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Phytotaxa 646 (3): 293–300
https://www.mapress.com/pt/
Copyright © 2024 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Peter J. de Lange: 15 Apr. 2024; published: 6 May 2024
https://doi.org/10.11646/phytotaxa.646.3.5
293
Two new species of Lomandra (Asparagaceae: Lomandroideae) from north
Queensland, Australia
JIAN WANG1,* & BRUCE GRAY2
1Queensland Herbarium & Biodiversity Science, Department of Environment, Science and Innovation, Brisbane Botanic Gardens, Mt
Coot-tha Road, Toowong, Queensland 4066, Australia
�
jian.wang@des.qld.gov.au; https://orcid.org/0000-0003-1558-1189
2Australian Tropical Herbarium, James Cook University, Cairns Campus, McGregor Road, Smithfield, Queensland 4878, Australia
https://orcid.org/0009-0002-5412-9320
*Corresponding author:
�
jian.wang@des.qld.gov.au
Abstract
Lomandra scabrifolia Jian Wang ter & B.Gray, sp. nov. and Lomandra tenuis Jian Wang ter & B.Gray, sp. nov. from north
Queensland, Australia are described, illustrated and compared to the morphologically and phylogenetically closely related
species. Notes are provided on their distributions including a map, habitat, phenology and affinities. Their conservation
status is also assessed.
Key words: flora of Australia, flora of Queensland, taxonomy, new species
Introduction
The genus Lomandra Labill. (Asparagaceae: Lomandroideae) includes 64 species to date (IPNI 2023; POWO 2023).
There are 62 species endemic to Australia, two occurring in Australia and New Guinea and one of the two extending to
New Caledonia (Lee & Macfarlane 1986; Wang 2023a, b). Lomandra has had various family placements over the past
decades, including Xanthorrhoeaceae in ‘Flora of Australia’ (Lee 1966; Lee & Macfarlane 1986), Dasypogonaceae
(Briggs 1986; Chase et al. 1995) and Laxmanniaceae (Chase & Stevens 1998; APG II 2003; Wang 2023c). The genus
is more often placed in a broadly circumscribed Asparagaceae under subfamily Lomandroideae Thorne & Reveal
(APG III 2009; Chase et al. 2009; APG IV 2016).
Lomandra was revised by Lee & Macfarlane (1986) for ‘Flora of Australia’, and they recognised 15 species and
four subspecies from Queensland. Currently, there are 25 recognised species in Queensland, as well as three non-
autonymic subspecies (Wang 2023a, b). A taxonomic review of Lomandra in Queensland is underway.
In recent research on Lomandra molecular phylogeny by Gunn et al. (2024), three entities referred to as L. sp.
Bamaga (BG 9947), L. sp. Stannary Hills (BG 9946, erred Lomandra sp. Stannary Hill BG 9246 in their publication),
and L. sp. Watsonville (BG 9948), are related to each other. According to their phylogenetic tree, L. sp. Bamaga (BG
9947) is related with both L. sp. Stannary Hills (BG 9946) and L. sp. Watsonville (BG 9948). The latter two have an
even closer relationship (Gunn et al. 2024). While the first entity L. sp. Bamaga (BG 9947) was formally described
and named as Lomandra grayi Jian Wang ter in March 2023 (Wang 2023a), the other two entities require further
examination. Based on detailed morphological characteristics, we confirmed that L. sp. Stannary Hills (BG 9946) and
L. sp. Watsonville (BG 9948) are distinct species and are described here.
Materials and methods
This study is based on morphological examination of Lomandra herbarium material at BRI and specimens received
on loan from CANB, CNS, DNA, JCT, MEL, NSW and NT. All measurements are based on both dried materials,
i.e. vegetative parts and inflorescence of non-floral parts, and spirit material, i.e. floral parts. For dried material, the
dimensions of flowers are based on material reconstituted with boiling water.
WANG & GRAY
294 • Phytotaxa 646 (3) © 2024 Magnolia Press
Common abbreviations in the specimen citations are: E (East), N (North), S (South), W (West), Ck (Creek), Mt
(Mount), Rd (Road), St (Street), CR (Conservation Reserve), FR (Forest Reserve), SF (State Forest) and NP (National
Park).
Taxonomy
Lomandra scabrifolia Jian Wang ter & B.Gray, sp. nov.
Type:—AUSTRALIA. Queensland: North Kennedy District, 3.8km from Silver Valley Rd towards Irvinebank, 17.4867°S, 145.2583°E,
920 m, 5 February 2023, B. Gray 10093 (holotype BRI!, isotypes CANB!, CNS!, K!).
Lomandra sp. Stannary Hills (BG 9946) (Gunn et al. 2024).
Diagnosis:—Lomandra scabrifolia resembles L. grayi Jian Wang ter, but differs in having bluish green, glaucous
and scabrous leaves; the leaf apex usually rounded to obtuse without teeth; the significantly shorter male and female
inflorescences, and male flower pedicels; the fewer male flowers, and the shorter female pedicel (Table 1).
Description:—Plants are tussocks with condensed ascending rhizomes, each tussock comprising up to 20 or more
tufts (stems). Each tuft is 3–7 mm in diameter at the base. Leaves firm and upright, arranged distichously or irregularly.
Leaf sheath margins at first membranous or cartilaginous, fraying into strips or fibres, 1.5–4 cm long, straw-coloured,
reddish brown or dark brown. Leaf blades bluish green and glaucous, scabrid, usually strongly involuted, flatten out
towards the tips; with (6–)8–16(–20) parallel veins on each surface; the margins coarsely serrulate; leaf apex rounded
to acute or obtuse without teeth. Leaves of male and female plants similar, 20–60 cm long, 1.1–2(–3) mm wide. Male
and female inflorescences a raceme. Male inflorescence 1–3(–5) per tuft, each 20–30(–45) mm long; scape flattened,
irregularly channelled and smooth, 10–20 mm long, 0.8–1.2 mm wide, white; rachis flattened, irregularly channelled,
10–15(–25) mm long, 0.5–1 mm wide, verrucate, same colour as scape; basal bracts between scape and rachis up to
10, long- to short-deltoid, 4–6 mm long and 0.5–2 mm wide at the base that is usually the widest point; membranous,
outer bracts with 2–3 veins. Male flowers 6–30(–40), solitary and alternating or rarely appearing paired or whorled;
each flower with a cucullate bract, 0.8–2.5(–4) mm long and 0.2–1 mm wide, membranous, usually half encircling the
pedicel. Flower pedicels terete, flattened or angular, 0.2–1.1 mm long, 0.15–0.2 mm wide, white to light brown. Male
flowers usually of similar age within each raceme. Flower buds globular, white to brown. Male perianth segments
6 with distinct outer and inner whorls; outer tepals (sepals) 3, cucullate, free except at the very base, uniform in
size, texture thick with slightly thinner margins, 1.4–1.8 mm long, 0.7–0.8 mm wide, greenish maroon; inner tepals
(petals) 3, triangular, free except at the base, uniform in size, texture thick with slightly thinner margins, 1.3–1.8 mm
long, 1.2–1.4 mm wide, cream except for whitish on the margins and inner surface. Stamens 6, all adnate basally to a
swollen centre surrounded by the inner tepals, 3 alternating with inner tepals, slightly higher position and 3 alternating
with outer tepals, slightly lower position; filaments not obvious, 0.1–0.2 mm long, c. 0.1 mm diameter; anthers all
similar, versatile, 0.3–0.35 mm long, 0.2–0.25 mm wide, bright yellow. Pistillode in the swollen centre usually
missing. Female inflorescence 1(–3) per tuft, 10–25(–40) mm long, usually completely hidden at leaf base; scape
flattened, slightly ribbed, creamy white, 5–25 mm long, 1.6–2 mm broad; the rachis flattened or irregularly angled
or channelled, smooth, 5–13 mm long, 0.5–1.3 mm wide. Female flowers (5–)10–25, solitary, alternate, overlapping,
rarely appearing paired; each flower has a bract with largest at base and smallest upwards along the rachis; bracts
membranous, cucullate, 0.5–2.5 mm long, 0.5–1 mm wide at the base widest point, pale brown, partially (1/3–1/2)
encircling the pedicel. Flower pedicels 0.3–0.4 mm long, 0.2–0.3 mm wide, terete or irregularly angled and channelled,
creamy to pale brown. Female flowers of similar age within each inflorescence. Flower buds globular, becoming cup-
shaped with age, yellowish brown. Female perianth segments 6 with distinct outer and inner whorls, yellowish brown;
outer 3 tepals (sepals) ovate, 1.5–2 mm long, 1.1–1.5 wide, adnate at the base; inner 3 tepals (petals) broadly ovate,
2.3–2.5 mm long, 1.8–2 mm wide, connate near base. Staminodes inconspicuous or sometimes absent, when present
lacking filaments and bearing vestigial anthers, 3 inserted on the middle part of inner tepals, 3 alternating with them on
the middle of lower part of each inner tepal. Pistil conspicuous, styles c. 0.2 mm long by 0.2 mm wide, stigmatic lobes
3, c. 0.2 mm long; ovary sessile, globular, c. 1 mm diameter, with 3 locules; ovules 1 per loculus. Fruiting pedicels 1–4
mm long, 0.4–0.5 mm wide. Fruiting styles 0.2–0.3 mm long, persistent. Capsules obovoid, 6–7 mm long, 5–6 mm
diameter, brown to dark brown usually with 4–5 transverse stripes when fresh; carpel brown to dark brown, pale brown
inside; the carpel margins slightly ridged; fruiting perianth segments 6, leathery, persistent, each 2.5–3.5 mm long,
TWO NEW SPECIES OF LOMANDRA FROM AUSTRALIA Phytotaxa 646 (3) © 2024 Magnolia Press • 295
2–3 mm wide, bracts usually persistent. Seeds 1 per locule, but 1 or 2 developed per fruit, narrowly ovoid to ovoid,
6–6.5 mm long, 2.2–3.2 mm wide, usually 2-angled on inner face, rounded on outer face, rough or slightly wrinkled,
translucent in appearance, whitish brown. (Figure 1)
FIGURE 1. Lomandra scabrifolia. A. Tufted plant. B. Leaf tips. C. Male flowers. D. Female flowers. E. Fruit. Photos by B. Gray.
Additional specimens examined:—AUSTRALIA. Queensland: Cook District, Stannary Hills, cultivated at
Hastie Rd, Atherton, 1 December 2017, Gray 9946 (CNS, MEL); Silver Valley to Irvinebank Rd 3.8km from Silver
Valley turn, 15 March 2022, Gray 10030 (BRI); Gray 10034 (BRI); 1.5km E of Silver Valley Rd, 12 September 2022,
Gray 10078 (BRI spirit only); 3.8km from Silver Valley Rd towards Irvinebank, 5 February 2023, Gray 10092 (BRI,
CNS); Gray 10094 (BRI); 1km SW of Watsonville, 15 March 2023, Gray 10111 (BRI including spirit, CNS, NSW);
1km SW of Watsonville, 15 March 2023, Gray 10112 (BRI, CNS).
Distribution and habitat:—Lomandra scabrifolia is endemic to northeast Queensland, occurring west and south
west of Herberton (Figure 3), where it is relatively common. Occurring on rocky hillsides among Themeda triandra
Forssk. and other grasses, along with Xanthorrhoea johnsonii A.T.Lee and sedges, under sparse eucalypt forest
consisting of Eucalyptus cloeziana F.Muell., E. crebra F.Muell., E. portuensis K.D.Hill, Corymbia citriodora (Hook.)
K.D.Hill & L.A.S.Johnson subsp. citriodora and C. rhodops (D.J.Carr & S.G.M.Carr) K.D.Hill & L.A.S.Johnson.
Phenology:—Male flowering was recorded from December to February. Female flowering, fruiting and mature
fruits were collected in March.
WANG & GRAY
296 • Phytotaxa 646 (3) © 2024 Magnolia Press
Discussion:—Lomandra scabrifolia belongs to the L. filiformis group of eastern Australia. It is putatively related
to L. grayi and L. tenuis (Gunn et al. 2024). The three species have similar-sized male and female flowers, fruits and
seeds.
Lomandra scabrifolia can be easily distinguished from L. grayi and L. tenuis by the leaves being bluish green,
glaucous and scabrous (green to yellowish green and smooth for both L. grayi and L. tenuis); Leaf apex, rounded to
obtuse for L. scabrifolia (minutely toothed for L. grayi, acuminate for L. tenuis); Inflorescences, male 2–3 cm long,
female 1–2.5 cm long for L. scabrifolia, ( male 20–55 cm long, female 14–30 long for L. grayi, male 12–27 cm long,
female l0–21cm long for L. tenuis). Pedicel, male 0.2–1.1 mm long and female 0.3–0.4 mm long for L. scabrifolia
(male 2–7 mm long and female 0.5–0.6 mm long for L. grayi, male 1.5–4 mm and female 1–2 mm long for L. tenuis).
(Table 1).
TABLE 1. Diagnostic characteristics of three morphologically and phylogenetically related Lomandra species.
L. grayi L. scabrifolia L. tenuis
Leaf surfaces Glabrous Scabrous Glabrous
Leaf colour Green to yellowish green Bluish green and glaucous Green to yellowish green
Leaf width 3–5 mm 1.1–2 mm 0.5–0.8 mm
Leaf apex Obtuse, near rounded, minutely
toothed Rounded to obtuse without teeth Acuminate
Male and female
inflorescences Dissimilar Similar Similar
Male inflorescences 20–55 cm long, unbranched or
with a few branches 2–3 cm long, unbranched (raceme) 12–27 cm long, branched
Male flower number 40–80 6–30 10–30
Male flower length 1.5–2 mm long 1.3–1.8 mm long 1.5–1.8 mm long
Male pedicels 3–4 mm long 0.2–1.1 mm long 1.5–4 mm long
Female inflorescences 14–30 cm long 1–2.5 cm long 10–21 cm long
Female flower number 10–25 10–25 3–7
Female flower length 1.6–2.8 mm long 1.5–2.5 mm long 1.7–2.6 mm long
Female pedicels 0.5–0.6 mm long 0.3–0.4 mm long 1–2 mm long
Conservation status:—Lomandra scabrifolia can be a common species where it occurs. Although not recorded
from any national park, it is not known to be at risk. Therefore, it is assessed as Least Concern (LC) using the IUCN
(2019) criteria.
Etymology:—From the Latin scaber, meaning rough; folium, meaning leaf. The epithet is in reference to the leaf
blades which are rough.
Lomandra tenuis Jian Wang ter & B.Gray, sp. nov.
Type:—AUSTRALIA. Queensland: Cook District, 1km SW of Watsonville, 17.3825°S, 145.3072°E, 880 m, 24 February 2023, Gray
10114 (holotype: BRI!, isotypes: K!, MEL!).
Lomandra sp. Watsonville (BG 9948) (Gunn et al. 2024).
Diagnosis:—Lomandra tenuis resembles L. grayi Jian Wang ter, but differs from the significantly narrower leaves; the
acuminate leaf apex; the significantly longer female pedicel; the fewer male and female flowers (Table 1).
Description:—Plants are tussocks with ascending rhizomes and rigid spreading roots, each tussock usually
comprising 1 to 10 tufts (stems). Each tuft is 3–6 mm in diameter at the base with leaves arranged distichously or
irregularly. Leaves firm and upright. Leaf sheath margins at first membranous or cartilaginous, fraying into short
TWO NEW SPECIES OF LOMANDRA FROM AUSTRALIA Phytotaxa 646 (3) © 2024 Magnolia Press • 297
to long strips or fibres 4–8 cm long, white, pale brown, straw-coloured to reddish purple. Leaf blades yellowish
green, glabrous, concavo-convex, flatten out towards the tips; with 3–9 parallel fine veins on each surface; margins
smooth, rarely minutely scabrous; leaf apex usually obtuse to acute without teeth. Leaves of male and female plants are
similar, 240–420 mm long, 0.2–0.7 mm wide. Male and female inflorescences similar in appearance being raceme or
paniculate. Male inflorescence 1 per tuft, usually shorter than longest leaf; 120–270 mm long; scape flattened, smooth
to minutely verruculose, 80–140 mm long, 0.5–0.8 mm wide, white, whitish brown or light purplish; primary rachis
flattened, slightly angled, longitudinally ribbed, usually twisted, smooth to minutely verruculose, 80–120 mm long,
0.4–0.7 mm wide, same colour as scape, bearing 0–5 branches; branches alternately arranged along the rachis, 6–20
mm apart, flattened, angled, or channelled or longitudinally ribbed, usually twisted, smooth to minutely verruculose,
10–55 mm long, 0.2–0.4 mm wide, longest branch usually at base, shorter upwards. Each branch has a single bract,
long- to short-deltoid, 1.5–3 mm long, 0.6–1 mm wide at the widest point, with single mid vein, longest usually at
the basal node of rachis, shorter upwards along rachis, usually half encircling the branch. All flowers usually similar
ages within each inflorescence. Male flowers usually 10–30(–40), solitary, alternate or very rarely appearing paired;
each flower has a bract, cucullate, 0.5–2 mm long, 0.4–1 mm wide, membranous, usually half encircling the pedicel.
Flower pedicels terete, flattened or angular, 1.5–4(–6) mm long, 0.15–0.2 mm wide, white to light greenish. Flower
buds ellipsoid, light green. Male perianth segments 6 with distinct outer and inner whorls; outer tepals (sepals) 3,
cucullate, free except on the very base, uniform in size, thick with slightly thin whitish margins in texture, 1.5–1.8
mm long, 1.1–1.3 mm wide, light green to light yellow; inner tepals (petals) 3, broadly cucullate, free except on the
bottom portion, uniform in size, thick with slightly thin margins in texture, 1.6–1.8 mm long, 1.7–1.9 mm broad,
cream to yellow except for whitish on the margins. Stamens 6, all adnate basally to a swollen centre surrounded by
the inner tepals, 3 slightly higher, alternating with inner tepals and 3 slightly lower, alternating with outer tepals;
filament 0.15–0.2 mm long, c. 0.15 mm diameter; anthers all similar, versatile, 0.4–0.5 mm long, 0.3–0.4 mm wide,
bright yellow. Pistillode in the swollen centre c. 0.15 mm long and 0.15 mm diameter. Female inflorescence 1 per
tuft, shorter than longest leaf; 100–210 mm long; scape flattened, smooth to minutely verruculose, 40–120 mm long,
0.4–0.8 mm wide, white, whitish brown or light purplish; primary rachis flattened, slightly angled, longitudinally
ribbed, usually twisted, smooth to minutely verruculose, 20–55 mm long, 0.4–0.8 mm wide, whitish brown or purplish,
bearing 0–3 branches; branches alternately arranged along the rachis, 6–16 mm apart, flattened, angled, or channelled
or longitudinally ribbed, smooth to minutely verruculose, 6–11 mm long, 0.2–0.4 mm wide, longest branch usually at
base, shorter upwards. Each branch has a single bract, long- to short-deltoid, 1.5–2.5 mm long, 0.6–1 mm wide at the
widest point, with single mid vein, longest usually at the basal node of rachis, shorter upwards along rachis. All flowers
usually similar ages within each inflorescence. Flowers 3–7(–11), solitary, alternate or very rarely appearing paired;
each flower has a bract, cucullate, 1–1.5 mm long, 0.3–0.7 mm wide, membranous, usually half to ¾ encircling the
pedicel. Flower pedicels terete, flattened or angular, 1–2 mm long, 0.15–0.2 mm wide, white to light greenish. Flower
buds ellipsoid, green to greenish brown. Female perianth segments 6 with distinct outer and inner whorls, green to
yellowish green; outer tepals (sepals) 3, cucullate, apex acute, free except on the very base, uniform in size, coriaceous
with strongly raised mid vein, 1.7–2.2 mm long, 1.1–1.5 mm wide; inner tepals (petals) 3, broadly orbicular with a
blunt apical extension, free except on the bottom portion, uniform in size, coriaceous with slightly thin margins in
texture, 2.45–2.6 mm long, 2.3–2.5 mm wide. Staminodes inconspicuous or sometimes absent, when present lacking
filaments and bearing vestigial anthers, 3 inserted on the middle part of inner tepals, 3 alternating with them on the
middle of lower part of each inner tepal. Pistil conspicuous, styles c. 0.2 mm long by 0.2 mm wide, stigmatic lobes 3,
c. 0.2 mm long; ovary usually sessile, triangular in cross section, c. 1 mm long by 1.2 mm diameter, with 3 locules;
ovules 1 per loculus. Fruit pedicels 1–1.5 mm long, 0.3–0.4 mm wide. Fruiting styles c. 0.1 mm long, persistence.
Capsules obovoid, usually 5.5–6.5 mm long, 5–6 mm diameter, brown to dark brown usually with irregular wrinkles;
carpel brown to dark brown, light or pale brown inside; the carpel margins smooth; fruiting perianths, 6, coriaceous,
persistent, bracts usually persistent. Seeds 1 per locule, but 1 or 2 developed, narrowly ovoid to ovoid, c. 3 mm long by
4.5 mm wide, usually flat on inner face, rounded on outer face, rough or slightly wrinkled, translucent in appearance,
whiteish brown. (Figure 2)
Additional specimens examined:—AUSTRALIA. Queensland, Cook District: Watsonville Cemetery, cultivated
at Hasties Rd, Atherton, 1 December 2017, Gray 9948 (CNS); Watsonville Range, 7 February 2023, Gray 10097
(BRI); Road opposite Watsonville Cemetery Rd, 8 February 2023, Gray 10098 (BRI, CANB, CNS, NSW); 1km
SW of Watsonville, 19 February 2023, Gray 10100 (BRI); 1km SW of Watsonville, 20 February 2023, Gray 10102
(BRI).
WANG & GRAY
298 • Phytotaxa 646 (3) © 2024 Magnolia Press
FIGURE 2. Lomandra tenuis. A. Tufted plant. B. Leaf tips. C. Male flower. D. Female flower. E. Fruits. Photos by B. Gray.
Distribution and habitat:—Lomandra tenuis is endemic to northeast Queensland, where it is restricted to the
west and south west of Herberton, and the Watsonville area where it is common (Figure 3). Plants occur scattered in
open forest and woodland dominated by Eucalyptus cloeziana F.Muell., E. portuensis K.D.Hill, E. crebra F.Muell.,
Corymbia citriodora (Hook.) K.D.Hill & L.A.S.Johnson subsp. citriodora, Callitris intratropica R.T.Baker & H.G.Sm.,
and Allocasuarina inophloia (F.Muell. & F.M.Bailey) L.A.S.Johnson with sparse understory of grasses and sedges.
Phenology:—Male flowering was mainly recorded from December to February. Female flowering and fruiting
were recorded in February.
Discussion:—Lomandra tenuis has the narrowest leaves among all known species in the genus. It belongs to the
L. filiformis group of eastern Australia and is closely related to L. scabrifolia and L. grayi (Gunn et al. 2024). However,
it is readily distinguished from the two, by its 0.5–0.8 mm leaf width (3–5 mm wide and 1.1–2 mm wide for L. grayi
and L. scabrifolia respectively), the acuminate leaf apex (Obtuse, near rounded with minute teeth or without teeth for
L. grayi and L. scabrifolia); Fewer (3–7) female flowers (10–25 for L. grayi and L. scabrifolia) and longer (1–2 mm
long) female pedicel (0.3–0.6 mm long for L. grayi and L. scabrifolia) (Table 1).
TWO NEW SPECIES OF LOMANDRA FROM AUSTRALIA Phytotaxa 646 (3) © 2024 Magnolia Press • 299
FIGURE 3. Distribution map of Lomandra scabrifolia and L. tenuis.
Conservation status:—Lomandra tenuis can be a common species where it occurs. Although it is not recorded
from a national park, it is not known to be at risk. Therefore, the category Least Concern (LC) is recommended using
the IUCN (2019) criteria.
Etymology:—From the Latin tenuis, meaning slender. The epithet is in reference to the leaf blades which are the
narrowest known in the genus.
Key to the Lomandra species of series Sparsiflorae in north Queensland
1. Leaves 3–5 mm wide. Male and female inflorescences dissimilar ......................................................................................... L. grayi
- Leaves 0.5–3 mm wide. Male and female inflorescences similar ......................................................................................................2
2. Leaf surface scabrous; leaf apex rounded to obtuse without teeth ................................................................................. L. scabrifolia
- Leaf surface glabrous; leaf apex acuminate or minutely 1–3-pointed ...............................................................................................3
3. Leaf apex minutely 1–3-pointed. Female inflorescences 2–9 mm long; 10–20 flowers .................................................. L. filiformis
- Leaf apex acuminate. Female inflorescence 10–21 cm long; 3–7 flowers .............................................................................L. tenuis
Acknowledgements
We are grateful to Mr Luke Hogan who helped in the preparation of distribution map. Mr Tony Bean, Prof. Peter de
Lange, and two anonymous reviewers are thanked for their detailed comments and valuable suggestions. We also wish
to thank the Directors of CANB, CNS, DNA, JCT, MEL, NSW and NT for providing loan specimens.
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