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A Review of Leptospermum Scoparium (Myrtaceae) in New Zealand

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A Review of Leptospermum Scoparium (Myrtaceae) in New Zealand New Zealand Journal of Botany ISSN: 0028-825X (Print) 1175-8643 (Online) Journal homepage: http://www.tandfonline.com/loi/tnzb20 A review of Leptospermum scoparium (Myrtaceae) in New Zealand J. M. C. Stephens , P. C. Molan & B. D. Clarkson To cite this article: J. M. C. Stephens , P. C. Molan & B. D. Clarkson (2005) A review of Leptospermum scoparium (Myrtaceae) in New Zealand, New Zealand Journal of Botany, 43:2, 431-449, DOI: 10.1080/0028825X.2005.9512966 To link to this article: http://dx.doi.org/10.1080/0028825X.2005.9512966 Published online: 17 Mar 2010. Submit your article to this journal Article views: 1624 View related articles Citing articles: 22 View citing articles Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=tnzb20 Download by: [119.224.93.141] Date: 13 December 2016, At: 11:44 New Zealand Journal of Botany, 2005, Vol. 43: 431–449 431 0028-825X/05/4302-0431 © The Royal Society of New Zealand 2005 A review of Leptospermum scoparium (Myrtaceae) in New Zealand J. M. C. STEPHENS Keywords Myrtaceae; Leptospermum scoparium; P. C. MOLAN manuka; tea tree; New Zealand; taxonomy; chemo- taxonomy; ecology; history; ornamental; essential B. D. CLARKSON oils; honey; pharmacology Department of Biological Sciences University of Waikato Private Bag 3105 Hamilton, New Zealand INTRODUCTION [email protected] Information about Leptospermum scoparium is spread throughout much literature. L. scoparium is the most widespread and important indigenous shrub Abstract Information about Leptospermum sco- species in New Zealand, and has probably undergone parium (Myrtaceae), the most widespread and im- the most varied development as an economic plant in portant New Zealand indigenous shrub species, is the indigenous flora. Bibliographies compiled by Or- reviewed. L. scoparium is a variable species, requir- win ( 1974) and Williams (1981) provide background ing more study of the genetically based differences material to this review. Whilst all material that refers between New Zealand populations and the affinity to L. scoparium in passing has not been included, of these populations to Australian populations and each section draws upon the principal publications. other closely allied Australian species. Improved Chemical control aspects are not included. understanding of the species' variation will assist The taxonomy, morphology, distribution, habi- both its conservation roles and economic uses, and tats and plant communities, and other biological the need to sustain genetically distinct varieties is associations in which it occurs are considered here, emphasised. Ecologically, the species has a dominant along with its historic and current uses as a source role in infertile and poorly drained environments, of essential oils and honey, and for ornamental shrub and a wider occurrence as a seral shrub species in development. The majority of early research consid- successions to forest where it may be regarded as a ered the ecological position of L. scoparium, either woody weed of pasture or a useful species for ero- as a dominant species where environmental stress is sion control, carbon sesquestration, and vegetation extreme, or as a seral species in disturbed habitats restoration. The main economic products derived where the species was recognised as a significant from the species are ornamental shrubs, essential weed of recently cleared forest for pasture develop- oils, and honey. The species' development as an ment. Recognition of the genetic and phenotypic ornamental plant and further definition of the phar- variation exhibited by the species led to studies macologically active components are recommended isolating various components of this diversity, rang- as priority areas for research. ing from morphological to chemotaxonomic treat- ments. Commercial development as an ornamental shrub and the identification of medicinal essential oil and honey components have motivated most of the recent research, and a collation of this material is warranted. We consider it timely to review the biology of L. scoparium as a basis for further economic develop- B04037; Online publication date 5 May 2005 ment of the species and conservation of genetic Received 16 September 2004; accepted 29 March 2005 variation. 432 New Zealand Journal of Botany, 2005, Vol. 43 BIOLOGY AND ECOLOGY of L. scoparium also support the species genus clas- sification. Taxonomy, morphology, anatomy, cytology A comprehensive taxonomic revision of the genus Leptospermum scoparium J.R. et G.Forst. (manuka, Leptospermum listed 79 species (Thompson 1989), kahikatoa, tea tree, red tea tree) is a member of the which has been increased to 83 with later additions Myrtaceae. This family contains at least 133 genera (Dawson 1997a). L. scoparium is one of 13 species and more than 3800 species, and has evolutionary included in the Leptospermum myrtifolium subgroup, centres in Australia, Southeast Asia, and Central the defining characteristics of which are deciduous and temperate South America. Myrtaceae are char- sepals and persistent strongly wooded fruit-valves acterised by a half-inferior to inferior ovary, usu- (Thompson 1989). The Australian species within ally numerous stamens, entire leaves containing oil this sub-group are extremely difficult to define; L. glands, internal phloem, and vestured pits on the continentale and L. rotundifolium were recently xylem vessels (Wilson et al. 2001). elevated by Thompson (1989) from L. scoparium Until recently Myrtaceae was divided into two varieties to species rank. The species L. juniperi- subfamilies, the capsular Leptospermoideae and the num and L. squarrosum have both been recorded as fleshy-fruited Myrtoideae. An extensive review of varieties ofL. scoparium (Thompson 1989), and the the Myrtaceae inflorescence structure confirmed this endemic TasmanianL. scoparium var. eximium could division; the Leptospermoideae contained seven al- be considered to warrant species status, displaying liances including the Leptospermum alliance, which lignotuber development which is not found in New was further subdivided into the Leptospermum and Zealand's L. scoparium (Bond et al. 2004). Calothamnus suballiances (Briggs & Johnson 1979). Initially three species of Leptospermum were re- However, cladistic analysis of morphological and corded as endemic to New Zealand; the widespread anatomical characters concluded that the subfamilies Leptospermum scoparium and L. ericoides, and L. should be discarded, as the fleshy-fruited Acmena sinclairii restricted to Great Barrier Island (Allan alliance did not group within the Myrtoideae sub- 1961). Revision of Leptospermum led to the transfer family (Johnson & Briggs 1984). A cladistic re- of L. ericoides to Kunzea as K. ericoides (A.Rich.) evaluation of non-molecular characters confirmed J.Thompson (Thompson 1983). L. sinclairii was a high level of homoplasy within Myrtaceae and included in synonomy to this species, and a new limited support for any clade (Wilson et al. 1994). name combination of K. sinclairii (Kirk) W.Harris Molecular analysis placed further doubt on the was later published without supporting material traditional taxonomic groupings. Sequences of the (Connor & Edgar 1987). Accordingly, L. scoparium chloroplast matK gene analysed in association with is now considered to be the only indigenous member nonmolecular data revealed that the Leptospermum of Leptospermum in New Zealand. The species is alliance was polyphyletic and, thus, an invalid taxo- not endemic to New Zealand as indicated by Allan nomic concept (Wilson et al. 2001). The sequencing (1961), as it also occurs naturally in mainland Aus- of two chloroplast regions for 31 species within the tralia from the southern coast of New South Wales Leptospermum suballiance revealed a monophyletic to western Victoria and is widespread in Tasmania grouping of eight genera and the suballiance was (Thompson 1989). considered a valid taxonomic unit (O'Brien et al. The time of arrival of L. scoparium in New Zea- 2000). However, the same study concluded that land is uncertain. Leptospermum pollen has been Leptospermum is polyphyletic and should be divided dated to the Paleocene (Fleming 1975), though the into at least four genera: the persistent-fruit group, representatives in the upper Cretaceous and older the East Australian non-persistent-fruit and West Tertiary beds should be interpreted to represent type Australian non-persistent-fruit groups, and Lept- pollen and not individual species (Couper 1953, ospermum spinescens separated as a fourth genus. 1960). Thompson (1989) suggested that Leptosper- L. scoparium was not included in this analysis but mum may have originated in the dry Miocene con- its fruit morphology allies it to the persistent-fruit ditions in Australia and that L. scoparium dispersal group. to New Zealand occurred relatively recently, as the Analysis of leaf anatomy of 40 species of Lept- species is not a primitive Leptospermum and cannot ospermum showed that L. scoparium has the typi- have been present earlier in New Zealand. War- cal xeromorphic structure of the genus (Johnson dle (1991) recorded L. scoparium as the only New 1980). The wood anatomy (Johnson 1984; Patel Zealand species to release seed overwhelmingly in 1994) and the pollen morphology (McIntyre 1963) concert after fire, a serotinous feature common in the Stephens et al.—Review of Leptospermum scoparium in NZ 433 Australian flora. Further evidence for the recent evo- but two triploid and one
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