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Journal ArticleDOI

Four new species of Syzygium (Myrtaceae) from Australia

01 Jan 2003-Blumea-Vol. 48, Iss: 3, pp 479-488
TL;DR: The following new species of Syzygium are described from Queensland, Australia: S. fratris, S. glenum, S., monimioides and S. monospermum.
Abstract: The following new species of Syzygium are described from Queensland, Australia: S. fratris, S. glenum, S. monimioides and S. monospermum. Notes are provided as to their putative relationships.

Summary (1 min read)

Jump to:  and [Specimens studied:]

Specimens studied:

  • Syzygium monospermum Craven, spec. nov. -Fig. 2a A S. erythrocalyx (C.T. White) B. Hyland planta semper cauliflora, foliis venis tertiariis obscure scalaribus, et fructu alba 1-seminali differt.
  • Branchlets compressed or terete, rounded, 1.4-2.8 mm diam.; bark dull, smooth, slightly striate or cracked.
  • Reproductive seasonal growth unit with a reproductive zone only.
  • Inflorescence leafless, on warty outgrowths on the trunk, 2-to few-flowered,.

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BLUMEA 48: 479 488
Published on 28 November 2003 doi: 10.3767/000651903X489438
FOUR NEW SPECIES OF SYZYGIUM (MYRTACEAE)
FROM
AUSTRALIA
L.A. CRAVEN
Australian National Herbarium, CPBR, CSIRO Plant Industry, G.P.O. Box 1600,
Canberra, ACT 2601, Australia
SUMMARY
The following new species of Syzygium are described from Queensland, Australia: S. fratris, S. glenum,
S. monimioides and S. monospermum. Notes are provided as to their putative relationships.
Key words: Acmena, Cleistocalyx, Piliocalyx, Syzygium, Myrtaceae, Australia.
INTRODUCTION
The Australian species of Syzygium Gaertn. and its closer generic relatives were recently
revised by Hyland (1983). Hyland accepted five genera within the complex, Acmena
DC., Acmenosperma Kausel, Eugenia L., Syzygium, and Waterhousea B. Hyland, treat-
ing the Australian species at that time referred to Cleistocalyx Blume within his concept
of Syzygium (Hyland, 1983). During work on an account of the generic complex for
Flora of Australia (Craven & Matarczyk, in press) it became apparent that to accept the
genera recognised by Hyland would require the description of several new genera to
accommodate anomalous species. These species were either unknown or inadequately
collected when Hyland’s work was published but at least one species treated by Hyland,
S. wesa B. Hyland, is incongruent within the genus to which he assigned it.
Eugenia and Syzygium often have been treated as congeneric (e.g. Henderson, 1949;
Kochummen, 1978) whereas others have maintained them as distinct genera (e.g. Mer-
rill, 1950; Kostermans, 1981). Schmid’s studies of morphological and anatomical data
indicated that two separate lineages were involved (Schmid, 1972a, b) and this view
has been supported by some recent molecular work. Analyses of matK sequence data
and non-molecular data have shown that Syzygium and Acmena form a clade with four
other Australasian genera (Anetholea Peter G. Wilson, Osbornia F. Muell., Backhousia
Hook. & Harv. and Choricarpia Domin) that is sister to a clade containing both Old and
New World genera (Archirhodomyrtus (Nied.) Burret, Pilidiostigma Burret, Rhodamnia
Jack, Calyptranthes Sw., Eugenia and Xanthomyrtus Diels, Wilson et al., 2001).
While the accumulated evidence that Eugenia and the Syzygium generic complex rep-
resent distinct evolutionary lineages may now have resolved that particular issue, there
remains considerable uncertainty as to the circumscription of Syzygium. In publications
of world-wide scope, Merrill (1950) accepted seven (and tentatively an eighth) genera
that belong to the Syzygium complex, Airy Shaw (1966) three, Kausel (1957) ten, and
Briggs & Johnson (1979) eight. In more geographically restricted works, often there has
© 2003 Nationaal Herbarium Nederland, Leiden University branch

BLUMEA Vol. 48, No. 3, 2003
480
been considerable variation as to which generic segregates are accepted. In a study of
Australian species, Hyland (1983) recognised Acmena, Acmenosperma, Syzygium (incl.
Cleistocalyx) and a fourth genus, Waterhousea, an Australian endemic. In contrast, in
an account of the syzygioid species in Fiji, Smith (1985) recognised Cleistocalyx as a
genus distinct from Syzygium, together with Syzygium and Piliocalyx Brongn. & Gris,
the last named genus occurring in New Caledonia, Vanuatu and Fiji. Chantaranothai
& Parnell (1994a, b) and Parnell (1999), in dealing with Thai species of the syzygioid
group, recognised Acmena, Cleistocalyx and Syzygium (incl. Acmenosperma). For
Malaya, Turner (1997) preferred to recognise Syzygium only (incl. Acmena, Acmeno-
sperma, Cleistocalyx). The issue was discussed by Craven (2001) who concluded that
recognition of a single genus, i.e. Syzygium, was the only practicable solution in the
light of present knowledge.
As a result of the work towards the accounts of the syzygioid genera for Flora of
Australia mentioned above, four undescribed species were identified from north-eastern
Queensland; these are newly described below and placed in Syzygium. Where relevant,
notes are given on the placement of the species relative to genera accepted by other
workers.
The reproductive seasonal growth unit that is described for each of the four species
in this paper is defined as the shoot or flush of growth that includes the inflorescence.
This growth unit may have a leafy zone in addition to the inflorescence per se or may
consist entirely of the inflorescence.
1. Syzygium fratris Craven, spec. nov. — Fig. 1a
A S. canicortex B. Hyland oribus sepalis discretis et petalis perfecte evolutis differt.
— Typus: L.A. Craven, R. Jensen & W. Cooper 10421 (holo BRI; iso A, BO, CANB, L,
LAE, MEL, P), Australia, Queensland, Mt Bartle Frere, NW Peak, 2 December 2001.
Tree to 10 m tall, to 10 cm dbh. Bark nondescript. Branchlets compressed or quadran-
gular, winged, the wings not joined at the base, 0.5–1 mm diam.; bark dull, smooth or
cracked. Leaf lamina elliptic, broadly elliptic or ovate, 3.5 8.5 by 1.33.4 cm, 1.9 2.9
times as long as wide, base cuneate or attenuate, apex long acuminate (tip of the acumen
rounded or obtuse), acumen plane or recurved, margin revolute, entire, midrib rounded
on the abaxial surface, primary and secondary venation generally similar with all or
nearly all secondaries joining the intramarginal vein, primary veins 10 –14 on each side
of the midrib, divergence angle more or less uniform, in the median part of the lamina
the veins at an angle of 55–70° and 1–3.5 mm apart, intramarginal vein present and
0.5–1.4 mm from the margin, secondary intramarginal vein present (obscure and only
visible after the removal of lamina surface tissues), oil dots visible to the unaided eye
in transmitted light, the dots small, sparse. Petiole 24.5 by 0.6 0.8 mm, axillary
enations often present, 0.4–1.4 mm long. Reproductive seasonal growth unit leafless.
Inflorescence among the leaves, terminal, few-flowered and then cymose or spicate or
1-flowered, main axis straight and rigid, 0.5–1.6 by 1– 2 cm, 0.5–1 mm thick at the mid
point; bracteate, the bracts deciduous; bracteoles subtending each flower, or subtending
the lateral flowers of a triad with the terminal flower ebracteolate, deciduous. Flowers
not calyptrate. Hypanthium dull, obscurely gland dotted, stipitate, stipitately narrowly
funnel-shaped or elongated-goblet-shaped, 5.5–7.5 by 2.5 4.5 mm, the stipe 1.5 2 mm

L. A. Craven: Four new species of Syzygium from Australia
481
Fig. 1. A. Syzygium fratris Craven, owering branchlets; B. Syzygium monimioides Craven, owering
branchlets (A: Craven, Jensen & Cooper 10421 (CANB); B: Forster, Sankowsky & Tucker 10750
(QRS)). — Scale bars = 1 cm.
long. Sepals 4, uniform in size or nearly so, transversely semicircular or transversely
narrowly semi-elliptic, persistent, not accrescent, 0.1–0.5 mm long. Petals 4, decidu-
ous, not coherent, 2 by 22.5 mm, margin entire or lacerate. Staminal disc unmodified
or prominent. Stamens all fertile, outermost stamens 24 mm long; anther sacs paral-
lel; anthers oblong, 0.30.5 mm long, dehiscing by longitudinal slits, connective not
glandular. Style 4 6.5 mm long at anthesis. Placentation axile-median, the placenta

BLUMEA Vol. 48, No. 3, 2003
482
weakly developed, narrowly oblong or sublinear (more or less flange-like). Ovules 4–6
per locule, pendulous, arranged in two longitudinal rows. Immature fruit smooth or
glandular-verruculose, barrel-shaped, perhaps less than 12 by 7 mm at maturity, with
the calyx rim not appreciably expanding in fruit, pericarp distinctly free from the seed
and without peg-like intrusions into the seed. Seed 1, spheroidal, perhaps c. 4 mm diam.
at maturity, testa absent, uni-embryonic; embryo with the cotyledons readily separable
and with no interlocking placental tissue present, equal in size or nearly so.
Distribution — Australia (Queensland, Mt Bartle Frere, NW Peak).
Habitat — Rain forest; 1400–1500 m altitude.
Notes 1. Flowers have been recorded in May and December. Fruits have been
recorded in March.
2. The most closely related species to S. fratris may be S. canicortex B. Hyland. In
both species the placentation is axile-median, the ovules are pendulous in two longitu-
dinal rows in each locule, and the leaves are fairly similar in shape and size. However,
the two species may be distinguished as follows: S. canicortex: flowers calyptrate (the
calyptra is sepaline in origin and the petals adhere to and fall with the calyptra) and
the fruit consequently lack sepals, leaves with axillary enations 0.20.5 mm long;
S. fratris: flowers with 4 sepals and petals, the sepals persistent through to the mature
fruit, leaves often with axillary enations, these 0.4–1.4 mm long.
3. The epithet is derived from the Latin frater, brother, in reference to the locality Mt
Bartle Frere which itself was named in honour of Henry Bartle Edward Frere, a former
Governor of Bombay and a former president of the Royal Geographical Society.
Specimens studied:
Cooper & Cooper 1735; Craven, Jensen & Cooper 10419, 10420, 10421 (type); Gray 3949; Gray
in Hyland 25917RFK; Hyland 12923; Jensen 805.
2. Syzygium glenum Craven, spec. nov. — Fig. 2c, d
A S. gustavioides (F.M. Bailey) B. Hyland ovario placentis axialiapicalibus, et semini-
bus cotyledonibus coalitis et contexto fungiformi intruso differt. — Typus: W. Cooper &
W. Cooper 979 (holo CANB; iso QRS), Australia, Queensland, Daintree River region,
Turpentine Road (N of Cow Bay), 21 June 1996.
Tree to 25 m tall, to 70 cm dbh. Bark reddish-brown-grey; flaky. Branchlets terete or
rounded-compressed, 1.32.2 mm diam.; bark dull, smooth or slightly striate. Leaf
lamina narrowly elliptic, elliptic or narrowly ovate, 8.4–17.9 by 3.2 6.7 cm, 2.43.4
times as long as wide, base cuneate, obtuse or attenuate, apex acuminate (tip of acumen
acute or occasionally rounded), acumen plane, margin revolute, entire, midrib angled
or rounded on the abaxial surface, primary and secondary venation distinctly differ-
ent with the secondaries relatively little developed and not joining the intramarginal
vein, primary veins 1942 on each side of the midrib, divergence angle more or less
uniform or with the distal veins more acute than the proximal, in the median part of
the lamina the veins at an angle of 55–70° and 3–11 mm apart, intramarginal vein
present and 1.66 mm from the margin, secondary intramarginal vein present and a
third is usually present, oil dots visible to the unaided eye in transmitted light, the dots
small, sparse. Petiole 7.5–16 by 1.3 2 mm, stipules and/or axillary enations absent.
Reproductive seasonal growth unit with distinct vegetative and reproductive zones or

L. A. Craven: Four new species of Syzygium from Australia
483
with a reproductive zone only. Inflorescence leafless, among the leaves, terminal or
distal axillary, few- to many-flowered, paniculate, main axis straight and rigid, 340
by1.5–17 cm, 1–2 mm thick at the mid point; bracteate, the bracts deciduous; bracte-
oles subtending the lateral flowers of a triad but with the terminal flower ebracteolate,
deciduous. Flowers calyptrate (the calyptra formed from fused sepals only, the petals
cohere to the calyptra and fall with it), the calyptra 0.75–1.5 mm long. Hypanthium
Fig. 2. A. Syzygium monospermum Craven, abaxial leaf surface; B. Syzygium erythrocalyx (C.T.
White) B. Hyland, abaxial leaf surface; C, D. Syzygium glenum Craven. C. Longitudinal section
through a fruit showing pericarp, embryo (the white tissue) and the intrusive mushroom-shaped
tissue; D. transverse section through a fruit (A: Hyland 2960RFK (CANB); B: Webb 945 (CANB);
C & D: Cooper & Cooper 979 (CANB)). — Scale bars = 1 cm.

Citations
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Journal ArticleDOI
01 Feb 2006-Taxon
TL;DR: Investigation of higher level relationships within the Syzygium group, using parsimony and Bayesian analyses of cpDNA sequences from the matK and ndhF genes and the rpl 16 intron, generated for a total of 87 species from the SyZombie group and eight outgroup taxa finds four major well supported clades.
Abstract: With as many as 1000 included species, Syzygium s.l. (including Syzygium and segregate genera such as Acmena, Acmenosperma, Cleistocalyx, Piliocalyx, and Waterhousea) comprises one of the major lineages within Myrtaceae, and is an important component of the Old-World tropical rainforest flora. As with other large genera, high species richness, an extensive distribution and relative homogeneity in morphology have hindered attempts to divide Syzygium s.l. Here, we investigate higher level relationships within the group, using parsimony and Bayesian analyses of cpDNA sequences from the matK and ndhF genes and the rpl 16 intron, generated for a total of 87 species from the Syzygium group and eight outgroup taxa. Within the ingroup, four major well supported clades are found, which form a basal polytomy along with S. wesa and monotypic Anetholea. Generally, the molecular data provide little support for traditional divisions of Syzygium s.l., and the recognition of segregate groups such as Acmena, Acmenosperma, Cleistocalyx, Piliocalyx and Waterhousea. While homoplasy amongst morphological characters has misled attempts to divide the group, detailed and critical assessments of placental, ovular and seed morphology may provide novel insights into evolutionary relationships, and are an important future step in the development of a sound higher level taxonomy for Syzygium s.l.

53 citations

Posted ContentDOI
05 Apr 2016
TL;DR: The Syzygium Working Group (SYZWG) presents a meta-modelling framework for estimating the phytochemical properties of the SyzyGium and its applications in agriculture, ecology, and the environment.
Abstract: 1 Recommended citation: SYZWG (2016) Syzygium Working Group 2 Faculty of Science & Natural Resources, Universiti Malaysia Sabah, Jalan UMS, 88400 Kota Kinabalu, Sabah, Malaysia. 3 The Mauritius Herbarium, Agricultural Services, Ministry of Agro-Industry and Food Security; R. E. Vaughan Building, Reduit, Mauritius. 4 Plant Gateway, 5 Talbot Street, Hertford, Hertfordshire, SG13 7BX, UK. 5 School of Biological Sciences, University of Aberdeen, Aberdeen, AB24 3UU, UK. 6 School of Biological Sciences, University of Adelaide, Adelaide, SA 5005, Australia. 7 Ecology & Ecosystem Research, Georg-August-University Gottingen, Untere Karspule 2, 37073 Gottingen, Germany. 8 Naturalis Biodiversity Center, Botany, P.O. Box 9517, 2300 RA, Leiden, The Netherlands. 9 Royal Botanic Gardens, Kew, Richmond, TW9 3AE, UK. 10 Department of Biosciences, University of Mauritius, Reduit, Mauritius. 11 Centre for Ecological Sciences, Indian Institute of Sciences, Bengaluru, 560012, India. 12 Ashoka Trust for Research in Ecology and the Environment (ATREE), Bengaluru, India 13 Department of Biology, Pittsburg State University, Pittsburg, Kansas, U.S.A. 14 College of Forestry, Guangxi University, Nanning, Guangxi 530005, PR China. 15 Royal Botanic Gardens, Mrs Macquaries Road, Sydney, NSW 2000, Australia

32 citations

Journal ArticleDOI
01 Jan 2006-Blumea
TL;DR: Based on morphological and molecular data, it is concluded that Acmena, Acmenosperma, Cleistocalyx, Piliocalyx and Waterhousea should be combined with Syzygium.
Abstract: Based on morphological and molecular data, we have concluded that Acmena, Acmenosperma, Cleistocalyx, Piliocalyx and Waterhousea should be combined with Syzygium The five genera are formally transferred to Syzygium and new names provided for those of their accepted representatives for which names are as yet not available in Syzygium, with the exception of the New Caledonian members of Piliocalyx as these currently are being revised

24 citations


Cites background from "Four new species of Syzygium (Myrta..."

  • ...After studying the morphological variation evident in syzygioid species in the Papuasian–Australian–new Caledonian region, Craven (1998, 2001, 2003) considered that only a single large genus, i.e. Syzygium, should be recognised....

    [...]

Journal ArticleDOI
22 Apr 2005-Blumea
TL;DR: It is concluded that Anetholea is better placed within Syzygium because of the occurrence of epigeal germination and dry fruitedness in the latter genus also and sequence data from nuclear ribosomal DNA and the chloroplast genome support this conclusion.
Abstract: The distinction between Anetholea and Syzygium is lessened due to the occurrence of epigeal germination and dry fruitedness in the latter genus also and it is concluded that Anetholea is better placed within Syzygium. Analysis of sequence data from nuclear ribosomal DNA and the chloroplast genome support this conclusion. The sole species of Anetholea, A. anisata, is transferred to Syzygium and the new combination, S. anisatum (Vickery) Craven & Biffin, made. Syzygium wilsonii subsp. cryptophlebium is reinstated at species rank, necessitating the new combination, S. cryptophlebium (F. Muell.) Craven & Biffin. Two new taxa of Syzygium are described from Australia, S. wilsonii subsp. epigaeum Craven & Biffin and S. maraca Craven & Biffin.

9 citations

Journal ArticleDOI
11 Aug 2010
TL;DR: Five new species of Syzygium are described from the island of New Guinea, four of which are considered Vulnerable, and one of which is considered Endangered, and the conservation status following IUCN guidelines is proposed.
Abstract: . Five new species of Syzygium are described from the island of New Guinea. One species, Syzygium neurophyllum, occurs in Papua Province, Indonesia. Four species, S. purpuricarpum, S. rambutyense, S. ridsdalei, and S. watutense occur in Papua New Guinea. Four of the five species are known only from the type gathering, but each is easily diagnosed from congeneric members of Syzygium occurring on New Guinea. The conservation status following IUCN guidelines is proposed for each new species, four of which are considered Vulnerable, and one of which is considered Endangered.

8 citations

References
More filters
Book
01 Jan 1919
TL;DR: A comprehensive and indispensable reference to the generic and family names of flowering plants and ferns can be found in the 8th edition of the Dictionary of Ferns as discussed by the authors.
Abstract: Willis's Dictionary is a famous publication in the world of botany and horticulture It is a comprehensive and indispensable reference to the generic and family names of flowering plants and ferns The entries attempt to cover all published generic names from 1753 onwards and published family names from 1789, together with a number of supra- and infra-familial taxa where these have not been based on family or generic names The generic names include many variant spellings and inter-generic hybrids Brief characters of subfamilies are usually given The treatment of the families and higher taxa of the Pteridophyta is based on the classification scheme proposed by Pichi-Sermolli The synopses of the Bentham & Hooker and Engler & Prand systems are retained This eighth edition is now published in paperback in order to make it available to a wider readership, not only of professional botanists, but also students and serious amateurs

1,203 citations

01 Jan 1979

286 citations


"Four new species of Syzygium (Myrta..." refers background in this paper

  • ...In publications of world-wide scope, Merrill (1950) accepted seven (and tentatively an eighth) genera that belong to the Syzygium complex, Airy Shaw (1966) three, Kausel (1957) ten, and Briggs & Johnson (1979) eight....

    [...]

Journal ArticleDOI
TL;DR: Two separate fleshy-fruited clades, the Acmena and Myrtoid groups, are identified, as are the Eucalypt and Tristania groups, and Psiloxylon and Heteropyxis are the first lineages to diverge in both analyses.
Abstract: Cladistic analyses are presented of matK sequence data as well as a nonmolecular database for an identical set of exemplar species chosen to represent the core genera or groups of genera in Myrtaceae. Eleven robust clades are recognized on the molecular data. Polyphyly of the previously recognized Metrosideros and Leptospermum alliances is confirmed, and several smaller informal taxonomic groupings are recognized from among the members of the former alliance, i.e., the Tristania, Tristaniopsis, Metrosideros, and Lophostemon groups. The nonmolecular analysis provides only limited resolution of relationships. A degree of congruence exists between the two analyses in that two separate fleshy-fruited clades, the Acmena and Myrtoid groups, are identified, as are the Eucalypt and Tristania groups, and Psiloxylon and Heteropyxis are the first lineages to diverge in both analyses. A combined analysis recognized all 11 clades that received strong support from the molecular data. A high level of homoplasy is revealed in many of the nonmolecular characters when they are examined against the combined estimate of phylogeny.

233 citations


"Four new species of Syzygium (Myrta..." refers background in this paper

  • ...Burret, Pilidiostigma Burret, Rhodamnia Jack, Calyptranthes Sw., Eugenia and Xanthomyrtus Diels, Wilson et al., 2001)....

    [...]

Journal ArticleDOI
TL;DR: Floral anatomy now provides additional, strong evidence confirming the distinctness of the mainly New World Eugenia s.
Abstract: A B S T R A C T Floral anatomy now provides additional, strong evidence confirming the distinctness of the mainly New World Eugenia s. s. and the strictly Old World Syzygium s. 1. Most significantly, species of Eugenia s. s. have a transeptal vascular supply to the ovules whereas those of Syzygium s. 1. have an axile one. Other features of floral histology and vasculature also support such a division. In addition, a review of the taxonomic literature revealed three hitherto neglected organographic criteria-nature of bracteoles, presence or absence of pubescence, and presence or absence of pseudopedicels-that sharply distinguish between Eugenia s. s. and Syzygium s. 1. An ensemble of these and other organographic criteria further demonstrates the basic disparity of these taxa. The organography and histology of flowers of Eugenia s. 1. are described in detail, with 26 characters contrasting the Old and New World species included in a table.

103 citations


Additional excerpts

  • ...Schmid’s studies of morphological and anatomical data indicated that two separate lineages were involved (Schmid, 1972a, b) and this view has been supported by some recent molecular work....

    [...]

DOI
01 Jan 1983
TL;DR: This study shows that [I ve genera are represented in Australia], including Acmena, Syzygium, Acmenosperma, Eugenia and Waterhousea gen.
Abstract: This study shows that [I ve genera are represented in Australia. Acmena: seven species and one subspecies, including two new species, A. mackinnoniana and A. resa, and one new subspecies, A. hemilampra subsp. orophila. Acmenosperma: two species, including one new species, A. pringlei. Eugenia: one species. Syzygium: 52 species and three subspecies, including 16 new species, S. alatoramulum, S. alliiligneum, S. argyropedicum, S. bamagense, S. boonjee, S. canicortex, S. dansiei, S. endophloium, S. macilwraithianum, S. papyraceum, S. pseudoJastigiatum, S. rubrimolle, S. sharonae, S. velae, S. wesa, S. xerampelinum, and one new subspecies, S. forte subsp. potamophilum; 23 new combinations also made. Waterhousea gen. nov.: four species, including two new species, W. mulgraveana and W. unipunctata; two new combinations also made.

49 citations


"Four new species of Syzygium (Myrta..." refers background in this paper

  • ...Hyland accepted five genera within the complex, Acmena DC., Acmenosperma Kausel, Eugenia L., Syzygium, and Waterhousea B. Hyland, treating the Australian species at that time referred to Cleistocalyx Blume within his concept of Syzygium (Hyland, 1983)....

    [...]

  • ...In a study of Australian species, Hyland (1983) recognised Acmena, Acmenosperma, Syzygium (incl....

    [...]

  • ...Syzygium monospermum appears closely related to S. erythrocalyx (C.T. White) B. Hyland and the specimens then available to Hyland were cited under the latter species in his revision (Hyland, 1983)....

    [...]

  • ...INTrODUCTION The Australian species of Syzygium Gaertn. and its closer generic relatives were recently revised by Hyland (1983)....

    [...]

Frequently Asked Questions (1)
Q1. What are the contributions in this paper?

Schmid ’ s studies of morphological and anatomical data indicated that two separate lineages were involved ( Schmid, 1972a, b ) and this view has been supported by some recent molecular work.