Professional Documents
Culture Documents
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
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
Contents
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
Conservation status......................................................................................................................................................................... 59
Prospects................................................................................................................................................................................................60
Contents
7. Species accounts............................................................................................................................................................................. 63
Acknowledgments............................................................................................................................................................................395
Photograph credits........................................................................................................................................................................ 396
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
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
M. williamsii subsp.
fletcheri
Callistemon gnidioides
M. sphaerodendra
Callistemon hemistictus
M. hemisticta
M. williamsii subsp.
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
Callistemon linearis
Callistemon rugulosus
M. rugulosa
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
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
M. viminalis subsp.
viminalis
Callistemon viridiflorus
M. virens
M. nervosa
Callistemon wimmerensis
M. wimmerensis
Melaleuca paludosa
M. glauca
Calothamnus suberosus
M. suberosa
Melaleuca smithii
M. quinquenervia
M. alsophila
M. squamophloia
Melaleuca acerosa
M. systena
M. irbyana
Melaleuca apodocephala
subsp. calcicola
M. calcicola
Melaleuca tamariscina
subsp. irbyana
M. pallescens
Melaleuca arenaria
M. tuberculata var.
arenaria
Melaleuca tamariscina
subsp. pallescens
M. dichroma
M. dempta
M. dichroma
M. longistaminea
M. quinquenervia
M. lutea
M. quinquenervia
M. eximia
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
Metrosideros pinifolia
Melaleuca halmaturorum
subsp. cymbifolia
M. halmaturorum
M. lanceolata
M. lanceolata
M. lanceolata
M. 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
13
Preface
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
15
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
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
17
Introduction to
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
19
Number of species
Bark
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
20
Figure 2.Bark types in Melaleuca: papery, as in (A) M. exuvia; and rough, as in (B) M. bracteata
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).
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.
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
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
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
26
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
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
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
31
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.
33
3. Uses
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
3. Uses
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
36
3. Uses
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
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
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
3. Uses
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
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
42
3. Uses
50
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
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
44
3. Uses
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.
47
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.
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.
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
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
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
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
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
55
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
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
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]).
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
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
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
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.