvesting and
concerns have arisen about the sustainability of brushwood
supply from native stands. The likely direct economic benefits, complemented by the indirect benefits of establishing
broombush on private lands for purposes such as shelter,
salinity control, soil erosion control, biodiversity and
diversification of farm income, have led to interest among
some landholders in the drier regions of the southern
states in developing commercial plantations of broombush.
It is estimated that a total plantation area in the order of
2,0002,500 ha would be required to fully meet demand on
a sustainable basis (Cameron 2003; AVONGRO 2007). The
total area planted is not known but about 900 ha had been
planted in the Avon catchment area of Western Australia
alone by 2007 (AVONGRO 2007).
Various landcare agencies in Western Australia, South
Australia and Victoria have published informative fact
sheets on growing broombush by either direct seeding or
planting tube stock and readers interested in this topic are
directed to these for detailed information (e.g. Bulman et
al. 1998; Cameron 2003; Robinson and Emmott, no date).
Until recently, broombush was regarded as being a single
species, M. uncinata. It is now known that there are several
species involved (Craven et al. 2004), a number of which
are suitable for brushwood production as mentioned above.
3. Uses
occurring salt-affected lands (Marcar and Crawford 2004).

The removal of native vegetation and development of annual
agricultural systems in southern Australia, leading to rising watertables carrying soil-borne salt to the surface, are
major contributors to salinisation of landscapes. Restoration
of deep-rooted perennial vegetation can make a significant
contribution to correcting this problem but it needs to be
on a large scale to control salinity (Pannell and Ewing 2004).
A range of Melaleuca species is suitably adapted to grow
on saline sites (e.g. Figure 17), ranging from moderately
(48 dS/m ECe) to extremely saline (>16 dS/m ECe) (Marcar et al. 1995; Department of Agriculture and Food Western
Australia 2004; Marcar and Crawford 2004).
Melaleuca halmaturorum is an example of a temperate
Melaleuca tree/shrub adapted to extremely saline conditions, while M. leucadendra is among the tropical tree-form
melaleucas adapted to highly saline conditions. Both species are tolerant of waterlogging. Unfortunately, few of the
presently recognised salt-tolerant melaleucas offer scope
to growers for direct financial benefits, thus large-scale
planting is not attractive. Melaleuca uncinata, one of the
source species for brushwood (see below), and its relative
M. atroviridis, are two exceptions. Melaleuca bracteata
might have potential in the future if an industry were to
develop around the production of betaines from its foliage.
Honey
Most melaleucas do not provide major honey crops. Clemson (1985), however, pointed out that many species assist
indirectly with honey production by providing nectar and
pollen, especially nectar, in sufficient quantities to stimulate brood-rearing and sometimes for use as stores. In this
way, colonies are maintained and built up for subsequent
major honey flows in other taxa.
Those which are important honey producers include
the broad-leaved melaleucas. Melaleuca quinquenervia is
a major source of honey in Australia and Florida (Blake
and Roff 1972; Robinson 1981; Clemson 1985; Geary 1988)
and similarly M. cajuputi in northern Australia and Vietnam (Brock 1988; Mulder 1992). Melaleuca leucadendra
is also said to be an important source of honey in its area
37
3. Uses
of natural occurrence (Roff 1966; Anderson 1993). The

honeys from these species are variously described as light
to dark amber in colour, with strong flavour and odour
and of low density so they granulate readily. Their pollens
are universally described as being a good source of protein
utilised by bees in building up colonies.
the cork from the bark has been used in infants pillows
and mattresses (Bootle 1983). The bark of M. cajuputi is
used in parts of Malaysia as a luting material in boatbuilding (Lum 1994; Lim and Midon 2001).
Bark
Fuelwood
The bark of Melaleuca species is still used today in the construction of traditional houses in Papua New Guinea. It is
used to line fernery baskets, for making bark paintings and
There is little reported on the fuelwood value of individual

species in the genus Melaleuca. Apart from the statement in Maiden (1889) that M. linariifolia wood made
Wood
Figure 18.Facets of the brushwood fencing industry: (A) the multistemmed habit of species in the broombush
complex (this is Melaleuca stereophloia); (B) a native broombush (M. uncinata) population; (C) broombush
bundles in the field ready for transport; (D) transporting the bundles to market; (E) constructing a brushwood
fence in situ; and (F) a typical brushwood fence
38
Posts, poles, stakes and sticks

The stems of the larger melaleucas like M. cajuputi, M. leucadendra, M. quinquenervia and many other species were
regularly used in the round or roughly fashioned for use
as posts, poles, piles, mine timbers and general construction materials (e.g. rafters for huts, fencing rails) in the
early days of settlement in Australia (Maiden 1889). Posts
were said to be durable in contact with fresh or salt water
(e.g. M. cajuputi, Blake 1968), although Cherrier (1981)
reported that durability of untreated M. quinquenervia
posts in the ground was high for 1 year but declined
thereafter, and replacement was necessary after about 3
years. In present-day Vietnam, there is widespread use
of the roundwood of the indigenous M. cajuputi and the
introduced M. leucadendra for piles, poles and general
construction materials (Figure 19).
A specialised industry exists in Western Australia to

supply sticks for use in the lobster-fishing and vegetablegrowing industries (Peter White, pers. comm. 2012).
Although other myrtaceous species may be used, such as
Kunzea sp., sticks derived from natural stands of M. viminea
are preferred due to their greater durability. It is believed that
lobsters enter pots made from natural materials more readily
than they do pots made from synthetic substances. In recent
years, about 1.2 million M. viminea sticks/year have been
used in making lobster pots. Additionally, large numbers of
M. viminea stakes are used each year in the vegetable-growing areas of the Gascoyne region for supporting climbing
beans. Presently, these markets are supplied from natural
stands but it may be that, in the future, the supply of sticks
and stakes could be augmented from plantation sources.
3. Uses
a first-class fuel, most published reports of the fuelwood

value of melaleucas refer to the larger-growing species
in the tropical broad-leaved M. leucadendra group. The
wood of M. quinquenervia, for example, is reported to be
an excellent fuel and converts into good-quality charcoal.
The reported calorific values for the wood and bark of this
species are around 18,400 and 25,800 kJ/kg (4,400 and
6,160 kcal/kg), respectively (Wang et al. 1981), but there
is great variability in these values between trees (Wang
and Littell 1983). The uniquely high heat of combustion
of M. quinquenervia bark (equivalent to some coals) is
due to the presence of a great amount of fatty substances
in the bark (Wang et al. 1982). Keating and Bolza (1982)
rated M. cajuputi and M. leucadendra as good fuelwoods,
although often difficult to split because of interlocking
grain. Gough et al. (1989) reported that the light wood of
M. leucadendra was quick to ignite, with sooty acrid smoke
initially produced from the burning bark.
Sawn wood
The wood of the broad-leaved paperbarks M. cajuputi,
M. leucadendra, M. quinquenervia and M. viridiflora has
yellowish sapwood, merging gradually into pinkish-brown/
red/grey heartwood. It has a high silica content (0.21.0%)
which blunts saws and planes. It is hard, heavy and of
moderate strength, with wood from native trees giving a
green density of c. 1,070 kg/m3 and an air-dry density of
c. 750800 kg/m3 (Keating and Bolza 1982; Bootle 1983).
Florida-grown wood of M. quinquenervia has a basic
specific gravity of 0.49, a density of 1,070 kg/m3 (green),
640 kg/m3 (air-dry) and 620 kg/m3 (oven-dry) (Huffman
1981). Collapse is slight, with shrinkage about 3.5% radial
and 7% tangential (Bootle 1983). Sawn timber tends to
check and warp but, if carefully seasoned, it is suitable
for general construction and flooring. Boards are difficult
to plane and mortice due to interlocking grain but glue
well and are good for joinery. Boat knees can be cut from
branches using their natural shape.
Figure 19.Production and marketing Melaleuca poles and piles in Long An province, Vietnam: (A) harvesting
and loading poles onto a barge for transport to market; and (B) stacks of melaleuca poles at a roadside market
39
3. Uses
Woodchips
Vietnam is the main producer of Melaleuca woodchips for
use in fibreboard production. In 2010, 100,000 t (bonedry) of Melaleuca woodchips were exported from Vietnam
to China, presumably for this purpose. Presently, there are
well-advanced plans to establish a medium-density fibreboard (MDF) plant in the Mekong Delta of Vietnam to
utilise the Melaleuca resource directly (Stephen Midgley,
pers. comm. 2012).
The kraft pulping potentials of 2-year-old paperbark
wood from Vietnamese plantations were reported by Chen
and Su (1998, as M. leucadendron). The low pulp yield and
high chemical consumption were unfavourable pulping
characteristics but the strength index was adequate and
bleachability excellent. The authors indicated that older
trees might have better pulp qualities.
Extractives
Organic chemicals produced and stored naturally in
plant tissues are numerous and chemically complex. By
definition, extractives are the organic chemicals that can
be removed from plant tissues by the action of water,
including steam, other inert solvents such as alcohol and
by mechanically crushing the source materials. The types
of extractive from selected Melaleuca species that are of
economic importance or have commercial potential fall
into two classes: non-volatile (e.g. betaines) extractives and
volatile (foliar essential oils).
Non-volatile extractives
The foliage of a range of Melaleuca species produces commercial levels (>2% fresh weight) of betaine (Naidu and
Cameron 1999). Betaines are non-volatile, water-soluble
compounds and comprise three methylated prolines:
N-methylproline, trans-4-hydroxy-N-methylproline and
trans-4-hydroxy-N,N-dimethylproline. They are osmoprotectants against stress (e.g. unfavourable temperatures,
drought, soil salinity) in tolerant plants and on application
(foliar and seed treatment) to stress-susceptible plants can
create acquired tolerance. Naidu (2003) believes that the
use of betaines to increase stress tolerance in Australian
agricultural crops would stabilise and even increase the
national income from agriculture.
Glycine betaine, a by-product of the sugar-beet industry, is currently sourced from Finland and a worldwide
shortage is predicted for this solute. Australian melaleucas
are a good alternative source of osmoprotectants. Naidu
(2003) found that M. bracteata, which accumulates a proline analogue, trans-4-hydroxy-N-methyl proline, had the
greatest potential of the melaleucas tested for commercial
development, because of its adaptability, vigorous growth
40
and high yields. Despite this potential, there have been no

recent reports of further development of this opportunity.
Research has also shown Melaleuca bark and leaves to
be a rich source of phenolic extractives (tannins) (Huffman
1981; Hussein et al. 2007) but no commercial exploitation
of phenolics from melaleucas for uses such as wood adhesives, leather tanning and as antimicrobial agents has been
reported.
Novel flavonoids have been identified in the leaf waxes,
seeds and honey of several Melaleuca species (e.g. Courtney
et al. 1983; Wollenweber et al. 2000; El-Toumy et al. 2001;
Yao et al. 2004; Yoshimura et al. 2008). Similarly, various triterpenes, some previously undescribed, have been extracted
from the leaves, bark, wood and seed of various melaleucas
(e.g. Ahmad et al. 1997; Lee and Chang 1999; Vieira et al.
2004; Bar et al. 2008). Habila et al. (2010), for example,
have reported the extraction of a triterpene, betulinic acid,
from the wood of M. bracteata. They tested this compound,
which they state is known for its anti-HIV and cytotoxic
activity against malignant versus non-malignant cancer cell
lines, and against a number of pathogenic organisms from
the genera Trichophyton, Candida and Microsporum. They
found that it had great potential as an antifungal drug.
Foliar essential oils

An essential oil is the (usually) hydrophobic liquid containing the volatile compounds that are found in the oil glands
or trichomes of a plant. These glands are usually associated with the leaves, although bark, wood or roots of plants
may also contain essential oil. The oil is usually obtained
by steam distillation, although the expressed oil (as in the
case of citrus peel) can also be used. Essential oils are usually associated with species in the families Myrtaceae and
Rutaceae, although they do occur in some other families.
Commercially important oils

Relatively few Melaleuca species have essential oils of commercial interest. One of the first species to be exploited
commercially for its foliar oil was M. cajuputi subsp. cajuputi in the Maluku archipelago of Indonesia, probably in
the first part of the eighteenth century. It was one of the
first products imported to Europe from South-East Asia
by the Dutch (Gildemeister and Hoffman 1961, cited in
Lowry 1973), because of its reputation as a panacea in the
treatment of all kinds of diseases. Cajuput oil is produced
currently in South-East Asian countries including Indonesia, Cambodia and Vietnam and annual production
potentially exceeds 600 t (Doran 1999a, b). This oil is rich in
1,8-cineole (typically 4060% of total oil) (Doran 1999a, b;
Pujiarti et al. 2011), as is medicinal Eucalyptus oil but at
slightly higher proportions (70% or more). Cajuput oil
acts as a mild antiseptic and is especially useful for treating respiratory ailments, but also finds use in a wide
range of personal-care (e.g. ointments and liniments) and
animal-husbandry purposes (Figure 20). Annual production in Australia of Australian tea tree oil is estimated to be
in the order of 400500 t, worth approximately A$1520
million at the farm gate.
The three abovementioned species/chemotypes provide
the bulk of the commercial production of essential oils
from the genus at present. In addition, there is sporadic
interest in the following oils.
Linalool-rich oil is sourced from specific provenances
of M. ericifolia. Linalool, with its fruity notes, is of value to
the flavour and fragrance industries and, although it can be
produced synthetically, there remains a market in aromatherapy where natural linalool is preferred (Coppen 1995).
E-nerolidol-rich oil can be extracted from the appropriate chemotype of M. quinquenervia. This compound,
presently sourced from a diminishing world supply of
cabreuva oil (Erich Lassak, pers. comm. 2007), has an
established market in perfumery where it is used as a base
note in many delicate, flowery odour complexes (Bauer
et al. 1997). Melaleuca quinquenervia was shown to yield
and coppice well in plantations in northern Queensland
(Doran et al. 2007) before the recent arrival of myrtle rust
to Australia (see Chapter 5).
E-methyl cinnamate has been derived from a northern
Queensland form of M. viridiflora (Hellyer and Lassak
1968; Brophy and Doran 1996). Methyl cinnamate is a
colourless, crystalline solid with a fruity, sweet-balsamic
3. Uses
household products (Doran 1999a, b; Lassak and McCarthy 2011). Niaouli oil from the 1,8-cineole-rich form of
M. quinquenervia (4080% 1,8-cineole) was, until recently,
produced in Madagascar from plantations yielding 1.52.0 t
of oil/year. This oil was used for similar purposes to cajuput
oil (Ramanoelina et al. 2008; Lassak and McCarthy 2011).
Production of niaouli oil in New Caledonia from natural
stands of M. quinquenervia (Trilles et al. 1999, 2006) has
now also ceased after many years of exploitation, although
there is new interest in producing this oil type in Vietnam
(Le Dinh Kha, pers. comm. 2012).
The basis for the commercial interest in, and development of, the Australian tea tree oil industryutilising
mainly plantations of M. alternifolia established in northern New South Wales and northern Queensland and
plantings made outside Australiacan be traced back
to the 1920s. It was then that the medicinal properties
of the oil were first studied and reported (Penfold and
Grant 1925). The terpinen-4-ol-rich oil produced from
M. alternifolia was found to be a powerful antimicrobial agent. It has demonstrated its ability to serve as an
antiseptic, antifungal, antiviral, antibacterial and antiinflammatory agent in multiple studies and is relatively safe
for topical applications (Southwell and Lowe 1999; RIRDC
2007; Lassak and McCarthy 2011). It is incorporated into
many personal-care and household products and is seeing
increasing use in products designed for agricultural and
Figure 20.A sample of the many products that utilise Australian tea tree oil
41
3. Uses
odour and its uses include as a flavour enhancer and in

perfumery (Bauer et al. 1997). A small market exists for
the natural product, but it can be produced artificially at
relatively little cost and there is strong competition from
other natural sources.
Platyphyllol, a b-triketone, is found in M. cajuputi. It
has been identified as having ultraviolet-blocking attributes
and insecticidal properties but it is not being produced
commercially at this time (Yaacob et al. 1989; Brophy and
Doran 1996; Doran 1999a, b).
Citral-rich oil is extracted from M. teretifolia (Southwell
et al. 2003, 2005). A small production of this oil has commenced from natural stands and plantations in Western
Australia. The producer claims the oil has perfumery and
therapeutic properties.
An objective of work undertaken for this book was, if
possible, to identify other species/chemotypes of melaleuca
with commercial potential. The results of this work are
summarised in the Species accounts (Chapter 7) and in
Appendix 1, which provides a quick reference to the oil
type(s) present in individual species.
Inter- and intra-specific variation

Essential oils from a plant species are not necessarily
chemically uniform. There can be, and usually is, variation
in the compounds contained in the oil and their relative
percentages. For this reason, chemists studying the essential oils of plants try to examine samples of the oils from
a number of plants, preferably from many different sites.
Variation in the essential oils may be continuous over a
geographical region, or it can be quite discrete. This latter
occurrence leads to the finding of different chemotypes of
the plant, i.e. plants which, though morphologically the
same, produce different essential oils. It is possible, because
of inadequate sampling, to find two different oil compositions within a species (i.e. two chemotypes) when, in fact,
they just represent the extremes of a continuous variation.
Examples of both types of variation as they apply to the
main (M. quinquenervia, M. alternifolia and M. cajuputi)
and one minor (M. ericifolia) commercial oil-producing
species are discussed below. The possible presence within
a species of these types of variation should be borne in
mind when interpreting the results given in the following sections.
The essential oil of M. ericifolia has been studied for over
50 years. A 2004 study examined its essential oil content
over its natural geographical range, which extends from
coastal regions near Newcastle, New South Wales, in the
north to Tasmania in the south (Brophy and Doran 2004).
The study looked particularly at the amount of 1,8-cineole and linalool in the oils and the results are shown in
Figure 21. Generally, the amount of 1,8-cineole increases
from Newcastle southwards to Tasmania, and the amount
of linalool decreases concomitantly. As the essential oil of
42
M. ericifolia is important because of its linalool content, it

is important to source the oil from plants in the north of its
range. But there is a continuous variation in oil contents so,
in this case, it is not correct to refer to a cineole chemotype
or a linalool chemotype.
Melaleuca quinquenervia, in contrast, is a species known
to contain chemotypes. There are two distinct chemotypes
of this species; chemotype I contains E-nerolidol as the
major component (in amounts of up to 95%) of the oil,
while chemotype II contains major amounts of either
1,8-cineole or viridiflorol (or both), as well as lesser
amounts of other terpenes. There is no plant producing
an oil containing significant amounts of all three oils
(E-nerolidol, 1,8-cineole and viridiflorol). Viridiflorol
production is catalysed by terpene synthase enzymes. The
genes coding for these enzymes are present in the genome
of all plants of the species, but are expressed only in the viridiflorol chemotype (Padovan et al. 2010). The Australian
distribution of the two chemotypes has been mapped and
is shown in Figure 22 (Ireland et al. 2002). The chemotype
containing major amounts of 1,8-cineole is the basis of the
niaouli oil industry.
Melaleuca alternifolia exists in several chemical forms
(chemotypes), although among them there are only three
principal forms (Butcher et al. 1994; Homer et al. 2000).
These three chemotypes contain terpinen-4-ol (up to 50%),
the chemotype on which the tea tree oil industry is based,
1,8-cineole (up to 60%) and terpinolene (up to 50%). Production of the oils of these three chemotypes is controlled
by three different enzymes (Keszei et al. 2010a, b). The
1,8-cineole chemotype is not used commercially, and the
oil presents a similar oil profile to a eucalyptus or niaouli
oil. This particular type of oil is very common in species of
Melaleuca (see Appendix 1). The terpinolene chemotype
found in M. alternifolia (Southwell et al. 1992) also occurs
in M. trichostachya (Brophy 1999).
Melaleuca cajuputi subsp. cajuputi, the basis of the
cajuput oil industry in South-East Asia, contains up to
approximately 60% of 1,8-cineole, together with lesser
amounts (approximately 10%) of limonene, a-terpineol
and viridiflorene, and spathulenol (up to 30% in some
cases). There are, however, a few isolated cases of this species, from northern Western Australia, producing an oil
containing large amounts of E-nerolidol and virtually no
1,8-cineole (J.J. Brophy et al., unpublished data).
Species by oil type

With 290 species in the genus Melaleuca, it is not surprising that their leaf oils have much variation, resulting
in many different types of oils. In this short section, we
will review this variation and show what a wide range of
chemicals is contained in Melaleuca leaf oils. The full data
from the analyses are provided on the internet, accessible
at <http://aciar.gov.au/publication/MN156>.
So far, only five species (M. bracteata, M. halmaturorum,

M. leucadendra, M. squamophloia and M. viridiflora) produce chemotypes whose oils contain a preponderance of
aromatic chemicals. Melaleuca bracteata gives four chemotypes in which methyl eugenol, E-methyl isoeugenol,
elemicin and E-isoelemicin are the principal components
of the oils. In a large collection survey in southern and
central Queensland, several collections (from Rolleston)

did not contain any aromatic components but were composed entirely of terpenoid components (Masunga 1998).
Melaleuca squamophloia, a species with a limited distribution, also produces oils containing either elemicin or
E-isoelemicin as the principal component (Brophy et al.
1999). Melaleuca leucadendra from the eastern part of its
range produces oil in which the principal components are
either methyl eugenol or E-methyl isoeugenol. The methyl
3. Uses
Species producing aromatic (in the chemical sense) oils
50
1,8-cineole content (% of total oils)
45
40
35
30
25
20
15
10
5
0
30
31
32
33
34
35
36
37
38
39
40
41
42
37
38
39
40
41
42
Latitude (S)
70
Linalool content (% of total oils)
60
50
40
30
20
10
0
30
31
32
33
34
35
36
Latitude (S)
Figure 21.Variation in the proportions of (A) 1,8-cineole and (B) linalool (% of total oils) in the essential oil of
Melaleuca ericifolia with latitude of occurrence (derived from Brophy and Doran 2004)
43
3. Uses
oil (Brophy et al. 2006a). Melaleuca nanophylla contains

a b-triketone, flavesone (44%), as a principal component
of its leaf oil, while M. deanei contains a homologous
b-triketone, leptospermone, in small amounts.
Species producing lemon-scented oils

There are only four Melaleuca species known to produce
lemon-scented oils, namely M. alsophila, M. citrolens,
M. stipitata and M. teretifolia. Melaleuca alsophila exists in
several chemical forms, but one form contains geranial as a
major component. In this oil there is a significant amount
of terpinen-4-ol.
Melaleuca citrolens, so named after the lemon scent
of the crushed leaves, exists in six chemical forms, three
of which have this lemon scent. These lemon-scented
forms contain (a) citronellal, as well as 1,8-cineole and
isopulegol, (b) 1,8-cineole, neral, geranial and citronellic
acid in significant amounts, and (c) neral, geranial and
methyl cinnamate. There is also a form that contains citronellol (2147%) and methyl citronellate (931%) which
has a pleasant fruity (but not necessarily lemon-scented)
odour. Melaleuca stipitata also contains neral and geranial
(totalling over 40%) as well as terpinen-4-ol (10%) in its
lemon-scented leaf oil, while one chemotype of M. teretifolia contains neral (29%) and geranial (39%) as principal
components of its oil.
Figure 22.Chemotypes of Melaleuca quinquen
ervia in Australia (Source: Ireland et al. 2002)
eugenol chemotype bred true, while the methyl isoeugenol

chemotype produced these two compounds in an approximate ratio of 3:1 (Brophy and Lassak 1988). The mix of
aromatic chemotypes occurs from Flying Fox Creek in the
Northern Territory eastward to Queensland. Westward of
Flying Fox Creek the oil is entirely terpenoid in content
with no aromatic compounds present (Brophy 1999).
Melaleuca halmaturorum produces an oil which,
although mainly terpenoid, contains about 30% of
2,4,6-trimethoxy-1-isobutyrophenone and small amounts
of other aromatic ketones (J.J. Brophy et al., unpublished
data). Melaleuca viridiflora has two chemotypes, one of
which has three terpene variants, while the other chemotype contains E-methyl cinnamate (82%) as its principal
component. The remainder of this oil is composed of
2,4,6-trimethoxy-1-isobutyrophenone (5%), a monoterpene, E-b-ocimene (12%), and other terpenoid compounds
(Hellyer and Lassak 1968; Brophy 1999).
Related to aromatic compounds are b-di-or b-tri
ketones, and several species produce these in significant
amounts in their leaf oils. Melaleuca triumphalis contains a
novel b-diketone, triumphalone, and its thermal rearrangement product, as by far the major component of its leaf
44
Species producing oils with significant amounts of

linalool
So far, about 10 species have been found to produce leaf
oils containing significant amounts of linalool. They can
contain up to 55% of linalool in their oils, sometimes
within a particular chemotype of that species. The main
species is M. ericifolia which, in the northern extent of is
range, produces an oil containing up to 55% of linalool
and, more importantly, in yields of 12%, based on fresh
leaf. In the other part of its range, its oil contains 1,8-cineole as the major component (Brophy and Doran 2004,
and references therein).
Other species containing linalool in significant
amounts are: M. bisulcata, 55% (0.5% yield); M. depressa,
37% (0.3% yield); M. exuvia, 1626% (2% yield);
M. hamata, 2834% (0.61.0% yield); M. parviceps, 41%
(0.2% yield); M. spicigera, 39% (0.1% yield); M. systena,
30% (0.2% yield); and M. tuberculata subsp. tuberculata,
57% (0.4% yield)all yields are based on fresh weight of
leaf. For any of these species to be useful commercially,
they would have to produce the oil at reasonable concentrations in the leaves (say >1%, weight for weight [w/w]
fresh weight), have a high percentage of linalool (say 50%
and above) in their oils and produce high yields of leafy
biomass from which to extract the oil. These constraints
rule out most of the above species except M. ericifolia. It
is probable that, in most of them, examination of more
Species producing oils with significant amounts of

terpinen-4-ol
Melaleuca alternifolia, and to a minor extent M. linariifolia
and M. dissitiflora, are used commercially for the production of Australian tea tree oil because of their high
percentage of terpinen-4-ol and good oil yield. Several
other species also produce terpinen-4-ol oils: M. alsophila,
1528% (0.10.3% yield); M. arcana, 2331% (0.61.0%
yield); M. calcicola, 33% (0.3% yield); M. concreta, 35%
(12% yield); M. exuvia, 2228% (2.02.3% yield);
M. foliolosa, 2340% (<0.1% yield); M. halophila, 44%
(1.7% yield); M. hamata, 2442% (0.72.0% yield);
M. nodosa, 1820% (0.81.3% yield); M. ochroma, 44%
(0.6% yield); and M. uncinata, 2731% (0.30.5% yield)
all yields being based on fresh weight of leaf.
As with the species rich in linalool mentioned above,
there would have to be a significant reason to consider
using these species to produce a tea tree oil. A reason could
be that the species grows in saline soils, e.g. M. hamata,
or contains other useful components, e.g. M. alsophila
(1219% geranial) and M. exuvia (1626% linalool).
Groundbreaking work on gene control of terpene

biosynthesis in melaleucas
Over the past decade, there has been a revolution in understanding the genes that control the production of essential
oils. Researchers at the Australian National University
(ANU) have been using new technologies developed
originally in the family Lamiaceae, and especially in mint,
to study the genes controlling both the profile and quantity
of essential oils in M. alternifolia. The primary objective
of this work is to improve efficiencies in the breeding of
M. alternifolia for better oil quality and greater in-leaf
oil concentrations (Keszei et al. 2010b). The latter has a
direct influence on off-paddock yields and the economics
of production in this commercially important essential

oilproducing species (Zhang et al. 2011).
Several conclusions can be drawn from the ANU work
on the genes controlling terpene synthesis in M. alternifolia that might be broadly applicable throughout the genus.
Six chemotypes described previously (e.g. Butcher et al.
1994; Homer et al. 2000), and one additional intermediate
chemotype, were identified from a reanalysis of existing
chemical data on the composition of M. alternifolia leaf
oils. It was confirmed that, as suggested by the chemotypes
present, as few as three terpene synthase genes produce
most of the monoterpenoid compounds in M. alternifolia
oil.
Keszei et al. (2010b) found that the gene that makes the
commercially important compound, terpinen-4-ol, likely
arose from a chance gene duplication event to an existing
gene that made 1,8-cineole, followed by a small number of
mutations. Thus, oil quality (high proportions of terpinen4-ol accompanied by low proportions of 1,8-cineole) in
this species, as demanded by industry, is based on a very
small number of genetic variants. These genes have been
characterised, thus allowing screening of seedlings at an
early age to indicate the chemotype of the mature plant.
The oil profiles of other species of Melaleuca (e.g. M. quinquenervia) are produced similarly (Padovan et al. 2010).
In contrast, the yield of oil in M. alternifolia is determined by the flux of precursor metabolites that are made
available for the terpene synthase enzymes. These precursors are produced by a complex series of enzymes in the
plant cell and, in high-yielding plants, nearly all of these
are up-regulated (Webb et al. 2013). Ongoing work aims to
identify the genetic variants that are associated significantly
with foliar oil concentration, raising the exciting possibility
of being able to efficiently screen plants at a young age for
their oil-producing capacity (Klheim et al. 2011).
Readers are directed to texts such as Sell (2010) for
general information on basic biosynthetic pathways for
terpenoid compounds in plants and to Southwell and Lowe
(1999) for information specific to Melaleuca oil biogenesis.
3. Uses
samples would show a range of linalool concentrations

and the existence of chemotypes within the species, so
some might be potentially useful.
45
Propagation, silviculture
and management
Propagation
There are many texts available on the propagation of Australian Myrtaceae, including
Melaleuca species, and readers embarking on a major propagating and planting of
melaleucas are directed to these for detailed information. Available texts include Doran
(1990, 1997); Wrigley and Fagg (1993, 2007) and Venning (1988).
Propagation by seed
Mass propagation of melaleucas is usually by seed, which germinate readily in moist,
warm conditions with no pretreatment. Seed should be sown under shade (optimum
temperature for germination is 2530 C) on a free-draining and sterilised medium and
covered very sparingly with inert material (e.g. sand). Germination should be complete
after 15 days and then shade can be reduced. After germination, the tiny seedlings
can be slow to develop at first, presumably while the roots establish. Once underway,
however, they grow quickly and the 36 months it takes for seedlings to reach plantable
size is similar to other fast-growing species such as eucalypts.
Young seedlings are easily damaged by overhead watering or rain, or may be killed if the sowing mix dries.
Growers in Vietnam have adopted the bog technique of
watering to avoid these problems in propagating M. cajuputi (Figure 23). This involves standing the base of the
germination tray permanently in water so that moisture
soaks up to the surface which is constantly moist but not
flooded. Seed is sown evenly over the surface at a density
of about 7,000 viable seeds/m2. An inflated plastic bag
is fitted over the germination tray to maintain a moist
environment. Once the seedlings are sturdy enough
to withstand overhead watering (c. 4 weeks), the container is removed from the water and handled normally.
The risk of fungal disease is high, so good hygiene is

essential.
Open-rooted seedlings are sometimes used in establishment of M. alternifolia plantations in northern
Queensland. Successful establishment of open-rooted
seedlings is very dependent on the weather at planting
time and/or the availability of irrigation. Container-grown
seedlings, although more expensive to produce than
open-rooted seedlings, suffer much less planting shock
and are less susceptible to the vagaries of the weather
(Colton et al. 2000).
There are two ways of producing container-grown seedlings commonly applied in the propagation of melaleucas:
47
4. Propagation, silviculture and management
Figure 23.Melaleuca propagation in Vietnam using the bog technique for germinating the
fine seeds
(1) the two-stage system where seeds are first sown into
germination trays or germination beds and the seedlings
later transplanted (an operation called pricking out); or
(2) the direct-to-container system where seeds (usually an
average of three per container) are sown directly into individual containers and thinned down to one per container
after germination is completed.
In the two-stage system, seedlings are transplanted
from the germination trays or beds at the second leaf-pair
stage (usually 23 cm tall at 48 weeks after sowing) to
containers (commonly tubes, bags or pots of about 550 cm3
filled volume, e.g. tubes of 65 mm diameter and 160 mm
depth) filled with sterilised potting mix (e.g. 1:1:1 coarse
river sand, perlite and cocopeat with the addition of a
slow-release fertiliser). Extreme care must be taken during
transplanting not to J-root (bend roots upward in a tooshallow planting hole) seedlings as this will cause retarded
growth and instability of the seedling after planting. Shade
cover is needed for the first week after transplanting after
which time plants should be fully exposed. This technique
is usually applied when only a relatively small number
of plants are required and/or seed is in short supply and
efficient capture of all available seedlings is a requirement.
Where very large numbers of seedlings are required, as
in the establishment of M. alternifolia plantations for oil
production with stocking levels commonly in the order of
30,000 plants/ha, the direct-to-container system is widely
applied. Cell-type trays of small individual cell volume
(c. 20 cm3) (e.g. speedling trays) are commonly used
48
in this system. A relatively sophisticated nursery infrastructure, including potting mix and sowing equipment,
plastic igloos or glasshouses, shadehouses and automated
watering systems, is usually employed to produce highquality planting stock at competitive prices for mechanical
planting. Seedlings are routinely topped at about 15 cm to
stop them becoming too tall and spindly and to encourage
a woody stem. Nursery duration under this system is in the
order of 1220 weeks.
Melaleucas form symbiotic mycorrhizal associations between the roots and various fungi. The roots of
M. quinquenervia trees growing on stream banks, or in
fresh or brackish waters in swamps and seepage areas of
New South Wales, Australia, were found to possess both
vesicular-arbuscular (VA) mycorrhizas and ectomycorrhizas (Khan 1993). Nurseries growing melaleucas, especially
where the soils are deficient in phosphorus, should attempt
to introduce appropriate mycorrhizas to the nursery soil.
Various delivery systems, including soil, spores, sporocarps
and vegetative mycelium, are described by Brundrett et al.
(1996) and Doran (1997).
Vegetative propagation
Many melaleucas can be propagated vegetatively from
stem cuttings (Figure 24) and grafts (Wrigley and Fagg
1993) and some have been successfully tissue cultured
(e.g. M. alternifolia; de Oliveira et al. 2010). To ensure the
genetic integrity of cultivars, it is essential that they be
propagated vegetatively.
Silviculture and
management
Melaleucas are used for a range of landcare, wood and
non-wood purposes. The silvicultural system adopted will
depend very much on the end use of the planting, although
it is clear from the lack of literature on the subject that little
is known about optimal stand establishment, tending and
management systems for melaleucas.
Hoang Chuong et al. 1996) and Florida, USA (1 1 m;

Geary 1988), appear appropriate for these end uses. Wider
spacings (e.g. 3 3 m up to 6 6 m) might be employed
where agroforestry is being practised or on sites where very
poor soils are being reforested (Geary 1988).
Practices that include good site preparation, fertilisation
when required and intensive weed control pay dividends
in the cultivation of melaleucas, as with other tree crops
like eucalypts. For example, intensive site preparation by
ploughing to a depth of 20 cm, addition of a nitrogen/
phorphorus/potassium (NPK) fertiliser and manual tending have been found to be beneficial to establishment and
early growth of Melaleuca plantations in the Mekong Delta
region (Simpson 1995). Although pruning is not usually applied in Melaleuca plantations, form pruning has
been advocated for garden specimens of M. leucadendra
(Hearne 1975).
Reported growth rates are reasonable without being
exceptional, even on good sites. For example, M. quinquenervia trees in Hawaiian plantations on good sites
average 18 m in height and 50 cm in diameter at 40 years
(NAS 1983). Annual increments in height of 12 m and in
basal diameter of 13 cm are typical of young plantations
of the broad-leaved melaleucas over a wide range of site
Prastyono et al. (2011) highlighted the potential of

clones in improving oil yields and qualities and, in turn,
the financial returns to producers of essential oil from
M. alternifolia plantations. Readers interested in the mass
vegetative propagation of melaleucas are directed to Chapter 22 in Eldridge et al. (1993) and Chapter 11 in Evans and
Turnbull (2004). Although mass vegetative propagation
of tropical eucalypts is the focus of these detailed descriptions, the methods are directly transferable to the related
genus Melaleuca.
The same principles used in mass vegetative propagation
can be applied on a much smaller scale. Readers interested
in the small-scale vegetative propagation of melaleuca cultivars are directed to the treatments by Wrigley and Fagg
(1993, 2007).
Plantations for wood production

Most interest in growing melaleucas for wood production
is in the tropics on difficult sites for tree growth where the
adaptive traits of the melaleucas give them a competitive
advantage over other, higher value tree crops. It is mainly
the broad-leaved species of the M. leucadendra complex,
such as M. cajuputi, M. leucadendra and M. quinquenervia, that are grown for this purpose in places such as
the Mekong Delta of Vietnam. An important advantage of
the broad-leaved melaleucas over other tree crops under
cultivation in this harsh environment for tree growth is
that they can be established successfully without expensive
and environmentally damaging soil mounding. Mounding
is required to cultivate alternative species and this exposes
the acid-sulfate soils. Species of the M. leucadendra complex are able to survive a fluctuating watertable, including
prolonged seasonal inundation and severe acidity. Other
important advantages in this environment are abilities to
withstand strong weed competition and dry-season fire.
Typically, these species are grown in plantations on
relatively short coppice rotations that maximise the production of small-size logs suitable for posts, piles, poles and
fuelwood. Conventional plantation spacings, such as those
used in trial plantings in Queensland, Australia (1.5 3 m
and 2 3 m; Ryan and Bell 1989), Thailand (2 2 m;
Pinyopusarerk 1989), Vietnam (1.5 2 m and 2 2 m;
Figure 24.Stem cuttings of Melaleuca alternifolia

displaying excellent rooting characteristics
49
types (Morton 1966; Lamb 1975; Ryan and Bell 1989; P.A.
Ryan and R.E. Bell, unpublished report, 1991; Gwaze 1989;
Pinyopusarerk 1989; Sun and Dickinson 1995; Hoang
Chuong et al. 1996). Rotation lengths as short as 35 years
are typical in Vietnam.
Plantations for production of essential oils

The silvicultural systems employed for the production of
essential oils from plantations fall broadly into two categories as highlighted in the two case studies given here.
The first case study is that of M. alternifolia plantations
Australian tea tree oil

Principal source: Plantations of Melaleuca alternifolia
(Maiden & Betche) Cheel (Figure 25) are the main
source of tea tree oil in Australia.
Species description: The mature plant is a shrub or
tree, 2.514 m tall. Its reddish-brown bark is papery,
peeling in long flakes; adult leaves are alternate, linear,
1032 mm long, 0.41 mm wide and short-petiolate
to subsessile, with glabrescent blades and dense oil
glands, more or less in rows (see the M. alternifolia
species account [Chapter 7] for more details).
Natural occurrence: The species occurs from the Stanthorpe district in Queensland, south and east into New
South Wales to the Lismore and Grafton areas, with a
southernmost disjunction near Port Macquarie. It is
found at elevations ranging from near sea level to 800 m.
Climate: Melaleuca alternifolia occurs in warm subhumid climates, with mean maximum temperatures
of hottest and mean minimum of coldest months of
2530 C and 19 C, respectively; frost incidence, low
to moderate (up to 50 at high elevation sites); and rainfall
of 7501,600 mm per year, with a summer maximum.
Topography and soils: The species is found on
coastal plains and adjacent ranges, where it grows on
seasonally inundated swamps and along watercourses.
It grows in soils that are mainly alluvial silty loams
while, in Queensland, soils are sandy loams derived
from granite (soil pH 4.55.5).
Essential oils: Six or more chemotypes have been
identified in the foliar oils of M. alternifolia of which
only one, a terpinen-4-olrich (3048%) type with
more than 100 components, qualifies commercially
as Australian tea tree oil (ISO standard no. 4730; see
ISO 2004). Leaf oil concentration is in the range of
36% (fresh weight).
50
in northern New South Wales and northern Queensland, Australia, which represents an intensive, high-cost
but high-return system. The second case study is that of
M. cajuputi subsp. cajuputi in Java, Indonesia, which is
a less intensive, lower cost but also lower return system.
This second case is representative of silvicultural systems
used in developing countries where Melaleuca plantations must provide a multitude of services for sustainable
development, such as inter-row cropping, rather than oil
production alone, as is the case with M. alternifolia in
Australia.
Uses: Efficacy, stability, oxidation and toxicity of terpinen-4-ol-rich tea tree oil have been closely studied
for many years. It is an effective antiseptic, antibacterial, antiviral, antifungal and anti-inflammatory
agent and is used in a wide range of antimicrobials
and cosmetics. It is also sold as pure oil or in 1015%
tea tree oil solutions.
Quality and prices: Contaminant-free oils with terpinen-4-ol levels of 40% or more in combination with
low levels of 1,8-cineole (i.e. <3%) are demanded by the
principal markets. Oil prices have fluctuated widely in
recent years but in 2013 are around A$30/kg, recovering from a low of A$12/kg in 2005.
Production and markets: Total annual world production of this oil type is in excess of 600 t (Australia
c. 400 t; China c. 200 t; and others). The main markets
are North America and Europe.
Plantations for tea tree oil production: Northeastern New South Wales and the Atherton Tablelands
of Queensland are hubs for production of Australian
tea tree oil from plantations totalling around 3,000 ha.
A typical Australian plantation will be established on
weed-free, level ground at a stocking rate of 30,000
35,000 plants/ha at row spacings that suit available
machinery. Row spacing of 1 m and 30 cm between
plants within rows is commonly applied. Managing
weeds, insect pests/diseases and crop nutrition, combined with use of carefully developed, higher yielding
seed lines, are paramount to optimising production.
Harvesting, distillation and oil storage: Mechanical
harvesting is used in Australian plantations, with the
first harvest taking place at 18 months and annually
thereafter. The best oil yields are in spring and summer. Steam distillation is used to extract the foliar
Yields: Oil yield is determined by three components:

yield of biomass harvested; proportion of leaf in the
total biomass; and oil concentration in the leaves.
Typically, the first harvest of 18-month-old seedlings
will give 50%, second harvest of 12-month-old coppice
75% and third harvest of 12-month-old coppice 100%
of the mature plantation yield. There is wide disparity
between growers in oil yields achieved in practice

due to the many interacting factors involved. Overall,
the average yield from mature coppice plantations in
Australia using unselected seed lines is an estimated
150 kg/ha. Selected seed lines now available from an
Australian industry breeding program have yielded
about 270 kg/ha and further substantial increases are
anticipated through breeding.
Further reading: Southwell and Lowe (1999); Colton
et al. (2000); Davis (2003); ISO (2004); Doran et al.
(2006); RIRDC (2007).
oils. Typical distillation times are 1.52 hours after

condensate starts to flow. Oil should be stored in cool,
dark, dry, air-free conditions to minimise the rate of
oxidation.
Figure 25.Steps in the production of Australian tea tree oil from Melaleuca alternifolia: (A) seedlings grown in
cell-type trays to produce plants well suited to mechanical planting; (B) mechanical planting into a cultivated,
weed-free, drained area; (C) newly planted seedlings being irrigated at establishment; (D) a plantation ready for
harvest; (E) mechanical harvesting into bins; and (F) oil separators in a modern distillery
51
Indonesian cajuput oil

Principal source: Plantations of Melaleuca cajuputi
Powell subsp. cajuputi (Figure 26) are the main source
of this oil in Indonesia.
growth, and moderately dense, obscure oil glands

(see the M. cajuputi species account [Chapter 7] for
more details).
Species description: The mature plant is a shrub or

typically an erect tree, (2)25(46 m) tall. Its bark is
grey to white and papery; adult leaves are alternate,
mainly narrowly elliptic, 40140 mm long, 726
mm wide and petiolate, with glabrescent blades
but silky-hairy on the branchlets and silvery new
Natural occurrence: The subspecies occurs in Indonesia (islands of Buru, Seram, Ambon, Tanimbar in
Maluku province and West Timor) and Australia (Top
End of the Northern Territory and north-western
Western Australia). It is found at elevations ranging
from near sea level to 400 m.
Figure 26.Steps in the production of Indonesian cajuput oil from Melaleuca cajuputi subsp. cajuputi plantations in Java: (A) seedlings grown in polyethylene bags; (B) plantation in Central Java; (C) leafy branches
delivered to a distillery; (D) four of the eight 1-t capacity pots in a cajuput oil distillery at Gundih; (E) a portion
of the dry, spent biomass being bundled for fuelling the distillery boiler; and (F) oil separators in a distillery run
by Perum Perhutani (Forestry Department)
52
Topography and soils: The subspecies is found mainly

on low swampy coastal plains but, in Maluku, mostly
pure stands extend inland on infertile gravelly ridges.
Soils are often highly organic alluvial clays of poor
drainage and low fertility.
Essential oils: There is wide variation in the chemical composition of cajuput oil. The commercial oil
usually contains substantial amounts of 1,8-cineole
(c. 4060%). Leaf oil concentration is in the range of
0.41.2% (fresh weight).
Uses: Cajuput oil is classified as non-toxic and nonsensitising. It is a common household medicine
throughout South-East Asia and is used internally for
treatment of coughs and colds and externally for relief
of pain, often in the form of ointments and liniments.
The oil is useful in treating roundworm and infections
of the genito-urinary system. It is used as a fragrance
and freshening agent in soaps, cosmetics, detergents
and perfumes.
Quality and prices: Contaminant- and adulterant-free
oils with 1,8-cineole levels of 5565% are preferred
by the principal markets. Oil prices at Indonesian
Government distilleries in 2013 are around A$15/kg.
Production and markets: Total world production
of 1,8-cineole-rich cajuput oil is estimated to be
c. 600 t/year, with most oil produced in Indonesia
(300 t from plantations and 90 t from natural stands)
and Vietnam (100 t from natural stands). The main
markets are in South-East Asia.
Plantations for cajuput oil production: The main

source of oil in Indonesia is from 20,000 ha of plantation established on degraded lands on the main island
of Java. Plantations are established at an average stocking rate of 2,000 seedlings/ha. Since 2002, genetically
improved seed from a government breeding program
has been deployed to improve oil yields and quality.
Plantations are intercropped with cassava, maize
and peanuts and participating farmers are required
to weed the cajuput trees when weeding their crops.
There are no major pests or diseases and fertiliser is
not routinely applied.
Harvesting, distillation and oil storage: At 4 years of
age, plants are pollarded at 1.1 m above ground during
the first harvest of essential oils. Thereafter, plants are
visited annually, when coppice shoots of greater than
1 cm diameter are selectively harvested and leaves
and twigs stripped into hessian bags for transport to
the distillery. Peak production in Java is from June to
October when oil yields are highest. Steam distillation
is used to extract the foliar oils. Distillation time is
usually 4 hours. Oil should be stored in cool, dark, dry,
air-free conditions to minimise the rate of oxidation.
Climate: Melaleuca cajuputi subsp. cajuputi occurs

in hot, humid climates, with mean maximum temperatures of hottest and mean minimum of coldest
months of 3133 C and 1722 C, respectively; frost
free; and rainfall of 6004,000 mm/year, monsoonal
with up to an 8-month dry season.
Yields: A plantation of 1 ha established using unimproved seed produces about 7.5 t of cajuput leaves
annually which in turn produces about 6065 kg of
oil. Through use of the improved seed available since
2002, future yields are expected to improve by more
than 20%.
Author: Dr Anto Rimbawanto, Centre of Forest
Biotechnology and Tree Improvement (CFBTI), Yogyakarta 55582, Indonesia.
Further reading: Doran (1999a, b); Susanto et al.
(2003, 2010).
53
Pests, diseases and
other limitations
Pests and diseases
A wide range of insects causing damage to leaves, stems and roots of various Melaleuca
speciesincluding suckers (e.g. bugs, psyllids, froghoppers, scales, galls and thrips)
and chewing pests (e.g. sawflies, caterpillars, beetles and borers)has been described
by Elliot and Jones (1982, 1983), Elliot et al. (1998) and Jones and Elliot (1986), who
also give methods of control.
Over 400 herbivorous insects were found in association with M. quinquenervia and
its close allies in Australia (Balciunas et al. 1993a, b; Burrows et al. 1994) but damage
was reported as localised. Coreid bugs attacked the growing tips of coppice growth
of M. quinquenervia in trials in northern Queensland, reducing yields of essential oil
and requiring application of insecticidal sprays (Doran et al. 2007). This species was
reported as suffering slight damage from sawflies (Marcar et al. 1995) and possessing heartwood that lacked resistance to damage by termites, marine borers and fungi
(Bultman et al. 1983). Damage to M. leucadendra by grasshoppers and leaf-rolling
caterpillars can be severe during the dry season in northern Australia (Hearne 1975).
Of the more than 100 insect species identified in native
stands and plantations of M. alternifolia in Australia,
five have emerged as significant pests in commercial
essential oilproducing plantations. They are pyrgo beetles (Paropsisterna tigrina), psyllids (Trioza spp.), mites
(Eriophyoid spp.), pasture scarabs (Diphucephala lineata)
and leafhoppers (including Erythroneura spp.) (Colton et
al. 2000). All eat flush new leaves or suck their sap and
can cause extensive damage. The sap-sucking leafhoppers also attract ants which in turn promote infestation
by black sooty mould. Economic losses through attack of
M. alternifolia plantations by African black beetle, mole
crickets, cutworms and a wide range of other minor insect

pests have also been reported.
Myrtaceae tip blight and leaf spots can attack melaleucas (Jones and Elliot 1986) and powdery mildew and
grey mould (Botrytris sp.) can develop on cultivated, ornamental melaleucas, especially when dry-region species are
cultivated in humid, subtropical climates.
An introduced disease of Australian plants of the
family Myrtaceae, Puccinia psidii sensu lato (synonym
Uredo rangelii), or myrtle rust as it is commonly called in
Australia, was first observed on the central coast of New
South Wales in April 2010 (Morin et al. 2012). This exotic
55
5. Pests, diseases and other limitations
pathogen (native to South America) has now spread from

Victoria to the Daintree River, north of Cairns in northern
Queensland. Myrtle rust is a form of guava/eucalyptus
rust which has had severe impacts on eucalypt plantations
in Brazil and has spread to other parts of the Americas
(South, Central and North), China and Japan. Melaleuca
quinquenervia, an invasive pest in the Florida Everglades, is
highly susceptible to guava rust in Florida and Hawaii (see
following section on Weediness/biological control) and
also highly susceptible to the rust in Australia. This disease
has so far been observed on 107 host species in 30 genera,
including Angophora, Asteromyrtus, Backhousia, Eucalyptus, Leptospermum and Melaleuca (Carnegie and Lidbetter
2012). There are expectations that many more species will
be found susceptible and this is causing much concern. The
young leaves and shoots of seedlings, the outer growing
tips of the crowns of saplings and, in some cases, adult
trees (e.g. M. quinquenervia) and coppice from stumps or
damaged trees are most vulnerable to attack by myrtle rust.
The rust causes spots or lesions on young leaves and shoots
that spread and develop masses of yellow powdery spores
(Figure 27). The rust can also infect floral buds and young
fruit, depending on the host. Infected leaves become curled
and distorted and severe infection can kill shoots, causing
these plants to become stunted after repeated infections.
In the worst cases, death of the whole plant can occur after
repeated destruction of new growth.
Melaleuca alternifolia in plantations in northern New
South Wales that at first appeared to be resisting the spread
of myrtle rust are now showing signs of greater damage
with the rust-induced death of flush growth and upper
stems becoming more common (Peter Entwistle, pers.
comm. 2012). Other fungal pathogens of M. alternifolia
plantations include stem blight (Dothiorella sp.), with pink
disease (Cylindrocladium sp.) that causes leaf drop, charcoal root disease (Macrophemena phaeseolina or Diplodia
sp.) and leaf scab (Elsinoe sp.) also causing damage (Colton
et al. 2000). These authors also reported grey mould (Botrytis cinerea), anthracnose (Colletotrichum sp.), rhizoctonia
(Rhizoctonia sp.) and damping off (Pythium sp.) to be
important diseases in nurseries growing M. alternifolia
seedlings. Some Western Australian melaleucas are prone
to the rootrot fungus Phytophthora cinnamomi (Wrigley
and Fagg 1993). Colton et al. (2000) reported that no bacterial or viral diseases of economic importance have been
identified in M. alternifolia plantations.
Other limitations
Weediness/biological control
Melaleuca species can seed profusely and there are
instances in Australia where they have escaped cultivation and naturalised to become invasive and troublesome
56
weeds, especially where periodic fires provide a suitable

seedbed. Species that are reported to have naturalised
include M. armillaris, M. bracteata, M. decussata, M diosmifolia, M. ericifolia (per root suckers), M. halmaturorum,
M. hypericifolia, M. incana, M. lanceolata, M. leucadendra,
M. linariifolia, M. microphylla, M. nesophila, M. parvistaminea, M. pentagona, M. quinquenervia, M. styphelioides,
M. viminalis and M. viminea (Lazarides et al. 1997; Randall
2002; Richardson et al. 2011; Wiersema and Len 2013).
Beyond Australia, M. quinquenervia has become a
United States federally listed noxious weed in southern
Florida and is also moderately invasive in the Caribbean (Bahamas and Puerto Rico) and Hawaii (Dray et al.
2006). Melaleuca quinquenervia was first introduced into
Florida as an ornamental and agroforestry species from
Australian and exotic sources. Dray et al. (2006) have
traced the earliest introduction back to 1886 in Sarasota
County, with the species becoming naturalised in southern Florida during the 1920s and spreading rapidly from
there. Since its introduction, the tree has invaded more
than 200,000 ha of Florida wetlands, including portions
of Everglades National Park (Turner et al. 1998). With
its prolific seed production, M. quinquenervia rapidly
invades moist, open habitats, both disturbed and undisturbed, and forms dense, impenetrable monocultures.
Unmanaged stands may have stocking densities of
7,00020,000 stems/ha, thus crowding out native vegetation and wildlife habitats (Geiger 1981; Loope et al. 1994).
Serbesoff-King (2003) reported that public agencies in
Florida had spent US$25 million in control efforts between
1989 and 1999. Serbesoff-King (2003) also gave estimates
of economic impacts of the invasive Melaleuca populations
on recreation, tourism, fires, loss of endangered species
and more. These ranged from US$168 million annually to
US$2 billion over a period of 20 years. It is currently being
suppressed using manual, mechanical, herbicidal and biological control management strategies (Martin et al. 2011).
A classical weed biological control program targeting M. quinquenervia in Florida was initiated in the late
1980s. Surveys in Australia for potential biological control
agents of M. quinquenervia for possible release in Florida
revealed several promising insect species (Center 1992;
Balciunas and Burrows 1993; Balciunas et al. 1993a, b;
Purcell and Balciunas 1994). One herbivore established
for biological control of M. quinquenervia in Florida is a
weevil, Oxyops vitiosa. It was introduced into Florida in
1997 (Christensen et al. 2011) and prefers to feed on the
nerolidol chemotype. Another is a psyllid, Boreioglycaspis
melaleucae, which was released in 2002 and prefers the
viridiflorol chemotype. Predation of M. quinquenervia
by these insects eventually results in partial defoliation
of mature trees, loss of reproductive ability and mortality of seedlings (Martin et al. 2011; Pratt and Arakelian
2011). Tipping et al. (2009), in a 5-year study of a cypress
in plant diversity) and facilitated the partial rehabilitation

of degraded habitats.
Fungi are also under investigation as potential biological
control agents of M. quinquenervia in Florida (Rayachhetry
et al. 1996a, b). Puccinia psidii, as detailed in the previous
section on Pests and diseases, is one possibility. In the
early 2000s, P. psidii was observed on M. quinquenervia
in Florida (Rayachhetry et al. 2001). It has now joined the
introduced herbivores as effective biological control agents
pine wetland in the West Everglades invaded by M. quinquenervia after a destructive crown fire, reported a 48%
decline in Melaleuca density over 5 years due to biological
control agents. Annual mortality ranged from 11% to 25%
and mean tree height declined by 31%. Rayamajhi et al.
(2009) found rapid reduction in Melaleuca density and
canopy cover, attributed to self-thinning accelerated by the
negative impact of the introduced insect pests, positively
influenced native plant diversity (two- to fourfold increases
Figure 27.Puccinia psidii sensu lato (synonym Uredo rangelii) (myrtle rust) spores on
Melaleuca quinquenervia in northern New South Wales: (A) yellow spores on a leafy
shoot; and (B) a badly deformed and stunt young plant after rust attack of its growing tips
57

58
of M. quinquenervia in Florida although there are resistant

individuals (Rayamajhi et al. 2010a, b). Regrettably, it has
been found to also attack some native American species,
including a threatened species.
Source of allergens
Earlier reports implicating M. quinquenervia in southern
Florida as the cause of serious allergic reactions and acute
respiratory problems in humans (Geary 1988) have been
shown to be false in a detailed medical study involving
more than 1,000 subjects (Stablein et al. 2002).
Conservation and
prospects
Conservation status
An estimated 100 million ha of the Australian landscape have been cleared for agriculture, urban development, mining and other pursuits. In addition to clearing of forests
and woodlands, drainage and flood mitigation measures, waterlogging from irrigation
and increased salinity have all adversely affected the extent of natural populations of
Melaleuca. Australia accounts for 20% of the worlds flora that has been classified as
presumed extinct and 15% of the worlds flora that has been recognised as threatened
(Briggs and Leigh 1995). It is somewhat surprising, therefore, to report that apparently
no species within this large plant genus have been classified as presumed extinct.
Wrigley and Fagg (1993) reported that M. arenaria, a species described in 1923 from
a specimen collected in the Western Australian wheatbelt in an area subject to much
clearing, was presumed extinct, but this species is now considered to be a variety of
the widespread M. tuberculata (see Species accounts [Chapter 7]).
Briggs and Leigh (1995) listed 48 Melaleuca taxa (including

Callistemon) in their compendium of rare and threatened
Australian plants. Melaleuca kunzeoides, from central
southern Queensland, M. sciotostyla, from south-western
Western Australia, and Callistemon sp. 1 (Boulia, L.Pedley
5297)now classified within the widespread M. viminalis
subsp. viminaliswere listed as vulnerable; with only
one of these species, M. sciotostyla, protected in reserves
in 1995. Eighteen species, 15 of which were in reserves or
National Parks in 1995, were classed as rare (Callistemon
acuminatus [= M. flammea], M. basicephala, M. cheelii,
M. chisholmii, M. cliffortioides, M. corrugata [= M. fulgens
subsp. corrugata], M. deanei, M. fissurata, M. flavovirens,
M. formosa, M. groveana, M. linearifolia, M. pauciflora,
M. pearsonii, M. pungens, M. pustulata, M. shiressii and
M. tortifolia); and the remaining taxa were placed in category K. Category K is for species known to be limited
in distribution but whose conservation status cannot be
reliably determined, either because the species has been
seldom collected or there is uncertainty about the level of
threat. The list of rare or threatened Australian melaleucas
needs to be revised, as many very localised and/or rare
species have been described since 1995.
Outside Australia, there have been concerns about the
decline of Melaleuca forests and woodlands of M. cajuputi
subsp. cumingiana in the wetlands of South-East Asia.
Clearing and draining of the Melaleuca forests for rice
production and other crops in places such as the Mekong
Delta region of Vietnam have led to environmental degradation, loss of biodiversity and social consequences for
59
6. Conservation and prospects

60
local peoples (Safford et al. 2009). While not to the extent

yet of endangering the survival of the species in these
wetlands, there are, nevertheless, compelling reasons to
rehabilitate selected areas of these forests and woodlands.
This has been a priority in Vietnams forest policy since
the mid 1990s.
Prospects
Opportunities for wider use
Reasonable growth rates in the face of extremely poor
environmental conditions for plant growth and a broad
range of uses are among the desirable attributes of the
Melaleuca species regularly deployed in reforestation,
land reclamation, amenity and ornamental plantings and
for production of essential oils. With a predominance
of species occurring in arid and semi-arid regions, but
with a range from the humid tropics to cool temperate
southern Australia and on highly variable soils and topography, it is possible to select species that are tolerant of
a wide range of unfavourable conditions (infertile soils,
poorly drained sites, continuous and periodic inundation,
coastal exposure, fire, frost, salinity and both high and
low soil pH). Uses, depending on species/provenances
or cultivars, include ornamental and amenity planting,
essential oils, fuelwood, woodchips, sawn timber, posts,
poles, rails, brushwood fencing, shade and shelter, honey,
land reclamation and improvement in biodiversity values.
In Appendix 2, we have endeavoured to highlight by end
use, best-bet species for planting/trialling in two broad
climatic zones: (A) subtemperate and (B) tropical and
subtropical.
Melaleucas are largely outbreeding, often with heritable and highly variable commercial traits (e.g. foliar
oil concentrations and various growth characteristics,
including inflorescence shape and flower colour). This
provides a huge opportunity for the tree breeder, whose
main task is to exploit this variability through exploration, evaluation, selection and breeding. Nowhere is this
more so than with the ornamental Melaleuca cultivars
that after manipulation by controlled pollination, either
within or between species, must be propagated vegetatively to capture desired characteristics (e.g. inflorescence
shape and colour). There is also great opportunity for
selection and breeding to improve oil yields and oil
qualities in the established essential oilproducing
species M. alternifolia, M. cajuputi subsp. cajuputi and
M. quinquenervia. These species all have distinctly different chemical variants of which only one (or two in
the case of M. quinquenervia) of several types found in
nature is suitable for commercial exploitation. So it is
very important to select the provenance(s) within species
that will reliably provide the required oil as well as the
ability to grow rapidly and coppice well so that oil yields

are maximised.
Once the best natural provenances are identified,
further economic gains can be achieved by selection
between and within families established in progeny trials and development of clonal or seedling seed orchards
to provide improved seed. This is well demonstrated by
a traditional, relatively low-cost, seed-based breeding
program for M. alternifolia in Australia (Figure 28). This
program has delivered to industry realised genetic gains
in oil yield from improvements in foliar oil concentration
and leaf biomass per ha in one generation of breeding:
55% from selections within the best natural provenance;
43% from a culled, broadly based seedling seed orchard;
and 83% from a clonal seed orchard established by
stem cuttings from selected individuals within the best
provenances in a progeny trial (Doran et al. 2006). Even
greater genetic gains are expected to be realised from the
second generation of selection and breeding in this species. Similar results have been achieved in the breeding
of M. cajuputi subsp. cajuputi in Indonesia where gains
in oil yield in excess of 20% are anticipated from the first
generation of selection and breeding in open-pollinated
seedling seed orchards (A. Rimbawanto, Indonesian
cajuput oil section in Chapter 4).
Caution
High on the list of undesirable traits, particularly when
introducing melaleucas to a new environment, is the
potential for their spread from cultivation to become
noxious weeds. This occurs through distribution of seed by
wind and water from canopies that hold a store of mature
fruit, often for many years, awaiting the right conditions to
stimulate release (e.g. fire) and also root suckering which
is a feature of some melaleucas with extensive root systems
(e.g. M. ericifolia, M. viridiflora). The experience with the
M. quinquenervia invasion of the Florida Everglades is a
classic example of an inappropriate species introduction
that has gone horribly wrong, with the aggressive, fastgrowing invader crowding out regeneration of native
species and destroying wildlife habitat. Thus, extreme
caution is warranted when introducing a Melaleuca to a
new environment for the first time, and particularly, it
seems, in swampy conditions. Another disadvantage is the
susceptibility of certain of the more tropical species, such
as M. leucadendra and M. quinquenervia, to fungal attack
at a young age by the rust, Puccinia psidii sensu lato. Insect
pests are also an impediment to the successful establishment and growth of some species (e.g. in the cultivation
of M. alternifolia for essential oil production), requiring
use of chemical sprays. Despite these disadvantages, there
will be localities where the genus Melaleuca can provide
the species-of-choice for the prevailing conditions and
intended end use.
Figure 28.Progeny trials (A, B), a young seedling seed orchard (C) and controlled crossing activities (DF) as
part of a tree breeding project aimed at improving oil yields in Melaleuca alternifolia in Australia
61

62
Advice is at hand
Prospects for wider exploitation of carefully selected
germplasm of Melaleuca species appropriate for intended
end use(s) both within and beyond their zones of natural
occurrence appear promising. When considering introduction of a Melaleuca species to a location for the first
time, plant risk analysis procedures should be applied and
the species rejected if the weediness risk is unacceptable.
The Australian Tree Seed Centre, CSIRO Plant Industry,
Canberra, holds seed stocks of a wide range of mainly
tree-form melaleucas and is a source of both seed and
information on cultivating species in the genus.

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MELALEUCAS

THEIR BOTANY, ESSENTIAL OILS AND USES

Joseph J. Brophy, Lyndley A. Craven

The Australian Centre for International Agricultural Research (ACIAR) was

ACIAR MONOGRAPH SERIES

Australian Centre for International Agricultural Research (ACIAR) 2013

2. Introduction to the genus Melaleuca............................................................................................................................19

Geographical distribution and ecology................................................................................................................................26

4. Propagation, silviculture and management........................................................................................................... 47

5. Pests, diseases and other limitations............................................................................................................................ 55

6. Conservation and prospects............................................................................................................................................... 59

Opportunities for wider use..................................................................................................................................................60

Appendix 1Oil types by species.........................................................................................................................................397

M. linearis var. pinifolia

Callistemon pungens subsp.

Callistemon pungens subsp.

M. linearis var. linearis

M. linearis var. linearis

Callistemon rugulosus var.

Callistemon macropunctatus M. rugulosa

Callistemon viminalis subsp. M. viminalis subsp.

Melaleuca minutifolia subsp. M. monantha

Callistemon viminalis var.

Melaleuca nervosa f. latifolia M. nervosa

Melaleuca nervosa subsp.

Melaleuca acacioides subsp.

Melaleuca styphelioides var.

Melaleuca calycina subsp.

Melaleuca urceolaris var.

Melaleuca cardiophylla var.

Melaleuca viridiflora var.

Melaleuca citrina Turcz.

Melaleuca coccinea subsp.

Melaleuca viridiflora var.

Melaleuca coccinea subsp.

Melaleuca densa var. pritzelii M. pritzelii

M. linearis var. pinifolia

Metrosideros quinquenervia M. quinquenervia

Melaleuca lanceolata subsp.

Melaleuca lanceolata subsp.

Melaleuca lanceolata subsp.

Melaleuca lateriflora subsp.

Melaleuca lateriflora var.

farmland, for rehabilitating salt-affected lands, for street

to northern Queensland. In susceptible plants, young

of Melaleuca species, George Benthams treatment of the

1. Taxonomic history and systematics

The related genus Callistemon was distinguished from

of Melaleuca in 1979 which gave an insight into arid-zone

agreement with the nuclear DNA phylogenies of Ladiges et

Current and future

1. Taxonomic history and systematics

of plants in detail for the first time. The consequences of

1. Taxonomic history and systematics

also occurs naturally in Papua province of Indonesia,

type conducted by Broadhurst et al. (2004) on members of

the genus Melaleuca

(2n + 2 = 24), triploidy (3n = 33), tetraploidy (4n = 44)

2. Introduction to the genus Melaleuca

Melaleuca, as circumscribed in this work, comprises 290

The bark of the majority of species is of the papery type.

Figure 1.A substantial

2. Introduction to the genus Melaleuca