Telopea : Journal of plant systematics

Telopea

Journal of Plant Systematics

Vo I 11(2) • 2006

Botanic Gardens Trust

SYDNEY

National Herbarium of New South Wales

Telopea is published by the National Herbarium of New South Wales, Royal
Botanic Gardens Sydney. It covers the field of systematic botany in Australia and the
Asia-Pacific region, specialising in the flora of New South Wales.

Telopea is published twice yearly.

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Cover Artwork

Telopea speciosissima (Sm.) R. Br., adapted by Helen Stevenson from the original by
David Mackay.

Telopea

Journal of Plant Systematics
1 1 ( 2 ): 99-245 • 2006

CONTENTS

Viola silicestris, a new species in Viola section Erpetion from Australia

Kevin R. Thiele and Suzanne M. Prober 99

Philotheca papillata (Rutaceae), a new endangered species from
north-eastern New South Wales Ian R.H. Telford and Lacldan M. Copeland 105

Pimelea creninophila (Thymelaeaceae), a new species from the New
England Tablelands escarpment of northern New South Wales

Lachlan M. Copeland and Ian R.H. Telford 111

Rediscovery of Prostanthera staurophylla F.Muell. and reinstatement
of P. terctifolia Maiden & Betche (Lainiaceae)

John T. Hunter, John B. Williams and Barry J. Conn 117

New combinations in Australasian Zostera (Zosteraceae)

Surrey W.L. Jacobs Donald H. Lesand Michael L. Moody 127

Two new species of Aponogeton (Aponogetonaceae), and a key to species
from Australia Surrey W.L. Jacobs, Donald H. Les, Michael L. Moody

and C. Barre Hellquist 129

Teucriutn thieleanum (Labiatae), a new species from Victoria, Australia

Barry J. Conn 135

Haloragodendron gibsonii (Haloragaceae), a new species from the

Blue Mountains, New South Wales Peter G. Wilson and Michael L. Moody 141

Lysimachia (Myrsinaceae) in New South Wales P.G. Kodela 147

Three new species of Nymphaea (Nymphaeaceae) in Australia

Surrey W.L. Jacobs and C. Barre Hellquist 155

An anatoino-ecological experiment in Austrostipa aristiglumis, a lowland
Stipoid species. Mirta O. Arriaga and Surrey W.L. Jacobs 161

New taxa and combinations for Oedogonium and Bidbocliaete
(Oedogoniales, Chlorophyceae) in Australia

Stephen Skinner and Timothy J. Entwisle 171

A review of the taxonomy and relationships of the Dendrobium speciosum
complex (Orchidaceae), and recognition of two new taxa

Peter B. Adams, Jacinta M. Burke and Sheryl D. Lawson 195

New combinations in the genus Vacliellia (Fabaceae: Mimosoideae)

from Australia Phillip G. Kodela and Peter G. Wilson 233

Corrigenda — Telopea 11(1) 245

TELOPEA 11(2): 99-245 - 2006

Scientific Editor
Elizabeth Brown
Editor of Publications
Gary Bridle

Typesetting and Production Assistance

Debby McGerty and Julia Sideris

Other members of Editorial Committee

Brett Summerell, Peter Wilson, Joy Everett and Darren Crayn

ISSN 0312-9764

Telopea 11(1) was distributed on 7 September 2005.

National Herbarium of New South Wales
Royal Botanic Gardens Sydney
Mrs Macquaries Rd
Sydney NSW Australia 2000

Telopea 11(2) 99-104

Viola silicestris, a new species in Viola section
Erpetion from Australia

Kevin R. Thiele* and Suzanne M. Prober

Centre for Plant Biodiversity Research, CSIRO, GPO Box 1600, Canberra, ACT 2601
* Author for correspondence

Abstract

Viola silicestris, a distinctive new species found in sandstone and rhyolite habitats in eastern New
Soutli Wales and far south-eastern Queensland, is described. It has previously been confused
with Viola sieberiana and V. hederacea, from which it differs in habit, leaf, flower and habitat
characters.

Introduction

Viola section Erpetion (Sweet) Benth. & Hook.f. comprises c. 12 taxa of endemic
Australian violets, and constitutes the majority of the Australian species of Viola.
Ongoing studies of members of the section are helping to resolve species boundaries
in what has traditionally been regarded as a difficult group, resulting in the description
of a number of new species (see Thiele & Prober, 2003). The present paper describes
Viola silicestris, a distinctive new species from sandstone and rhyolitic habitats on the
coast and ranges of New South Wales and south-eastern Queensland.

Viola silicestris K.R.Thiele & Prober sp. nov.

Affinis Viola sieberiana Spreng. sed foliis plerumque latioribus, basi late cuneata, et
dentibus minoribus aequioribusque differt. Affinis Viola hederacea Labill. sed foliis
latioribus, floribus concoloribus, pallide caeruleo-violaceis, glandibus antherarum
angusto-sinuatis, ct seminibus atro-purpureis differt.

Type: Australia, New South Wales, Carrington Falls, 3 Jan 1998, K.R.Thiele 2568 &
S.M. Prober (Holo: CANB; Iso: MEL, NSW, BRI)

Perennial herb spreading by stolons; rootstock not or scarcely swollen at the stem
bases, with a slender, ± woody taproot. Stems varying from contracted so that the
leaves form rosettes (in exposed plants) to elongate and scrambling to 0.4 m high with
caulescent leaves (in sheltered plants). Leaves broad, the largest (4—)6—10(—15) mm
long, (5—) 10—25(—38) mm wide, (1—)2—4(—5) times wider than long, usually truncate
at base (the basal angle c.l80°), occasionally sub-reniform or ± tapering; lamina with (5-)
7—12(—14) obscure teeth, the apical tooth not prominently larger than the rest, glabrous
or occasionally sparsely short-pubescent when young, green above and beneath; petioles
2-11 cm long; stipules scarious, narrowly triangular with several irregular, elongate,

© 2006 Royal Botanic Gardens and Domain Trust

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Telopea 11(2): 2006

Thiele and Prober

Fig. 1. Viola silicestris. a-c, habit xl. a, caulescent plant from sheltered site; b, non-caulescent
plant from sheltered site; c, non-caulescent, small-leaved plant from exposed site; d, Stipule x5;
e, Flower X4; f, Ventral anther showing gland XlO. (K.R. Thiele 2542, CANB).

Viola silicestris, Section Erpetion

Telopea 11(2): 2006

101

glandular teeth on each side. Flowers prominently displayed, on scapes longer than
the leaves, concolorous pale mauve-blue (rarely somewhat discolorous); anterior petal
(5—)6—7(—8) mm long, 3-4 mm wide, ± rectangular to ovate, usually emarginate, with a
small green blotch at the base then concolorous or with a whitish base, without a white
apex, ± regularly triplinerved with darker veins ; lateral petals spreading, 6-8 mm long,
twisted usually to c. 90° and usually reflexed, concolorous; beard absent or present,
comprising pale hairs; dorsal petals 6-7 mm long, 2-3 mm wide, narrowly obovate,
erect to reflexed, concolorous. Anthers 1.5-2.5 mm long, violet-tinted, the terminal
appendages straw-coloured, with short, irregular hairs on the outer margins of the
anther cells; anther glands pale, slightly shorter than the anther cells, irregular, narrow;
pollen and interior margins of the anther cells pale cream. Ovary and fruit pale green
with or without purple spots or blotches; style distinctly kinked at its insertion on the
ovary. Seeds 1.5-1.7 mm long, glossy purplish-black when mature. Figure 1.

Derivation of name: from the Latin silice (silica), and -estris (inhabiting), in reference
to the characteristic occurrence on siliceous, sandy soils.

Suggested common name: Sandstone Violet

Distribution and habitat: Viola silicestris occurs in scattered localities on the coast and
Great Dividing Range in eastern New South Wales and far south-eastern Queensland.

Fig. 2. Distribution of Viola silicestris

102

Telopea 11(2): 2006

Thiele and Prober

There are three main centres of distribution - from the northern Budawang Ranges
and Jervis Bay to Bundanoon and Robertson, in the Blue Mountains from Bowen
Mountain to the Newnes Plateau, and in northern New South Wales and south-eastern
Queensland from Werrikimbe National Park to Lamington National Park (Fig. 2).

Viola silicestris is typically found in seasonally moist (but often summer-dry) sites in
sclerophyllous, heathy vegetation on acid, sandy soils derived from sandstones or silica-
rich igneous intrusives, particularly rhyolite but possibly also on related rocks such as
porphyries and adamellite. It appears to be absent from better-quality soils derived
from non-sandstone sedimentaries, granites etc. (although it may be found on the
contact zones where the influence of silica-rich parent materials is high).

Conservation Status: Viola silicestris is a relatively common species with a wide
distribution including secure populations in several National Parks. It is not under any
known threat.

Notes: Viola silicestris may be distinguished from all other Australian violets by its
combination of very broad leaves with ± even teeth across the apex, concolorous, pale,
bluish-violet flowers and (in suitable sites) caulescent habit.

Like other species in section Erpetion, leaves of Viola silicestris are highly plastic and
vary greatly depending on their conditions of growth (Fig. 3). Plants growing in
exposed positions, such as on track verges and in other areas with high sunlight, have
reduced leaves which are much smaller and narrower than leaves from plants growing
in more sheltered positions amongst vegetation. It is often possible in the field to find
a complete gradation from small, narrow, reduced leaves in exposed positions to large,
broad, typical leaves in adjacent sheltered sites. Most herbarium specimens, however,
fail to represent such local variation adequately. This is a problem throughout the

Fig. 3. Range of leaf shapes in three species of Viola x 1. a-g, Viola silicestris ; h-m, V. sieberiana
(Sydney population); n-s, V. hederacea s.s. Leaves to the left are from exposed plants with small
leaves, leaves to the right are from sheltered plants with fully expressed leaf shapes.

Viola silicestris, Section Erpetion

Telopea 11(2): 2006

103

section, with reduced leaves of all species converging in shape (e.g. Fig. 3c cf Fig. 3o),
while fully developed leaves from sheltered sites are usually highly distinctive.

Specimens of Viola silicestris with small leaves have often been placed in V. sieberiana ,
while specimens with broad leaves have generally been placed in Vi hederacea sens,
lat ., largely because the broad leaves rule out Vi sieberiana and Vi hederacea has been
incorrectly regarded as highly polymorphic. Specimens with intermediate leaves
(e.g. NSW522386, NSW522398) have sometimes been regarded as hybrids between
Viola hederacea and Vi sieberiana. There is no evidence that such hybrids exist.

Leaves of Viola sieberiana and Vi silicestris differ in that the former are always narrowly
spathulate, even in sheltered sites (see Fig. 2h-m) and have an acute terminal tooth that
is distinctly larger than the lateral teeth, whereas even reduced leaves of Vi silicestris are
generally more or less flat-topped with the central tooth no larger than the lateral teeth.
Leaves of Viola hederacea differ from both species in being ± semicircular, at least when
fully developed (Fig. 3n-s).

Flowers of Viola silicestris and V. sieberiana are similar in shape, size and colour;
together, (lowers of these two species are very different from any other species in section
Erpetion. They are usually concolorous pale bluish-violet, without distinct white tips to
the petals. The anterior petal is narrowly oblong, with a very small green blotch at the
base, a whitish region traversed by pale bluish veins in the lower third or half, grading to
a concolorous pale bluish-violet apex. In contrast, V. hederacea and most other species
(with the exception of the very small-flowered V. fuscoviolacea and Vi cleistogamoides)
have broader anterior petals with a large green blotch at the base, a pale to bright violet
middle section, and a distinct and often sharply contrasting white apex.

Viola silicestris plants often develop a caulescent growth habit when growing in sheltered
sites amongst other vegetation, with elongate, weakly erect, scrambling aerial stems to
30 cm high and scattered leaves. Plants in exposed sites have contracted stems and
fasciculate leaves. Viola hederacea and Vi sieberiana , by contrast, never develop such a
habit, even when growing amongst dense vegetation.

As with other species in the Viola hederacea species complex, it is not unusual to find Vi
silicestris growing allopatrically with other species. It has been found growing more or
less intermingled with Vi hederacea near Robertson, NSW, and with V. sieberiana and
V. fuscoviolacea near Clarence. In such situations, Vi silicestris is usually found in drier
sites on poorer, sandier soils. No hybrids between V. silicestris and any other species
have been seen.

Viola silicestris was included in the key to known Australian species in section Erpetion
(Thiele & Prober 2003) as Viola sp. nov. A. It appears to correspond with Viola hederacea
forma A of James (1990), although specimens at NSW have also been annotated as
formas B, C and D.

Morphologically, Viola silicestris appears to be most closely related to Viola sieberiana.
However, relationships in section Erpetion are largely unknown, and will probably
require DNA sequence data and chromosome counts to elucidate.

Selected specimens examined: New South Wales: Northern Tablelands: About 0.5 km
along Minyon Falls walking track, Whian Whian State Forest, 5 Oct 1987, R. Johnstone
57 (NSW209787); Gibberagunyah Mountain, Whian Whian State Forest, Sep 1967, W.T. Jones
3630 (CANB210906); Peach Mountain, Whian Whian State Forest, 15 miles N of Lismore, 3 Nov

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Telopea 11(2): 2006

Thiele and Prober

1967, K. Grieves s.n. (NSW137263); Cangai Rd, Cangai Boards Flora Reserve, Dec 2000, NSW
NPWS s.n. (NSW444497); Branch of Middle Ck, c. 5 km NE of Glenreagh, Sherwood Nature
Reserve, 13 Mar 1997, P. Gilmour 7722 (NSW434270); Grass Tree Swamp, Werrikimbe National
Park, 7 Nov 1992, T.A. James 1328 (NSW423018); Central Tablelands: Nayook Ck, Deep Pass
east of Glow Worm Tunnel Rd, Newnes Plateau, Wollemi National Park, 13 Jan 1993, P. Hind
6579 (NSW362405); 14.2 km WNW along Culoul Range Rd towards Colo River, 8 Apr 1977,
R. Coveny 9222 (NSW522386); 1.5 km from Bilpin along Mountain Lagoon Rd, 1 Nov 1988,
P. Hind 5703 (NSW212907); c. 3 km W of Bell, 25 Feb 1968, R. Coveny s.n. (NSW 137315); Track
to Tabaraga Rill, c. 2.5 km NW of Bowen Mt, 9 Feb 1988, T.A. James 916 (NSW213497); End
of Lt. Bowen Drive, Bowen Mountain Settlement, 9 Mar 1987, T.A. James 867 (NSW196953);
Southern Tablelands: Butlers Swamp, Tourist Rd, 7 km N of Robertson, 4 Dec 1992, T.A. James
(NSW273260); Bundanoon, 27 Oct 1987, M. O’Neill s.n. (NSW216079); Fitzroy Falls, 30 Nov
1930, PA. Rodway (NSW137314); 3.6 mis. [5.5 km] from The Jumps turnoff toward Tomerong
Nerriga Rd, 12 Nov 1972, M. McMillan 72/87 (CBG50924); 4 km NNE of Mount Tianjara,
2 May 1981, K. Paijmans 3980 (CANB333663); Boolijah Creek, ca 4 km ESE of Sassafras, 15
Oct 1982, LG. Adams & R. Pankhurst 3897 (CANB376712); c. 4 miles SW of Corang Peak,
Budawang Range, 2 Feb 1974, T.G. Hartley14243 (CANB245755); Tianjara Falls, near Nerriga,
1961, Walker ANU1032 (CANB122254); 10 miles [16 km] S of Sassafras, 24 Oct 1965, T. & J.
Whaite s.n. (NSW522398); The Castle, Northern Budawang Range, 31 Jan 1998, l.R. Telford
12381 (CBG9800212); 4.3 km E of Tianjara Falls along Turpentine Road to the north, 27 Oct
1988, S. Donaldson 32 (CBG8803775); South Coast: Currowan State Forest: ca 18 km NNW of
Nelligen, 8 Mar 1973, L.G. Adams 3138 (CANB381723); I iuskisson, Jervis Bay, 13 Sep 1931, PA.
Rodway 507 (NSW137280); 400 m along Stoney Creek Road heading towards Steamers Beach,
29 Sep 1993, M. Kennedy 587 (NSW277654). Queensland: Daves Creek Country, Binna Burra,
23 Nov 1961, R. Jones s.n. (AQ478594); Lamington National Park, c. % mile 11.2 km] towards
Binna Burra from Coomera Falls on the Coomera Track, 15 Jan 1960, R. Scltodde 1153
(AQI15410); Upper Tallebudgera Creek, Dec 1917, C.T. White s.n. (AQ115411).

Acknowledgments

We would like to thank the directors and staff of BRI, CANB, MEL, NSW and AD for
access to their collections, Rob Price and Alan Fairley for assistance with field work,
and Laurie Adams for discussions on Viola and for the Latin diagnosis. Chris Allen is
thanked for his work on the distribution map.

References

James TA (1990) Violaceae. Pp. 435-441 in Harden GJ (ed.) Flora of New South Wales , vol. 1.

(New South Wales University Press: Kensington)

Thiele KR & Prober SM (2003) Two new species and a new hybrid in the Viola hederacea species
complex. Muelleria 18: 7-25.

Manuscript received 22 August 2005, accepted 24 November 2005

Telopea 11(2) 105-110

Philotheca papillata (Rutaceae), a new
endangered species from north-eastern
New South Wales

Ian R.H. Telford and Lachlan M. Copeland

Botany, Centre for Ecology, Evolution and Systematics, The University of New England, Armidale,

NSW, 2351

Abstract

Philotheca papillata I.Telford & L.M.Copel., apparently endemic to Sherwood Nature Reserve,
north-eastern New South Wales, is described as new. Notes on its distribution, habitat and
conservation status are provided. The species is illustrated, its distribution mapped and some of
its attributes compared with related taxa.

Introduction

In 2000, while collecting material of Homoranthus floydii in Sherwood Nature
Reserve near Glenreagh for a revision of that genus, one of us collected material of
an associated species of Philotheca, then thought to be the widespread and variable P.
salsolifolia. Recent revisionary studies in Philotheca sect. Philotheca (Wilson 1998) and
the treatment of Philotheca for Flora of New South Wales (Weston & Harden 2002),
however, present no taxon with the unique combination of attributes of our material.
The species which appear closest morphologically are P. reichenbachii Sieber ex Spreng.,
particularly similar vegetatively, and P. salsolifolia (Snr.) Druce.

Methods

Specimens of Philotheca reichenbachii and P. salsolifolia representing the known range
of the taxa in New South Wales were borrowed from the National Herbarium of New
South Wales (NSW) to augment the collections held in the N.C.W. Beadle Herbarium
(NE). In this study, eight populations of P. reichenbachii and 21 of P. salsolifolia were
scored. For morphological observations and measurements, dried herbarium material
was reconstituted. Leaf characters (direction relative to axis of stem, length, shape
in transvese section, presence of enlarged oil glands and indumentum) and floral
characters (pedicel length, petal length, petal indumentum, petal colour, anther shape
and indumentum of anther apicula) were recorded.

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Telopea 11(2): 2006

Telford and Copeland

Discussion

Philotheca reichenbachii is restricted to the Sydney region (Wilson 1998, Weston &
Harden 2002). Wilson (1998) recognised two subspecies of P. salsolifolia with their
distributions recorded (Wilson 1998, Weston & Harden 2002) as follows. P. salsolifolia
subsp. salsolifolia is widespread in coastal and near-coastal New South Wales from
near Taree south to near Bega, inland to Pilliga and West Wyalong. P. salsolifolia subsp.
pedicellata Paul G.Wilson is restricted to the Angourie district of coastal north-eastern
New South Wales.

The Philotheca population closest geographically to the locality of the new collection
is of P. salsolifolia subsp. pedicellata at Sandon, c. 80 km to the north. This taxon differs
from the Glenreagh plants in its triquetrous, non-verrucose, spreading leaves, long
pedicellate flowers, glabrous keel of the petals and anther shape. P. salsolifolia subsp.
salsolifolia has its closest population at North Haven, c. 180 km to the south. Specimens
from here and other coastal populations differ in almost terete, non-verrucose leaves,
petal indumentum and anther shape (Table 1). Inland populations ol P. salsolifolia
show considerable morphological differences and will not be discussed further here.
This morphological variability and the disjunct distribution pattern of this species
suggest a species complex in need of further investigation. Attributes scored for
P. salsolifolia subsp. salsolifolia in Table 1 are for coastal ‘type’ populations from between
Port Macquarie and Bega.

Comparison of the Glenreagh collections with P. reichenbachii show a remarkable
similarity in vegetative attributes. In floral characters they differ in bearing smaller
flowers with white to pale pink petals, the petal keel pubescent, anther shape and
anther apicula lacking hair tufts (Table 1). It could be postulated that this new species
and P. reichenbachii constitute a vicariant pair, but testing of relationships must await
elucidation of end taxa. The Glenreagh material clearly represents a new species,
described below.

Philotheca papillata I.Telford & L.M.Copel. sp. nov.

P. reichenbachii similis sed petalis minoribus albidis vel subroseis atque pubescentibus
in mediano abaxiali et antheris ad apicem giabris differt.

Type: New South Wales: North Coast: Sherwood Nature Reserve, I.R. Telford 12786,
/./. Bruhl & L.M. Copeland, 14 Sep 2004 (holo: NSW; iso: BRI, CANB, HO, K, MEL,
MO, NE, PERTH). (Specific locality details withheld for conservation purposes)

Shrub, erect, multistemmed, to 60 cm tall, bearing root suckers. Branchlets pilose, pale
green beneath the white indumentum. Leaves incurved, narrow-elliptic, 9-12 mm long,
1-1.5 mm wide, acute, the margins recurved, crenate, verrucose with 4 or 5 glands on
each side of lower surface, both surfaces papillate. Stipules minute, dark purple to black.
Flowers solitary, terminal, on pedicels c. 0.5 mm long. Sepals 5, suborbicular, 1.5-2 mm
long, pubescent. Corolla of 5 free petals, white to pale pink. Petals elliptic, 7-10 mm
long, pubescent on both surfaces, the abaxial surface sparsely verrucose. Stamens 10,
6.5-8 mm long. Filaments fused at base for 4-5 mm. Anthers ovate, apiculate, c. 1.2 mm
long, the apicula glabrous or minutely papillate. Gynoecium of 5 basally-fused carpels,
the carpels tomentose, pale green; style terete, broadening towards the base, c. 4 mm
long, pilose on lower three-quarters; stigma capitate, minutely 5-lobed. Disc obscure.
Fruit not seen. (Fig. 1).

Philotheca papillata (Rutaceae)

Telopea 11(2): 2006

107

Fig. 1. Philotheca papillata a, flowering branch; b, leaf; c, flower; d, androecium; e, gynoecium.
Scale bar: a = 10 mm; b, c = 5 mm; d, e = 2 mm. (all from L.M. Copeland 2605 & P.R. Sherringham).

Additional specimens examined: New South Wales: North Coast: Sherwood Nature Reserve,
L.M. Copeland 2605 & P.R. Sherringham, 2 Sep 2000 (NE); L.M. Copeland 3758, 4 Jul 2004
(CANB, NE, NSW); l.R. Telford 12787, J.J.Bruhl & L.M.Copeland, 14 Sep 2004 (NE). (Specific
locality details of all specimens withheld for conservation purposes).

Distribution: Philotheca papillata is known only from the type locality in Sherwood
Nature Reserve, east of Glenreagh (Fig. 2). In spite of widespread searches in areas
of similar geology and geomorphology (sandstone cliff lines and rocky slopes of the
Grafton Formation and Kangaroo Creek Sandstone) between Chambigne Nature
Reserve, Whitemans Creek, Coaldale and Flaggy Creek Nature Reserve, no further
populations were discovered. The dissected sandstone country between Glenreagh
and Woolgoolga provides habitat for several other narrowly endemic species including
Homorantlnts floydii, Boronia umbellata, B. hapalophylla and an undescribed species of
Lasiopetalum.

Flowering: flowers have only been observed in September although the species is likely
to flower from mid August through to at least early October.

Habitat: this species occurs in a heath community with Banksia oblongifolia,
Leptospermum tr'mervium, Phcbalium wootnbye, Bossiaea rhombifolia, Xanthorrhoca
johnsonii and Philothrix deusta with occasional emergent malleed Eucalyptus
planchoniana. The substrate is a shallow sandy soil over sandstone (Grafton Formation)
along escarpment cliff tops at an altitude of c. 350 m. The site appears to have a high fire
frequency and the root-suckering habit of the species probably assists in maintaining
the population.

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Telopea 11(2): 2006

Telford and Copeland

Table 1. Comparison of some distinguishing attributes between Philotheca papillata,
P. reichenbachii and P. salsolifolia s.s.

Character

P. papillata

P. reichenbachii

P. salsolifolia s.s.

Leaf lamina

incurved

± incurved

± straight

Leaf surface

papillate

hispid, papillate

glabrous-sparsely ciliate

Leaf margin

verrucose

verrucose

smooth

Petal length

7-8 mm

8-13 mm

6-10 mm

Corolla outer surface

keel tomentose

keel ± glabrous

keel glabrous

Corolla colour

white-pale pink

purple

pink-purple

Anther apex

glabrous

with hair tuft

glabrous

Anther shape

ovoidal

cylindroidal

narrow-ellipsoidal

Conservation status: Philotheca papillata is apparently restricted to Sherwood Nature
Reserve where a single population is known of c. 150 individuals along c. 200 m
of escarpment edge. An inappropriate fire regime could present a major threat but
resprouting by root suckers could potentially allow survival. The population is also
close to an informal lookout and a series of obscure walking tracks. Alt hough all known
plants occur within the reserve, the species should still be considered endangered due
to its highly restricted distribution and small population size. Following the criteria of
Briggs and Leigh (1996) a conservation code of 2ECit is recommended.

Etymology: the epithet papillata is from the Latin ( papillatus=bearing papillae), in
reference to the distinctive leaf surfaces.

Fig. 2. Approximate location of Philotheca papillata (denoted by ★) in Sherwood Nature Reserve,
north-eastern New South Wales.

Philotheca papillata (Rutaceae)

Telopea 11(2): 2006

109

Acknowledgments

We would like to thank Jeremy Bruhl for assistance in the field and constructive
comments on the manuscript. Paul Sherringham of the NSW National Parks and
Wildlife Service is also thanked for field assistance and jointly discovering the new
species. Peter Wilson kindly provided the Latin diagnosis.

References

Briggs )D & Leigh JH (1996) Rare or Threatened Australian Plants, revised edition. (CSIRO Pub¬
lishing: Collingwood)

Weston PH & Harden GJ (2002) Philotheca. Pp. 292-299 in Harden GJ (ed.) Flora of New South
Wales, vol. 2, 2nd edition. (New South Wales University Press: Kensington)

Wilson PG (1998) A taxonomic review of the genera Eriostemon and Philotheca (Rutaceae: Bo-
ronieae). Nuytsia 12: 239-265.

Manuscript received 31 May 2005, accepted 24 November 2005

Telopea 11(2) 111-116

Pimelea cremnophila (Thymelaeaceae), a new
species from the New England Tablelands
escarpment of northern New South Wales

Lachlan M. Copeland and Ian R.H. Telford

Botany, Centre for Ecology, Evolution and Systematics, The University of New England,

Artnidale, NSW 2351

Abstract

Pimelea cremnophila L.M.Copel. & I.Telford, a rare new species endemic to the Macleay Gorges
east ofWalcha, is described. Notes are given on its distribution, habitat and conservation status.
Although all plants arc known from a conservation reserve the species is considered to be
endangered due to its restricted distribution and small population size.

Introduction

The gorge country of the eastern escarpment of the New England Tableland is well
known as a ‘hot spot’ for endemism. Narrowly endemic taxa in the Macleay and Guy
Fawkes River gorges include Hakeafraseri, Pltebalium squamulosum subsp. verrucosutn,
Leucopogon trichostylus, Zieria floydii and Bertya ingramii. Other taxa thought to be
endemic to the area include undescribed members of Westringia, Persoottia, Acalypha,
Zieria, Callistemon, Eucalyptus, Leionema and Olearia (Briggs & Leigh 1996; Copeland
1997). Recent fieldwork in Oxley Wild Rivers National Park has yielded a species of
Pimelea which did not fit any currently known taxa in the treatments of Threlfall
(1983), Rye (1990) and Harden (1990). An examination of Pimelea specimens held in
CANB, NSW and NE also supported the recognition of the Macleay Gorges Pimelea as
distinct (herbarium abbreviations follow Holmgren et al. 1990). This paper describes
the new species and gives notes on its distribution, habitat and conservation status.

Pimelea cremnophila L.M.Copel. & I.Telford sp. nov.

P. umhratica similis sed indumenta foliari caulinoque longiore densioreque et filamentis
staminum distinctis differt.

Typ e: New South Wales: Northern Tablelands: Oxley Wild Rivers National Park,
c. 40 km ENE ofWalcha, 30” 55' S, 151" 52' E, L.M. Copeland 3816, I.R. Telford &
P.J. Lupica, 13 Oct 2004 (holo NSW; iso BRI, CANB, CHR, HO, K, MEL, MO, NE).
(Specific locality details withheld for conservation purposes).

Erect shrub to 2.5 m tall. Stems red-brown, hirsute with strigose white antrorse hairs to 3
nun long, glabrescent with age. Leaves opposite, petiolate; petioles c. 1 nun long, densely

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hairy; lamina narrow-elliptic to narrow-ovate, acute, 10-37 mm long, 2.5-6 mm wide,
secondary venation indistinct, the adaxial surface glabrous or sparsely hairy mainly
along the midvein, the abaxial surface paler and hirsute with scattered white, strigose
hairs, the hairs denser and longer, to 2.5 mm long, on the margins. Inflorescences axillary
or terminal, extending up to 15 nodes below shoot apex, 1-4 flowered, condensed
racemes 2-3 mm long; peduncle c. 1 mm long, strigose; bracts leaf-like, c. 2.5-5 mm
long, caducous. Flowers functionally male, bisexual or functionally female; subsessile.
Hypanthium antrorse hairy outside, glabrous inside, greenish-white; sepals 4, narrow-
ovate. Male flowers with hypanthium 6-8 mm long; sepals 3-4 mm long; stamens 2,
rarely 3, inserted near summit of hypanthium; filaments c. 1 mm long; anthers narrow-
oblong, c. 1.7 mm long; pistillode c. 1 mm long. Bisexual flowers protandrous, with
hypanthium 4-6.5 mm long; sepals 3-4 mm long; stamens similar to male flowers;
ovary c. 1.5 mm long, with erect hairs at apex; style eventually exserted; stigma brush¬
like. Female flowers with hypanthium 3-4.5 mm long, circumsdssile c. 1 mm below
sepal attachment; sepals 1.5-2.5 mm long; staminodes minute; gynoecium similar to
bisexual flowers. Fruit dry, ovoid, enclosed in the persistent base of the hypanthium,
pale green. Seed ovoid, 3-3.5 mm long, c. 2 mm wide, with minute longitudinal, foveate
furrows, red-brown (Fig.l).

Additional specimens examined: New South Wales: Northern Tablelands: Oxley Wild
Rivers National Park: c. 40 km ENE of Walcha, 30°55'S, 151°52'E, L.M. Copeland 3444 &
P. lupica , 24 Oct 2002, (BRI, CANB, MEL, NSW, NE); c. 40 km ENE ofWalcha, 30°55'S, 151 °52'E,
L.M. Copeland 3608 & S. Dank, 8 Oct 2003, (CANB, MEL, NE, NSW); rim of gorge of Spring Ck.,
c. 38 km E ofWalcha, 30°55‘ S, 151°51'E, L.M. Copeland 3735, J.J. Bruhl & l.R. Telford, 4 May
2004, (BRI, CANB, NSW, NE); c. 38 km ENE ofWalcha, 30°55'S, 151°51'E, L.M. Copeland 3819,
l.R. Telford & P.J. Lupica, 13 Oct 2004, (AD, NSW, NE); c. 37 km ENE ofWalcha, edge of gorge
of Redmans Ck, 30°56'S, 151°5TE, L.M. Copeland 3822, l.R. Telford & P.J. Lupica, 13 Oct 2004,
(AD, AK, BRI, CANB, CHR, HO, MEL, NSW, NE). (Specific locality details of all specimens
withheld for conservation purposes).

Distribution: apparently confined to gorge rims in the southern part of Oxley Wild
Rivers National Park, approximately 40 km E of Walcha. Several small populations
are scattered along a 5 km stretch of gorge rim. A large area of similar habitat exists
within the park and further searches of this area of gorge rim could potentially yield
additional populations.

Habitat: all plants observed grow in a shallow, skeletal loam over metasediments
on exposed cliff tops or more sheltered cliff-side sites with south-westerly to south¬
easterly aspects. Altitude ranges from 1050-1090 m. Associated species include
Allocasuarina littoralis. Eucalyptus retinens, E. campanulata. Acacia blakei subsp.
diphylla, Maytenus silvestris, Prostanthera rhombea, Dodonaea rhombifolia, Astrotricha
longifolia, Ozothamnus obcordatus, Persoonia media, Callistemon sp. nov., Correa reflexa
var. reflexa, Lepidosperma elatius s.l., L. laterale, Rhodanthe sp. nov. and Notodanthonia
longifolia.

Flowering: flowers have only been observed in early to mid October, but the presence
of unopened floral buds and young fruits on specimens suggests that the species is
likely to flower throughout spring.

Conservation status: the species is currently known from fewer than 100 individuals and
relatively few juveniles have been observed. Potential threats include an inappropriate
fire regime and grazing by feral goats. Several mature individuals appeared to die during

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Fig. 1. Pimelea cremnophila a, flowering branch; b, bisexual flower prior to elongation of style,
with part of hypanthium and one sepal removed; c, bisexual flower; d, functionally female
flower; e, seed. Scale bar: a = 10 mm; b-d = 5 mm; e = 1 mm. (a, b, e, from L.M. Copeland 3608
& S. Doak; c, d, from L.M. Copeland 3816 , ill. Telford & P. Lupica)

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Fig. 2. The distribution of Pimelea cremnophila (denoted by ★) in northern New South Wales.

the period 2002-2003, presumably due to the extreme drought conditions endured
at the time. Although all known plants are reserved in Oxley Wild Rivers National
Park, the species should still be considered endangered due to its highly restricted
distribution, small population size and the potential threats. A ROTAP code of 2ECit is
recommended following the criteria of Briggs & Leigh (1996).

Etymology’: the specific epithet cremnophila is derived from the Greek cremnos (cliff)
and philos (loving), in reference to its habitat.

Comparison with similar species: Pimelea cremnophila belongs to section Epallage
(Endl.) Benth. and appears to be most similar to P. umbratica. It differs from
P. umbratica by its longer, denser indumentum, less prominent secondary leaf venation,
predominantly axillary inflorescences, and smaller anthers on distinct filaments (Table 1).

Sexuality in sect. Epallage requires further study. Threlfall (1983) stated that in P. umbratica,
male, female and bisexual flowers may occur on the same individual. However, Threlfall
included P. leptospermoides under P. umbratica and did not cite which specimens had
been studied for that observation. Rye (1990) claimed P. umbratica has bisexual and
female flowers. We have observed apparently functionally male (with pistillodes) and
functionally female (with staminodes) flowers on the same plant. In P. cremnophila,
plants appear to be polygamous, with functionally male and bisexual flowers on the
same plants and only functionally female on others.

Note that in Harden (1990) the description of P. umbratica follows Threlfall (1983)
with P. leptospermoides (a central Queensland serpentinite endemic) included in
the circumscription. The leaves of P. leptospermoides are alternate while those of
P. umbratica are in fact opposite.

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Table 1. A comparison of the distinguishing features between Pimelea cremnophila and P. umbratica.

Character

P. cremnophila

P. umbratica

Secondary venation

obscure

prominent

Secondary venation angle

25-35°

40-50°

Inflorescence

mostly axillary

mostly terminal

Bisexual flower hypanthium length

4-6.5 mm

5-7.5 mm

Bisexual flower sepal length

3-4 mm

2-2.5 mm

Stamens

filaments present

anthers subsessile

Anther length

1 .7 mm

2-2.5 mm

Seed colour

red-brown

black

Acknowledgments

We are grateful to Sam Doak and Patrick Lupica, rangers from the NSW National Parks
and Wildlife Service, for their assistance in the field. Patrick Lupica is also thanked
for producing the distribution map. Jeremy Bruhl also assisted in the field and gave
constructive comments on the manuscript. We would like to thank Peter Wilson for
assisting with the Latin diagnosis. Directors of CANB, NSW and NE are also thanked
for allowing access to specimens held within their herbaria.

References

Briggs JD & Leigh ]H (1996) Rare or Threatened Australian Plants, revised edition. (CSIRO Pub¬
lishing: Collingwood)

Copeland LM (1997) Rare or Threatened Plants (ROTAPs) occurring in Oxley Wild Rivers Na¬
tional Park. Unpublished report produced for the NSW National Parks and Wildlife Serv¬
ice.

Harden GJ (1990) Thymelaeaceae. Pp. 375-387 in Harden GJ (ed.) Flora of New South Wales,v ol.
1. (New South Wales University Press: Kensington)

Holmgren PK, Holmgren Ni l & Barnett l.C (1990) Index Herbariorum 8th Edition. (New York
Botanical Garden: New York)

Rye BL (1990) Thymelaeaceae. Pp. 122-215 in George AS (ed.) Flora of Australia, vol. 18. (Aus¬
tralian Government Publishing Service: Canberra)

Threlfall S (1983) The genus Pimelea (Thymelaeaceae) in eastern mainland Australia. Brunonia
5: 113-201.

Manuscript Received 31 May 2005, accepted 24 November 2005

Telopea 11(2) 117-126

Rediscovery of Prostanthera staurophylla
F.Muell. and reinstatement of P. teretifolia
Maiden & Betche (Lamiaceae)

John T. Hunter 1 , John B. Williamsf 2 and Barry J. Conn 3

'School of Human & Environmental Studies, The University of New England,
Armidale NSW2351, Australia

2 Botany, Centre for Ecology, Evolution and Systematics, The University of New England,
Armidale NSW2351, Australia

'National Herbarium of New South Wales, Royal Botanic Gardens and Domain Trust,
Mrs Macquaries Road, Sydney NSW 2000, Australia

Abstract

A population of Prostanthera staurophylla was recently rediscovered in the Mt Mackenzie area
near Tenterficld on the Northern Tablelands of New South Wales. A detailed morphological
comparison of these plants with the previously known population occurring in the Torrington
area revealed that the populations were morphologically sufficiently distinct from each other
to be recognised as separate species. Prostanthera teretifolia is reinstated as a distinct species,
separate from P. staurophylla. The former species occurs near Torrington, whereas, P. staurophylla
is restricted to the Mt Mackenzie area near Tenterficld. Descriptions, botanical illustrations and
comments on the conservation status of each species are provided.

Introduction

The staff of the Northern Tablelands Region of the New South Wales National Parks and
Wildlife Service (Department of Environment and Conservation) have been conducting
floristic and vegetation inventories of the reserves within their care since 1997. These
surveys have resulted in extensions of ranges for a number of highly restricted species
and the discovery of many new taxa (Hunter et al. 1996; Hunter 1997; Hunter 1998;
Hunter & Copeland 2001). During a flora survey of Mt Mackenzie Nature Reserve
(by one of us - JTH), specimens were collected of a Prostanthera growing amongst heath
plants within crevices of a single granite outcrop. Although this plant was tentatively
identified as Prostanthera staurophylla (sensu lato — as circumscribed by Conn 1992),
an initial comparison of the morphological features of these plants, with those that
occur in the Torrington area, suggested that they were distinct.

t Deceased 31 July 2005

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The plant collector Charles Stuart originally made collections of Prostanthera
staurophylla from the Tenterfield area (Northern Tablelands region, NSW) while he
was a gardener on a property in that region (Orchard 1999). These collections were
sent to Ferdinand von Mueller who described the taxon in Fragmenta Phytographiae
Australiae (Mueller 1875). However, since Stuart made these original collections,
definitive material conspecific with the type has not been recollected. In 1908,
based on collections by Richard Cambage in the Torrington area, Prostanthera
terctifolia was described (Maiden & Betche 1908). Using the herbarium collections
available and limited field knowledge, Conn (1992) incorrectly assumed that
P. staurophylla represented a precociously flowering juvenile growth stage of
P. terctifolia since the juvenile leaves of this latter species are superficially similar to
the adult leaves of P. staurophylla. Thus, the two taxa were considered synonymous
and combined under P. staurophylla, which had nomenclatural precedence (Conn
1992). However, the rediscovery of populations conforming to the type material of
Prostanthera staurophylla sens. str. has clarified the distinctions between these two
taxa and provided evidence that Prostanthera terctifolia (setisu stricto ), occurring near
Torrington, should not have been reduced to the synonymy of P. staurophylla.

Terminology

The terminology used in this paper follows Conn (1984), with modifications to
inflorescence terminology defined in Conn (1995).

Key to species

An excerpt from the Key to species of Prostanthera (Conn 1992, p. 647, couplet
27) is amended to incorporate the reinstatement of P. terctifolia as distinct from
P. staurophylla.

27 Branches densely covered with short curled hairs; bracteoles not persistent

27a Adult leaves grey-green, strongly revolute such that lamina appears subterete, 5-16 mm
long; margin entire or deeply 2- or 3-fid, densely covered with sessile glands;
branches densely covered with sessile glands; anther appendage absent or minute
.. P. terctifolia

27a*Adult leaves lime- to dark green, oblong or linear, if lobed, then lobes linear, never
terete-like, 2-9 mm long, deeply (2-)3(-7)-fid, with scattered glandular hairs adaxially;
branches densely covered with podiate glandular hairs; anther appendage present

. P. staurophylla

27* Branches glabrous between nodes or sparsely to densely covered with ± straight, spreading
to appressed hairs; bracteoles persistent . [to couplet 28]

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Taxonomy

Prostanthera staurophylla F.Muell., Fragmenta Phytographiae Australiae 9: 73 & 74
(1875).

Lectotype (here designated): New South Wales: Northern Tablelands: ‘12 miles SW
of Tenterfield’, C. Stuart (MEL 43669); probable isolectotypes: ‘New England’, C. St.
(K);‘Nova Anglia’,‘Prostanthera trilobata Fv M ined.’, C. St., 1869 (K); ‘New England’,
C. Stuart (NSW 134503) (refer ‘Typification’, below).

Erect to spreading shrub 1-1.8 m high. Branches ± terete to sub-quadrangular, densely
glandular-pubescent with sub-sessile glands, tubercular hairs throughout, non-
glandular hairs multicellular, curled or crisped, 0.5-1.2 mm long, white. Leaves lime-
to dark green, strongly aromatic (when crushed); petiole distinct, 0.6-1.2 mm long;
lamina elliptic to oblong or more commonly widely trullate or cross-shaped, deeply
(1-) 3 (-7)-lobed, 2-9 mm long, 0.9-7 mm wide, discolorous, abaxial surface paler
and densely covered with curled or crisped non-glandular hairs, particularly along
mid-vein and lateral veins towards lobes where distinct and raised, margin recurved
but never revolute; adaxial surface sparsely glandular when young, less dense towards
petiole, sparsely scabrous particularly on margins and apices, mid-vein depressed.
Flowers axillary, solitary. Podium 0.7-1.5 mm long, sparsely hairy and glandular.
Prophylls persistent, inserted below the calyx base (propodium to anthopodium ratio
3-4), elliptic to narrowly ovate, 0.3-0.4 mm long, 0.15-0.25 mm wide, glandular.
Calyx 3-4 mm long, lime-green, sometimes with a purple-maroon tinge; outer surface
sparsely glandular; inner surface glabrous; margin fimbriate with multicellular white
hairs 0.2-0.3 mm long; abaxial lobe broadly ovate, 1.5-2.5 mm long, 2-2.5 mm wide,
apex rounded; adaxial lobe broadly ovate, 1.2-1.6 mm long, apex obtuse to rounded.
Corolla 15-18 mm long, bluish mauve with darker markings; outer surface glabrous;
inner surface sparsely hairy; tube 8-10 mm long; abaxial median lobes transversely
broad-spathulate, 5-6 mm long, 6-7 mm wide, apex irregular, rounded and bilobed;
lateral lobes elliptic, 3-4 mm long, 3-4 mm wide, apex rounded; adaxial median lobe-
pair very depressed ovate, 1-2 mm long, 9-10 mm wide. Stamens inserted 3-5 mm
above base of corolla; filaments 3-6 mm long; anthers 0.8-1.4 mm long, connective
extended to form a basal appendage 1-1.4 mm long, terminating in narrowly triangular
trichomes. Pistil 7.5-9 mm long (Fig. 1).

Selected specimens: New South Wales: Northern Tablelands: Mt Mackenzie Nature Reserve, SW
of Tenterfield, J.T. Hunter 14377-80 ,12 Mar 2001 (all NSW); loc. cit. J.T. Hunter s.n.,01 Nov 2004
(NSW, spirit); loc. cit. 1.13. Williams, 29 Sep 2004 (NE, NSW).

Notes: the discovery of the Mt Mackenzie population of P. staurophylla is the first
record of this species ( sensu stricto) for 130 years. The affinities of Prostanthera
staurophylla are unclear; however it is expected that it will prove to be closely related to
P. teretifolia from the Torrington area. This species differs from P. teretifolia by its adult
leaves being 3-7-lobed, not terete; the lime- to dark green coloured leaves as opposed
to grey-green; the presence of a distinct anther appendage (lacking or minute in
P. teretifolia); and the habit which is taller in mature specimens and usually develops
into an erect, sometimes leafless, single stem (P. teretifolia is an erect to spreading, multi-
branched from base, leafy shrub). Since the leaves of P. staurophylla are not terete, the
distinctly glandular abaxial leaf surface of this species is visible (abaxial leaf surface
not visible in P. teretifolia). The flowers of P. staurophylla have distinct markings on the
inner surface of the corolla, whereas these markings are absent in P. teretifolia. These
and other useful distinguishing features are summarised in Table 1.

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Fig. 1 . Prostcmthem stciurophylla F.Muell. a, branchlet; b, entire leaf (abaxial surface); c, 3-lobed
leaf (abaxial surface); d, detail of nodal area of branchlet and base of leaves; c, open flower
showing corolla and stamens; f, adaxial (upper) stamen, showing dorsal surface of anther,
connective and appendage; g, adaxial stamen showing ventral surface of anther, and connective
appendage extending from behind; h, abaxial (lower) stamen showing dorsal surface of anther,
connective and appendage; i, flower showing dots on corolla tube and stamens (ventral view);
j, base of calyx, showing prophylls, with one prophyll detached (from J.T. Hunter s.tu, 1
Nov 2004, NSW). Scale bar: a=20 mm, b & c=6 mm, d, f, g, h & j=2.4 mm, e & i=10 mm.
Illustration by Lesley Elkan.

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Etymology: in reference to the cross-shaped leaves, referring to the divided nature of
the leaves which in some cases resemble a cross.

Typification: the protologue of this species states ‘In rupibus graniticis Novae Angliae
inter vicum Tenterfield et flumen Severn; C. St.’ (Mueller 1875, p. 73). The label on the
single collection of P. staurophylla held at MEL (MEL 43669) also refers to ‘Granite
rocks’ and summarises the locality as ‘12 Miles SW of Tenterfield’. Together with the
morphological similarities of these collections to the protologue, this collection is
here selected as the lectotype. The MEL collection does not provide the name of the
collector(s); however, the label is thought to be handwritten by Charles Stuart, with
identification ‘Prostanthera staurophylla EM.’ (at the bottom of the label) added at
some later time by Ferdinand von Mueller.

The two collections of P. staurophylla by Charles Stuart, held at K (as listed above), are
here regarded as probable isolectotypes. Both of these collections refer to the collection
locality as ‘New England’ or ‘Nova Anglia’. The protologue refers to the locality in the
Latin form ‘Novae Angliae’. Both of these collections use the abbreviation ‘C. St.’ for the
collector Charles Stuart. This abbreviation is also used in the protologue. The single
collection of this species by Charles Stuart from ‘New England’, on ‘Phytologic Museum
of Melbourne’ label (handwritten at some later date by Ernst Betche) (NSW 134503) is
also regarded as a probable syntype. A note added by Nerida Ford (ex NSW; in her own
hand) to this latter collection suggests that it is ‘possibly a para-type’. Flowever, since
Mueller did not specifically designate a holotype, this collection should more correctly
be regarded as a probable syntype (according to Article 9.5, Greuter et al. 2000). She
also added that the ‘Type locality was New England, from the vicinity of Tenterfield to
the Severn River’ but this information is merely a reference to the Protologue (Mueller
1875). All three collections morphologically correspond with the description provided
in the protologue. The NSW collection and the two specimens held at K are here
regarded as probable isolectotypes.

Distribution: scattered plants within a single diffuse population of Prostanthera
staurophylla were found at Mt Mackenzie Nature Reserve, Northern Tablelands, New
South Wales (Fig. 3).

Habitat: Prostanthera staurophylla grows in open and exposed situations within
crevices of a granitic outcrop at an elevation of 1200 m above sea level. This species
is a co-dominant with Kunzea opposita F.Muell. and Leptospermum novae-angliae Joy
Thomps., with an understorey dominated by Leucopogon neoanglicus F.Muell. ex Benth.,
Lomandra longifolia Labill., Lomandrafiliformis (Thunb.) Britten, Gahnia aspera (R.Br.)
Spreng., Themeda triandra Forssk. and Brachyscome stuartii Benth.

Conservation status: Prostanthera staurophylla has a very restricted occurrence being
only known from the Mt Mackenzie Nature Reserve. The population size is estimated
to be less than 500 individuals, occurring in an area of about 200 metres long and 80
metres wide. Although fire is unlikely to have occurred in this area for over 30 years;
many seedlings were noted underneath mature stands. This suggests that germination
is occurring naturally without the stimulus of fire, which is typical of plants endemic to
granitic outcrops (Hunter 2003). This species should be considered as endangered and
warrants listing as threatened in the schedules of the New South Wales Threatened Species
Conservation Act 1995 (TSC 1995, TSC 2002, TSC 2004), the Environment Protection
and Biodiversity Conservation Act 1999 (EPBC 1999), and as Critically Endangered
(CR Bla + B2a) in the IUCN Red List ofTheatened Species (Baillie et al. 2004).

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Fig. 2. Prostnnthem teretifolin Maiden & Betche a, branchlet; b, leaf showing adaxial surface;
c, leaf showing abaxial surface; d, detail of nodal area of branchlet and base of leaves; c, open
flower showing corolla, stamens and gyneociunt; f, stamen, showing ventral surface of anthers,
locular appendages and minute connective appendage (visible between locules of anther); g,
stamen showing dorsal surface of anthers, locular appendages and connective; h, fruiting calyx
and persistent style and stigma (lateral view); i, flower bud (lateral view) (from Cawbnge s.n.,
29 Sep 1907, NSW 134430). Scale bar: a=20 mm, b, c, h & i=6 mm, d, f & g=2.4 mm, e=10 mm.
Illustration by Lesley Elkan.

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Prostanthera teretifolia Maiden & Betche, Proc. Linn. Soc. New South Wales 33: 313 &
314 (1908).

Holotype: New South Wales: Northern Tablelands: Torrington, R.H. Cambage s.n.,
29 Sep 1907 (NSW 134430).

Erect to spreading shrub 0.3-1 m high, multi-branched from near base; branches
quadrangular, tending to become sub-terete, very densely glandular with sub-sessile,
semi-hemispherical glands, densely with multicellular curled (eglandular) hairs (hairs
0.1-0.3 mm long) (refer details in ‘Notes’ below). Leaves grey-green, strongly aromatic
(when crushed); petiole distinct or indistinct, 0.6-1.8 mm long; lamina dimorphic;
juvenile lamina, rarely present, entire to deeply 3-lobed, 5-10 mm long, 1-9 mm wide.
Adult lamina linear, subterete (refer details in ‘Notes’ below), entire or 2- or 3-lobed,
5-16 mm long; 0.5-0.9 mm wide, abaxial and adaxial surfaces densely glandular with
sub-sessile, semi-hemispherical glands and with occasional crisped hairs (particularly
on younger leaves). Inflorescence racemiform on leafy branches; uni florescence
monadic with flowers occurring singly in the axils of leaves. Podium 0.6-0.9 mm
long, densely hairy and glandular. Prophylls not seen, soon deciduous, inserted near
midpoint of podium (propodium to anthopodium ratio c. 1.3). Calyx 4-5 mm long,
green, adaxial surface often purplish and/or with a purple-maroon tinge; outer surface
densely glandular with sub-sessile, semi-hemispherical glands, sometimes with a few
scattered hairs; inner surface glabrous; lobes depressed-ovate; abaxial lobe 1.5-2 mm
long, apex obtuse to rounded; adaxial lobe 1-1.2 mm long (shorter than abaxial lobe),
apex almost truncate. Corolla 8-9 mm long, bluish purple, violet or white; outer and
inner surface glabrous; tube 4.5-5 mm long; abaxial median lobes broadly spathulate,
c. 4 mm long, 4-4.4 mm wide, apex slightly irregular, rounded, shortly bilobed; lateral
lobes elliptic, 3-4 mm long, 3-4 mm wide, apex rounded; adaxial median lobe-pair
very depressed ovate, 2.2-3 mm long, 6.5-8 mm wide. Stamens inserted 2.2-2.5 mm
above base of corolla; filaments ! .2-2.5 mm long; anthers 0.8-1.3 mm long, with base
of lobes extended to form a locular appendage 0.2-0.3 mm long; connective extending
to form an indistinct, minute appendage (up to 0.2 mm long) or appendage absent.
Pistil c. 6 mm long (Fig. 2).

Selected specimens: New South Wales: Northern Tablelands: Torrington,/.L. doorman s.n., Jan
1911 (BM, K, NSW); J.B. Williams s.n., 20 Nov 1965 (NSW); J.B. Williams s.n., 29 September
2004 (NE, NSW); c. 1 km N of Torrington township, A.]. Whalen 109 6- R.G. Coveny, 18 Oct
1993 (BR1, MEL, NSW); 4 miles NW of Torrington, H. Wissnum s.n., Apr 1967 (NSW). North
Western Slopes: Eastern Binghi, N of Torrington,/. Benson s.n., 3 Aug 1991 (NSW).

Etymology: in reference to the ‘terete’ (subterete) and linear foliage. The vernacular
name for this species has been once recorded as ‘Turpentine bush’ (J.B. Williams s.n.,
NSW 134377).

Typification: the protologue (Maiden & Betche 1908) cites the habitat as ‘in glabris
saxosis graniticis’. This is in agreement with the notes (in Cambage’s hand) on the
holotype (lower left of herbarium sheet) that describes the plant as growing‘On bare
acid granite rocks’. 1 he handwritten label (lower right) cites the locality as ‘Torrington
n[ea]r Deepwater’; however, this additional information is not supported by the
protologue.

Distribution: found commonly on rocky granite outcrops within the Torrington area
of New South Wales (Fig. 3).

124

Telopea 11(2): 2006

Hunter, Williams and Conn

Fig. 3. Distribution map of Prostanthera staurophylla (A) and P. teretifolia (+), Northern
Tablelands region of New South Wales, Australia

Table 1. Diagnostic characters separating Prostanthera staurophylla from P. teretifolia.

Feature

P. staurophylla

P. teretifolia

Leaves dimorphic

No

Yes

Adult lamina margin

Recurved

Revolute (lamina
subterete)

Adult lamina length

2-9 mm

5-15 mm

Lamina adaxially densely glandular

No

Yes

Adult lamina non-glandular indumentum

Stiff & scabrid

Weak & crisped

Adult lamina abaxial surface visibility

Visible

Obscured

Adult lamina lobes

0 or (2—)3(—7)

0 or 2-3

Leaf colour

Lime- to dark green

Grey-green

Stems branching

Intricate & very densely branched

Loosely branched

Corolla inner surface markings

Present

Absent

Corolla length

15-18 mm long

8-9 mm long

Anther appendage

1-1.4 mm long, with
triangular trichomes

Absent or minute

Prostanthera staurophylla

Telopea 11(2): 2006

125

Habitat: Prostanthera teretifolia is also a species endemic to granite outcrops. It is
found in open and exposed situations within crevices in heathlands and shrublands
from elevations of 800-1200 m above sea level (Fig. 3). It is commonly associated
with Leucopogoit neoanglicus F.Muell. ex Benth., Kunzea bracteolata Maiden & Betche,
Leptospermum novaeangliae |oy Thomps., Leucopogoit melaleucoides A.Cunn. ex
DC., Acacia viscidula Benth., Brachyloma saxicola J.T.Hunter and Micromyrtus sessilis
J.W.Green.

Notes: the curled hairs of the branchlets are especially common at the nodes and in two
narrow zones on the internodes, with each zone on opposite ‘sides’ of the branchlets,
extending from the leaf axil region to the next more distal nodal region between the
opposite leaf bases.

The lamina of the leaves is not truly terete, but rather, the strongly revolute margin
obscures the abaxial lamina surface, including the mid-vein, such that the lamina
appears subterete.

Conservation status: Prostanthera teretifolia is restricted to the Torrington area.
However, it is very common within the granite landscapes and has been described as a
dominant species epitomizing shrublands of granitic outcrop communities of the New
England Batholith (Hunter & Clarke 1998). Thousands of plants exist with many of
these occurring within the Torrington State Conservation Area under the protection
of the New South Wales National Parks Service. Currently the species is listed under
the ROTAP (Briggs 8< Leigh 1996) code of 2RC-. This schedule should be changed
slightly based on current information as the conservation status is considered adequate
(2RCa).

Acknowledgments

The authors wish to thank Alan Hill (Department of Environment 8< Conservation
(N.S.W.), Parks and Wildlife Division) for assistance in the field when the Prostanthera
staurophylla population was re-discovered. Joy Everett (NSW) provided guidance in
regard to the submission of this article. Katy Sommerville and Catherine Gallagher
(both MEL) for kindly providing additional information on the type material of
P. staurophylla as held at MEL. We are grateful to Peter Jobson (NSW) for his advice
on hand-writing on various collections. We thank Lesley Elkan (NSW) for skillfully
illustrating the two species.

References

Baillic JEM, Hilton-Taylor C and Stuart SN (eds) (2004) 2004IUCN Red List of Threatened Spe¬
cies. A Global Species Assessment, (http://www.iucn.org/themes/ssc/red_list_2004/
GSA_book/Red_List_2004_book.pdf) (Downloaded 16 June 2005).

Briggs JD & Leigh JH (1996) Rare or Threatened Australian Plants, revised edition. (CSIRO Pub¬
lishing: Collingwood)

Conn BJ (1984) A Taxonomic Revision of Prostanthera Labill. Section Klanderia (F.v. Muell.)

Benth. (Labiatae). Journal of the Adelaide Botanic Gardens 6(3): 207-356.

Conn BJ (1992) Prostanthera. Pp. 646-662 in Harden GJ (ed.) Flora of New South Wales, vol. 3.
(New South Wales University Press: Sydney)

126

Telopea 11(2): 2006

Hunter, Williams and Conn

Conn BJ (1995) Description of inflorescence axes in the genus Logania R.Br. (Loganiaceae). Kew
Bulletin 50: 777-783.

EPBC (1999) Environment Protection and Biodiversity Conservation Act 1999. (http://www.
deh.gov.au/epbc/index.html)

Greuter W, NcNeill J, Barrie FR, Burdet 1 l-M, Demoulin V, Filgueiras TS, Nicholson DH, Silva
PC, Skog JE, Trehane P,Turland NJ 8c Hawksworth DL (2000) International Code of Botani¬
cal Nomenclature (St Louis Code). Regnutn Vegetabile 138. (Koeltz Scientific Books: Konig-
stein)

Hunter IT (1997) Acacia williamsiana (Fabaceae: Juliflorae): A new granitic outcrop species from
northern New South Wales. Journal of the Royal Society of Western Australia 80: 235-237.

Hunter JT (1998) Two new rare species of Homoranthus (Myrtaceae: Chamelaucieae) from the
Northern Tablelands of New South Wales. Telopea 8: 35-40.

Hunter JT (2003) Persistence on insclbergs: the role of obligate seeders and resprouters. Journal
of Biogeograpliy 30:497-510.

1 lunter JT 8c Clarke PJ (1998) The vegetation of granitic outcrop communities on the New Eng¬
land Batholith of eastern Australia. Cunninghatnia 5:547-618.

Hunter JT 8c Copeland L (2001) Homoranthus binghicnsis (Myrtaceae) a new species from the
North Western Slopes of New South Wales. Telopea 9: 431-433.

Hunter JT, Quinn FC 8c Bruhl JJ (1996) Micromyrtus grandis (Myrtaceae) a new species from
New South Wales. Telopea 7: 77-81.

Maiden H 8c Betche F. (1908) Labiatae. Prostanthera teretifolia , n. sp. Proceedings of the Linnaean
Society of New South Wales 33: 313 Sc 314.

Mueller F. von (1875) Labiatae. Prostanthera staurophylla. Fragmenta Phytographiae Australiae
9: 73 8c 74.

Orchard AE (1999) A History of Systematic Botany in Australia. Pp 11-105 in Orchard AE 8c
Thompson HS (eds) Flora of Australia, vol. 1, 2 nd edition (ABRS/CSIRO Australia: Mel¬
bourne)

TSC (1995) Threatened Sped es Conservation Act 1995 No 101 in ‘In Force Legislation’, (http://
vvww.legislation.nsw.gov.au/maintop/search/inforce)

TSC (2002) Threatened Species Conservation Amendment Act 1995 No 78 in ‘In Force Legisla¬
tion’ (http://www.legislation.nsw.gov.au/maintop/search/inforce)

TSC (2004) Threatened Species Conservation Amendment Act 1995 No 88 in ‘In Force Legisla¬
tion’. (http://www.legislation.nsw.gov.au/maintop/search/inforce)

Manuscript Received 22 June 2005, accepted 24 November 2005

Telopea 11(2) 127-128

New combinations in Australasian Zostera

(Zosteraceae)

Surrey W.L. Jacobs 1 , Donald H. Les 2 and Michael L. Moody 3

'National Herbarium of New South Wales, Mrs Macquaries Road, Sydney, NSW 2000, Australia;
Author for correspondence: surrey.jacobs@rbgsyd.nsw.gov.au
department of Ecology and Evolutionary Biology, University of Connecticut, Storrs,
Connecticut 06269-3043, USA

Indiana University, Department of Biology, Jordan Hall 142, 1001 E. 3rd St., Bloomington,

Indiana 47405, USA

Abstract

The new combinations Zostera muelleri subsp. capricorni (Ascherson) S.W.L.Jacobs,Z. tnuelleri
subsp. mucronata (Hartog) S.W.L.Jacobs and Z. tnuelleri subsp. ttovazelandica (Setchell)
S.W.L.Jacobs are provided for species recently recommended to be treated as a single species.

Introduction

Les et al. (2002) provided ITS, trnK and rbcL sequences of numerous samples of the
family Zosteraceae, concentrating on the Australasian taxa. These were supported by a
cladogram derived from morphological data. All results indicated that:

(i) there was no support for retaining Hetcrozostera as a distinct genus unless subgenus
Zosterella (which contains all of the other Australasian Zostera taxa) was raised to genus
level (Tomlinson & Pozluszny 2001); and

(ii) there was no support for maintaining the species Zostera capricorni, Z. tnuelleri,
Z. mucronata and Z. ttovazelandica and that these could not be reliably and consistently
distinguished from each other. The recommendation was that they be treated as one
species, Z. capricorni (though Z. tnuelleri has priority and is the name that should have
been recommended).

Neither conclusion is novel (Phillips & Menez 1988). Les et al. (2002) present the
most detailed analysis of characters and the most recent review. The results provide
no support for the solution suggested by Tomlinson 8< Pozluszny (2001) and suggest
some sorting of specimens on a geographical basis. While some of these loose groups
have morphological markers, they do not all have useful markers. Those markers that
are discernible are not completely reliable and there is some breakdown between them,
especially where two groups overlap. Nonetheless experienced practitioners can and
do make distinctions most of the time, except for the indicated molecular differences
within Z. tnuelleri s. str. In response to those results (Les et al. 2002) and to reduce

© 2006 Royal Botanic Gardens and Domain Trust

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128

Telopea 1 1(2): 2006

Jacobs, Les and Moodyr

the probability of losing valuable field information, we here recognise the four taxa
as subspecies of Z. muelleri, the earliest available name for the group. Subspecies is
an appropriate level of recognition as the taxa are sorted geographically and there is a
considerable (?uncomfortable) degree of character overlap between them.

Zostera muelleri Irmisch ex Asch., Linnaea 35: 168 (1867)

Type (fide Hartog 1970:90): “Melbourne, at the mouth of the river Yarra, “Australia
Felix” fl., November 1852, F. von Mueller (Type, MEL, P)”.

Zostera muelleri subsp. muelleri

Zostera muelleri subsp. capricorni (Asch.) S.W.L.Jacobs, comb, et stat. now

Basionym: Zostera capricorni Ascherson, Sitzungsber. Ges. Naturf. Freunde Berlin: 11
(1876)

Type (fide Hartog 1970:85): “Moreton Bay, 10-10-1875, Naumann (Type, UC)”.

Zostera muelleri subsp. mucronata (Hartog) S.W.L.Jacobs, comb, et stat. nov.

Basionym: Zostera mucronata Hartog, Verb. Kon. Ned. Akad. Wetensch., Afd. Natuurk.
Tweede Reeks 59 (1): 91 -92 (1970)

Type citation: “Mandurah [W.A., small bay near the bridge], in sandy estuary, 16-8-
1950, G.G. Smith 274 ” (holo L; iso PERTH-U!).

Zostera muelleri subsp. novazelandica (Setch.) S.W.L.Jacobs comb, et stat. nov.
Basionym: Zostera novazelandica Setchell (1933: 816).

Type citation: “Bluff, [lower littoral zone,] N.Z., W.A. Setchell , Mar 11 1927 Herb U Calf
313091” (holo UC).

A discussion of lectotypes is currently in press with Flora of Australia (John Kuo pers.
comm.).

References

Hartog, C den (1970) The sea-grasses of the World. Verhandelingen der Koninklijke Nederlandse
Akademie van Wetenschappen, afdeeling Natuurktmde Tweede Reeks 59, no. 1.

Les DH, Moody ML, Jacobs SWL & Bayer RJ (2002) Systematics of seagrasses (Zosteraceae) in
Australia and New Zealand. Systematic Botany 27: 468-484.

Phillips RC & Menez EG (1988) Seagrasses. Smithsonian Contributions to the Marine Sciences
no. 34.

Setchell WA (1933) A preliminary survey of the species of Zostera. Proceedings of the National
Academy of Sciences Washington 19: 810-817.

Tomlinson PB & Pozluszny U (2001) Generic limits in the seagrass family Zosteraceae. Taxon
50: 429-437.

Submitted 20 September 2005, accepted 06 December 2005

Telopea 11(2) 129-134

Two new species of Aponogeton
(Aponogetonaceae), and a key to species
from Australia

Surrey W.L. Jacobs 1 , Donald H. Les 2 , Michael L. Moody 3
and C. Barre Hellquist 4

’National Herbarium, Royal Botanic Gardens, Mrs Macquaries Rd, Sydney, NSW 2000, Australia
Author for correspondence: surrey.jacobs@rbgsyd.nsw.gov.au
'Department of Ecology and Evolutionary Biology, University of Connecticut,

Storrs, Connecticut 06269-3043, USA

Indiana University, Department of Biology, Jordan Hall 142, 1001 E. 3rd St.,
Bloomington, IN 47405, USA

‘‘Department of Biology, Massachusetts College of Liberal Arts, North Adams,

MA 01247-4100, USA

Abstract

Two new species, Aponogeton cuneatus (removed from A. clongatus) and Aponogeton tofus
(removed from A. euryspermus) are described from Australia. A key is provided for all native and
naturalised Aponogeton species in Australia, and new descriptions are provided for A. clongatus
and A. euryspermus.

Introduction

Hellquist and Jacobs (1998) revised the Australian species of Aponogeton, describing
six new taxa. Even though there had been a substantial increase in the number of
specimens available for examination since Bruggen’s (1969) pioneering work, our
paper stimulated further collecting and further good quality specimens have become
available. While collecting DNA samples for a phylogenetic study of the family (Les
et al. 2005) we observed some material that indicated to us the possibility that there
may be at least one additional new species. The results of the analyses in Les et al.
(2005) clearly demonstrate that there actually are two new species, one (identified
previously as A. elongatus) that molecular data resolved as a distinct sister species to
A. queenslattdicus, and another (identified previously as A. euryspermus) that resolved
as a distinct sister species to the A. euryspermus/A. kimberleyensis clade. Examination
of morphological characters has confirmed their status by demonstrating that these
species are not only well-defined by their degree of molecular divergence but also are
distinctive morphologically. 1 he two new species of Aponogeton are described here,
with modified descriptions of the species in which they were formerly included also
provided, along with a key to all Australian species.

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Jacobs, Les, Moody and Hellquist

Key to native and naturalised species of Aponogeton in Australia

1. Proliferous offshoots formed at tip of peduncle; plants rarely flowering. A. proliferus

1* Proliferous offshoots not formed on plants; plants regularly flowering. 2

2. Inflorescence branched with two spikes. 3

2* Inflorescence unbranched with one spike. 5

3. Flowers white (rarely pink); tepals 1, with 13 or more veins. *A. distachyos

3* Flowers yellow, yellow-green, or green; tepals 2-6, 1 -veined. 4

4. Tepals 6; seeds (2.6-)5.1-6.2(-7.0) mm long; leaves all floating. A. hexatepalus

4* Tepals 2; seeds 2.1—3.2 mm long; leaves all submersed. A. lancesmithii

5. Leaves all submersed, slightly to strongly bullate. 6

5* Leaves submersed and/or floating, not bullate. 7

6. Spathe persistent, to 2.5 cm long; inflorescence to 5.5 cm long; peduncle about same diameter

as inflorescence rachis. A. bullosus

6* Spathe caducous, to 1.5 cm long; inflorescence to 34 cm long; peduncle much broader than
inflorescence rachis. A. lancesmithii

7. Plants mainly producing floating leaves; floating leaves mostly cordate at base; seeds with

obvious double testa. A. queenslandicus

7* Plants mainly producing submersed leaves or submersed and floating leaves; floating leaves
mostly obtuse, some cuneate or truncate at base; seeds with a single testa or, if double, the
second seed coat closely adhering and difficult to detect. 8

8. Submersed leaves narrow, 0.9-3 mm wide; seeds 0.7-1.4 mm long. A. kimberleyensis

8* Submersed leaves broad, 5-60 mm wide; seeds 1.2-4.9 mm long. 9

9. Fruit oval and thick-coated; seed coat closely adhering to embryo and difficult to detect, or

tightly-fitting and very thin; floating leaves rarely formed. 10

9* Fruit elliptic and papery; seeds with a loose outer coat, easily separated from embryo; floating
leaves commonly formed... 11

10. Perianth segments spathulate; south of the Tropic. A. elongatus

10* Perianth segments cuneate; north of the Tropic. A. cuneatus

11. Seeds with a distinct knob at one end, cylindrical to narrowly elliptic, 0.7-1.6 mm wide.
. A. vanbruggenii

ll*Seeds without a knob at one end, broadly elliptic to almost cylindrical, nearly oval,
(1.1—) 1.5—2.8 mm wide. 12

12. Seeds broadly elliptic, 2-2.8 mm wide; Kimberley Region. A. euryspermus

12*Seeds almost cylindrical to narrow elliptical, 1.5-2 mm wide; eastern Northern
Territory. . A. tofns

* = introduced species

Aponogeton cuneatus S.W.L.Jacobs sp. nov.

A. elongato similis sed segmentis perianthii cuneatis, distributione geographica tropica
differt.

Holotype. Queensland: Mowbray River, S of Port Douglas, 16° 33.844'S 145° 22.596'E,
S.W.L. Jacobs 9319 & C.B. Hellquist, 15 Apr 2005 (NSW). Isotypes: BR1, NASC.

Two new species of Aponogeton (Aponogetonaceae)

Telopea 11(2): 2006

131

Submersed aquatic perennial. Tuber ovate to elongate, to 4 cm long, 1.5 cm wide.
Leaves submersed and rarely floating; submersed blades narrow to broadly-elliptic
to elongate, 10-50 cm long, 2-5 cm wide, green or dark green to maroon-green;
margins sometimes slightly undulate; base tapering gradually into petiole, sometimes
asymmetrical; apex blunt-cuneate or obtuse; 5-9-veined; 0—2(—4) rows of lacunae on
each side of the midrib; petiole to 30 cm long. Floating leaves not seen. Peduncle to
c. 90 cm long, 2-4 mm wide at base broadening to 5-6 mm wide at base of inflorescence.
Spathe to 1.5 cm long, caducous. Inflorescence yellow, emergent or floating, single,
tightly-flowered or spaced along the rachis; rachis 1.1-3.1 mm wide. Flowers turned
in all directions. Perianth segments 2, cuneate, 1-veined, yellow, 1-1.4 mm long,
1.4-2 mm wide. Stamens 6; anthers 0.3-0.4 mm long, 0.3-0.6 mm wide; filaments
0.8—1,7(—2.7) mm long, broader at base. Carpels (2—)3(—6). Fruit rounded, thick-
coated, 2.5-5.8 mm long, 1.8-5.0 mm wide; beak mostly lateral, occasionally terminal,
to 1.5 mm long. Seeds 2-5 per fruit, 1.2—3.2(—4.2) mm long, 0.5-1.1 (-1.6) mm wide,
elliptic, with a single testa, closely adhering to embryo or very thin.

Distribution: coastal Queensland north of the tropic.

Selected specimens examined: Queensland: Portland Roads Road between Tozers Gap and the
Middle Claudie Crossing, Clarkson 2923, 25 Feb 1980 (DNA, MEL); Mclvor R., N of Hopevale-
Cooktown rd, Jacobs 8195, Hellquist & Wiersma, 1 Aug 1997 (NSW, BRI, NASC); Stone Creek, nr
Innisfail, Jacobs8571 &Les, 180ct 1999 (NSW,CONN).

Notes: permanent creeks and rivers, often shaded.

This species is named for the cuneate perianth segments characteristic of the species.

A. cuneatiis is superficially similar to A. elongcitus, but presents a gestalt that always
made us suspect that it was a different taxon. No clear distinction could be made
with seed characters so it was retained in A. elongatus. The use of comparative DNA
sequence data (Les et al. 2005) indicated that, despite its superficial resemblance,
A. cuneatiis is quite distantly related to A. elongatus and actually most closely related to
A. queenslandtcus. Subsequently, better spirit collections have allowed us to see that a
conspicuous difference exists in the shape of the perianth segments, with A. cuneatiis
having cuneate tepals (perianth segments) and A. elongatus having spathulate tepals.

Aponogeton tofus S.W.L.Jacobs sp. nov.

Aponogeton elongatus forma strigosus H. Bruggen (1969: 133). syn. nov.

Holotype. Northern Territory: Edith Falls Road, c. 3 miles [5 km] E of Stuart Highway,
L.G. Adams 1658 (L). Isotype NSW.

A. euryspermo et A. vanbruggenii similis,sed ab A. euryspermo seminibus angustioribus
cylindricis differt: ab A. vanbruggenii seminibus extremo etoruloso differt.

Holotype. Northern Territory: Cultivated by Dave Wilson, Howard Springs,
12° 30.982'S 131° 03.239'E, S. Jacobs 9258, 13 Mar 2005, originally from 100 m
downstream (S side) of the crossing of the Bulman to Gove rd over the Goyder River,
13° 01.68'S 134° 58.6'E (NSW).

Rooted, submersed perennial aquatic. Tuber ovate-elongate, 1-3 cm long, 1-2 cm
wide. Leaves mostly submersed, sometimes floating. Submersed blades linear, linear-
lanceolate, ovate, or elliptic, to 23(-35) cm long, (0.5—) 1.4—2.5(—6) cm wide; margins
flat or slightly undulate; base narrowly to broadly cuneate, obtuse or attenuate; apex

132

Telopea 11(2): 2006

Jacobs, Les, Moody and Hellquist

obtuse, broadly obtuse, blunt-cuneate to acute-cuneate, truncate, or rarely refuse;
(3—)7—9(—1 l)-veined; 1-5 rows of lacunae on each side of midvein; petiole to
23(—50) cm long. Floating leaves ovate or elliptic, to 11 cm long, green to maroon-green;
margins flat; base cuneate, obtuse or attenuate; apex obtuse; midrib with 2-4 parallel
veins on both sides; petiole to 64 cm long. Peduncle to 29(-107) cm long, 0.6—1.8 mm
wide at base broadening to (7—)22—48 mm wide at base of infloresence. Spathe to
1.5 cm long, caducous. Inflorescence yellow, emergent or floating, single, rarely
branched, to c. 17 cm long, tightly flowered or spread out along rachis; rachis 0.9-3
mm wide. Flowers turned in all directions. Perianth segments 2, spathulate, 1-veined,
yellow, c. 1.5 mm long, 0.7-1.5 mm wide. Stamens 6; anthers c. 0.2-0.5 mm long,
0.4-0.5 mm wide; filaments 1.2-2.1 mm, broader at base. Carpels (2—)3. Infructescence
moderately dense. Fruit 4-6 mm long, 3.3-4 mm wide, beak to 1 mm long, lateral,
reflexed or straight. Seeds cylindrical, almost ovoid, 2-5 per fruit, usually 1 per small fruit,
3.5-4.5 mm long, (1.1—) 1.5—2 mm wide, thick outer testa easily removed, a second, if
present, tightly adhering and difficult to remove.

Distribution: northern portion of the Northern Territory and Arnhem Land.

Selected specimens examined: Northern Territory: Edith Falls rd, c. 3 miles [5 km] E of Stuart
Hwy, Adams 1658 11 Dec 1966 (NSW);c. 7 km NE Katherine on the Katherine River, Jacobs 3810
3 May 1980 (NSW); Cultivated by L. Smith, Kelso, Qld, jncobs 8268 & Hellquist , 18 Aug 1997,
originally from the Northern Territory (NSW).

The following specimens were identified as A. euryspermus in Hellquist and Jacobs (1998) and
almost certainly belong here, but the specimens have not been reannotated. Arnhem Land,
Mt Gilruth area, 13° 03'S, 133° Ol'E, Henshall 1869 , 6 Jun 1978 (DNA, MEL); lagoon at Jabiru,
12°40'S, 132° 53'E , Craven 2277, 17 Feb 1973 (DNA, MEL); Kakadu Nat. Park, 13°32'S, 132° 29'
Russell-Smith 8559, 7 Jan 1992 (DNA).

Notes: grows in freshwater streams, rivers and lagoons, usually in or near sandstone
country, especially the Arnhem Land Plateau. Flowering and fruiting December to
February and May to July.

This species is named for the sandstone of the Arnhem Land Plateau area where most
of the specimens have been collected. Tofus’ is literally translated as ‘tufa’, which in
modern usage is applied to a relatively porous or spongy rock composed of chemically
or biologically precipitated CaCO,. In older usage, and Latin, ‘tufa’ (or tofus) was used
for a sedimentary rock deposited in layers, a definition including sandstone.

Many specimens of Aponogeton from the tropics originally were assigned to
A. elongatus by Bruggen (1985) and others but can now be assigned to A. euryspermus,
A. vanbruggenii or A. tofus. These taxa initially produce submerged leaves in flowing or
clear water but later in the year plants often develop floating leaves. Species in this group
can be separated by their seed shape and size; A. euryspermus has large broad seeds that
can become very thick when mature, A. tofus has narrower almost cylindrical seeds,
while A. vanbruggenii has even narrower seeds with a distinct knob at one end. All have
seeds with an outer testa that is easily removed (thick in A. euryspermus and A. tofus
- thin in A. vanbruggenii). All species are confined to the tropics with A. euryspermus
more westerly in its distribution, A. vanbruggenii more easterly and A. tofus growing
between the two but overlapping with A. vanbruggenii. DNA sequence data (Les et al.
2005) indicate that indeed, A. tofus is closely related to A. euryspermus (as it had been
previously identified); however, it is distinct and quite divergent at the molecular level
from that and other species.

Two new species of Aponogeton (Aponogetonaceae)

Telopea 11(2): 2006

133

Aponogeton elongatus RMuell. ex Benth., FI. Austr. 7: 188 (1878)

Submersed aquatic perennial. Tuber ovate to elongate, to 4 cm long, 1.5 cm wide.
Leaves submersed and rarely floating (except for subsp. fluitans ); submersed blades
narrow to broadly-elliptic to elongate, 7-34(-42) cm long, 0.8-3.7(-6.7) cm wide,
green or dark green to maroon-green; margins slightly undulate to undulate;
base tapering gradually into petiole, broadly cuneate, obtuse, or rarely truncate,
sometimes slightly asymmetrical; apex blunt-cuneate or obtuse; (5—)7—9(— 1 l)-veined;
0—2(—4 ) rows of lacunae on each side of the midrib; petiole to 17(—35) cm long. Floating
leaves not commonly formed, narrowly elliptic to slightly obovate, 9-19 cm long,

2.2- 3.5 cm wide; base obtuse, broadly cuneate, cordate, or truncate; apex obtuse;
7-9-veined; petiole to 60(-90) cm long. Peduncle to c. 90 cm long, 6-8 mm wide at base
broadening to 5-8 mm wide at base of inflorescence. Spathe to 1.5 cm long, caducous.
Inflorescence yellow, emergent or floating, single, tightly-flowered or spaced along the
rachis; rachis 1.1-3.1 mm wide. Flowers turned in all directions. Perianth segments 2,
spathulate, 1-veined, yellow, 1.8-2.5 mm long, 0.7-1.3 mm wide. Stamens 6; anthers
0.3-0.4 mm long, 0.3-0.6 mm wide; filaments 0.8-1.7(-2.7) mm long, broader at
base. Carpels (2—)3(—6). Fruit rounded, thick-coated, 2.5-5.8 mm long, 1.8-5.0 mm
wide; beak mostly lateral, occasionally terminal, to 1.5 mm long. Seeds 2-5 per fruit,

1.2— 3.2(—4.2) mm long, 0.5-1.1 (-1.6) mm wide, elliptic, with a single testa, closely
adhering to embryo or very thin.

Distribution: coastal Queensland south of the Tropic and coastal northeastern New

South Wales.

1. Plants rarely or never producing floating leaves . 10a. subsp. elongatus

1* Plants commonly producing floating leaves . 10b. subsp. fluitans

Aponogeton euryspennus Hellq. & S.W.L.Jacobs, Telopea 8: 16 (1998)

Rooted, submersed perennial aquatic. Tuber ovate-elongate, to 1-3 cm long, 1-2 cm
wide. Leaves submersed and/or floating. Submersed blades linear, linear-lanceolate,
ovate, or elliptic, to 23(—35) cm long, (0.5-)1.4-2.5(-6) cm wide; margins flat or
slightly undulate; base narrowly to broadly cuneate, obtuse, or attenuate; apex obtuse,
broadly obtuse, blunt-cuneate to acute-cuneate, truncate, or rarely refuse; (3—)7—9
(-ll)-veined; 1-5 rows of lacunae on each side of midvein; petiole to 23(-50) cm
long. Floating leaves ovate or elliptic, to 11 cm long, green to maroon-green; margins
flat; base cuneate, obtuse, or attenuate; apex obtuse; midrib with 2—4 parallel veins on
both sides; petiole to 64 cm long. Peduncle to 29(—107) cm long, 0.6-1.8 mm wide
at base broadening to (7—)22—48 mm wide at base of inflorescence. Spathe to 1.5 cm
long, caducous. Inflorescence yellow, emergent or floating, single, rarely branched, to
c. 17 cm long, tightly flowered or spread out along rachis; rachis 0.9-3 mm wide.
Flowers turned in all directions. Perianth segments 2, spathulate, 1-veined, yellow,
1.1-2 mm long, 0.7-1.5 mm wide. Stamens 6; anthers c. 0.2-0.5 mm long, 0.4-0.5 mm
wide; filaments 1.2-2.1 mm, broader at base. Carpels (2—)3. Infructescence moderately
dense. Fruit 3-6.2 mm long, 2.4—5 mm wide, beak to 1 mm long, terminal or lateral,
reflexed or straight. Seeds broadly elliptic, almost ovoid, 1—3(—4) per fruit, usually 1 per
small fruit, 3.2—4.4 mm long, (1.1—)2—2.8 mm wide, thick outer testa easily removed, a
second, if present, tightly adhering and difficult to remove.

Distribution: the Kimberley Region of Western Australia.

134

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Jacobs, Les, Moody and Hellquist

Acknowledgments

We thank Karen Wilson for the Latin diagnoses, and Dave Wilson for continued
cooperation by collecting and cultivating specimens.

References

Bruggen HWE van (1969) Revision of the genus Aponogeton (Aponogetonaceae): III. The spe¬
cies of Australia. Blutnea 17: 121-137.

Hellquist CB & Jacobs SWL (1998) Aponogetonaceae of Australia with descriptions of six new
taxa. Telopea 8: 7-19.

Les DH, Moody ML & Jacobs SWL (2005) Rhylogeny and systematics of Aponogeton (Aponoge¬
tonaceae): the Australian species. Systematic Botany 30: 503-519.

Submitted 20 September 2005, accepted 07 December 2005

Telopea 11(2) 135-140

Teucrium thieleanum (Labiatae), a new species
from Victoria, Australia

Barry J. Conn

National Herbarium of New South Wales, Mrs Macquaries Road, Sydney NSW 2000 Australia

Abstract

Teucrium thieleanum is an erect subshrub that is moderately to densely glandular hairy, with
simple, multicellular podiate glands, subsessile hemispherical glands, as well as long and short
eglandular hairs, ovate to broadly ovate leaves with crenate to lacerated margin. This paper
provides a full description of Teucrium thieleanum, habitat notes, botanical illustration, and a
key to distinguish this species from other named Teucrium species in Victoria.

Introduction

This species is only known from two localities in East Gippsland, Victoria (Australia).
The earliest collection being by Neville Walsh and David Albrecht, gathered in 1988 from
the Marble Gully area, with later collections by Kevin Thiele, gathered in November 2004
and January 2005, both from Little River Gorge. With the description of this new species,
in this paper, there are now five known species of Teucrium occurring in Victoria.

Terminology follows Conn (1984). Inflorescence terminology follows Briggs and Johnson
(1979), except for modifications by Conn (1995).

Teucrium thieleanum B.J.Conn, sp. nov.

T. corymboso foliis latioribus (24-45 mm), margine crenato vel lacerato, prophyllis
latioribus (0.6-0.8 mm), indumento glandifero longo subsessilique (vice subsessili
modo) differt.

Type: Victoria: East Gippsland: Little River Gorge above 'BBC Camp’, K.R. Thiele 2992,
24 Nov. 2004 (holo NSW 677435; iso MEL 2264890).

Erect subshrub 1-1.5 m high, branches above the base (Thiele 2992). Branches ±
quadrangular, slightly ridged on angles, densely hairy; hairs simple, straight to slightly
curved, multicellular, glandular, ± spreading (patent to slightly antrorse or retrorse);
internodes with short and long hairs, short hairs 0.1-0.2 mm long, translucent throughout
or with gland white to pale golden-coloured, !onghairs0.5-0.9 mm long; nodes with longer
spreading hairs 1-1.5 mm long, moderately glandular with hemispherical, subsessile
glands. Leaves aromatic, with petiole 7-8 mm long; lamina ovate to broadly ovate, less
frequently elliptic, greyish green, 40-50 mm long, 24-45 mm wide, with length 1-1.7
times width, distance from base of maximum width 0.3-0.5 times total lamina length;

© 2006 Royal Botanic Gardens and Domain Trust

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Barry J. Conn

abaxial surface sparsely glandular hairy (average density 8-10 hairs per mm 2 ), hairs ±
patent, c. 0.1 mm long, with longer hairs on veins and margin (hairs 0.2-0.4 mm long),
very densely covered with subsessilc glands (as per branches); adaxial surface moderately
glandular hairy (average density c. 12 hairs per mm 2 ), hairs ± patent, 0.8-1.3 mm long,
densely covered with subsessile glands (as for branches); base decurrent; margin crenate
to lacerate, slightly recurved, with apex of teeth rounded to subacute (when lacerate);
apex subacute; venation distinct, midvein and secondary veins slightly raised on abaxial
surface. Inflorescence frondose-bracteose, many-flowered; uniflorescence dichasial,
(5-)7-15-flowered (7-flowered distally). Podium 2—5.5(—7) mm long, densely hairy (as
for branches), hairs 0.1-0.3 mm long. Prophylls ± persistent, inserted near middle of
podium (propodium 2-3 mm long; anthopodium 1.8-2 mm long, with propodium
1-1.5 times length of anthopodium), opposite to subopposite; lamina narrowly
triangular, 1-1.5 mm long, 0.6-0.8 mm wide, slightly concave lengthwise to almost flat,
densely hairy with short hairs (as for branches). Calyx dull green, campanulate; outer
surface densely hairy with short hairs (as for branches), midveins prominent, ridged,
with slightly longer glandular hairs; tube 1-1.5 mm long, inner surface glabrous; lobes
equal, narrowly triangular, c. 2 mm long, 0.8-1 mm wide at base, with length 2-2.5
times width, inner surface sparsely hairy (hairs as for branches), apex subacute. Corolla
6-7.5 mm long, white; outer surface of lobes moderately hairy, eglandular hairs 0.1-0.2
mm long, translucent to white, simple, multicellular, spreading to antrorse, sparsely
to moderately glandular with subsessile hemispherical glands; inner surface sparsely
hairy in tube, with hairs simple, spreading, c. 0.1 mm long; tube 1.5-2 mm long; lobes
glabrous, or with an occasional eglandular hair, inconspicuously glandular with minute,
subsessile hemispherical glands; abaxial median lobe spathulate, slightly involute-
conduplicate, 3-3.6 mm long, 2-2.5 mm long, with length c. 1.5 times width, apex
rounded; lateral lobes and adaxial median lobe pair slightly elliptic to oblong, 2—3 mm
long, each lobe 1-1.3 mm wide, with length 2-2.3 times width, apex obtuse. Stamens
inserted 0.5-0.7 mm above base of corolla; filaments arching towards adaxial inner
surface of corolla, 6-8 mm long, sparsely hairy near base (as for inner surface of corolla
tube), abaxial filament-pair longer than adaxial filaments; anthers 0.5-0.6 mm long, c. 0.3
mm wide, connective with a line of hemispherical, white glands. Pistil 10-11 mm long;
ovary cylindrical to slightly obovoid, 0.3-0.5 mm long, diameter 0.6-0.7 mm, lobes c. 0.1
mm long, moderately hairy distally, with hairs eglandular, white, ± curled, moderately
glandular (glands subsessile, hemispherical); style c. 9 mm long, strongly curved (towards
adaxial plane); stigma lobes 0.7-1 mm long, slightly unequal, soon becoming ± strongly
reflexed. Fruiting propodium and anthopodium extended to c. 3 mm long. Fruiting calyx
enlarged, 3-4 trim long, mouth 2-2.5 mm diameter (compared to c. 1 mm diameter
in flower), becoming pale brown, densely glandular. Mature mericarps 1.5-2 mm long,
surface light to dark brown, with irregular longitudinal ridges (corrugated), moderately
hairy and glandular (as for ovary). Figure 1.

Other specimens examined: Victoria; East Gippsland: Little River Gorge below ‘Farm Creek’,
K.R. Thiele 2994 , 5 Jan 2005 (MEL n.v., NSW); Marble Gully area, 200 m S of Old Hut Creek,
N.G. Walsh 2089 & LIE. Albrecht, 30 Jun 1988 (AD, MEL, NSW)

Distribution: this species is only known from East Gippsland region of Victoria.

Habitat: this species is recorded as growing in Eucalyptus albens grassy woodland on
loamy soils derived from Snowy River volcanics ( Thiele 2992 & 2994 ) and from the
margin of dense Pomaderris sp. aff. oraria - Helichrysum adnatum shrubland on marble
substrate ( Walsh 2089).

Teucrium thieleanum (Labiatae), a new species

Telopea 11(2): 2006

137

Fig. 1. Teucrium thieleanum B.J.Conn a, branchlet, showing flowers and flower buds; b, detail
of adaxial leaf surface showing eglandular hairs, plus podiate and hemispherical glands; c, detail
of branchlet showing indumentum with long multicellular podiate glands at node between leaf
bases; d, leaf with margin lacerate; e, leaf, showing prominent venation of abaxial surface and
margin crenate; f, flowers, lateral view, showing calyx, corolla, exserted style, stigma and stamens;
g, corolla, ventral view, showing position of stamens and stigma; h, anther, dorsal view, showing
line of hemispherical glands on connective; i, anther, ventral view; j, open corolla, showing hairy
inner surface of tube and androecium, and comparative lengths of stamens; k, gynoecium,
showing indumentum on distal surface of ovary, displaced alignment of style (towards adaxial
inner surface of corolla), and slightly unequal stigma lobes, (a, b, e-k from living material of
Thiele 2994 ; c from herbarium material of Thiele 2994 , d from Thiele 2992). Scale bar: a = 50
mm; b = 2 mm; c = 4 mm; d & e = 40 mm; f & g = 10 mm; h & i = 2.5 mm; j & k = 7.5 mm.

138

Telopea 11(2): 2006

Barry J. Conn

Etymology: the specific epithet thieleanum recognises the significant contribution
that Kevin Thiele has made to systematic botany. He is also the collector of the type
material.

Notes: this species has multi-branched botryoidal superconflorescences, with all
axes anthotelic, consisting of thyrsoid con florescences. Each conflorescence usually
consists of 7-flowered simple dichasia throughout much of its length, whereas
15-flowered dichasia are present towards the base. However, fewer-flowered dichasia are
occasionally present, especially when the plant is beginning to flower. The inflorescence
is leafy throughout much of its length, with a gradual reduction in leaf size from base
to apex. However, the bracts at the distal end of the a, internode of the uniflorescence
are slightly lacerated, like normal leaves. Ultimately, the leaves are reduced such that
they are prophyll-like distally. Teucrium thieleanum is morphologically very similar to
T. corymbosa. The characters that distinguish these two species are summarised in the
identification key (see below).

Conservation status: although the conservation status of this species is unknown, it
has been recorded as locally ‘uncommon’ (Walsh 2089) and ‘infrequent’ ( Thiele 2992
and 2994).

Identification of Teucrium thieleanum from other Victorian species

The identification key provided by Conn (1999, p. 457) to the species of Teucrium
occurring in Victoria is here modified to distinguish this new species from the other
species.

1 .

1 .

2 .

2 .

3.

3.

4.

4.

Leaves entire, or trifoliate with linear entire leaflets; plants greyish from a very dense, short

indumentum . 2

Leaves toothed or deeply lobed almost to midrib; plants green . 3

Leaves trifoliate (or sometimes the lowermost entire) . T. albicaule

Leaves simple throughout . T. racemosum

Leaves entire, deeply lobed; hairs mostly branched . T. sessiliflorum

Leaves toothed, not deeply lobed; hairs simple, never branched . 4

Leaves with margin serrate, lamina 6-20 mm wide; corolla with abaxial median lobe (lip)
6-9 mm long; prophylls narrow (1-1.3 mm long, c. 0.2 mm wide); branches with simple,

short, antrorse eglandular hairs and subsessile hemispherical glands . T. corytnbosutn

Leaves with margin crenate to slightly lacerate, lamina 24-45 mm wide; corolla with abaxial
median lobe (lip) 3-3.6 mm long; prophylls broad (1-1.5 mm long, 0.6-0.8 mm wide);
branches with simple, short and long eglandular hairs, straight to slightly curved, and multi-
celled, podiate glands and subsessile hemispherical glands . T. thieleanum

Acknowledgments

The botanical illustration of Teucrium thieleanum was skillfully drawn by Lesley Elkan
(NSW). Kevin Thiele (CANB) brought this taxon to my attention. Peter Wilson (NSW)
kindly corrected the Latin diagnosis.

Teucrium thieleanum (Labiatae), a new species

Telopea 11(2): 2006

139

References

Briggs BG & Johnson LAS (1979) Evolution in the Myrtaceae - Evidence from infloresence
structure. Proceedings of the Linnean Society of New South Wales 102:157-256.

Conn BJ (1984) A taxonomic revision of Prostanthera Labill. Section Klanderia (F.v.Muell.)

Benth. (Labiatae). Journal of the Adelaide Botanic Gardens 6: 207-348.

Conn BJ (1995) Description of inflorescence axes in the genus Logania R.Br. (Loganiaceae). Kew
Bulletin 50: 777-783.

Conn BJ (1999) Teucrium. Pp. 456-459 in Walsh NG & Entwisle TJ (eds) Flora of Victoria, vol.
4. (Inkata Press: Melbourne)

Manuscript received 28 October 2005, accepted 9 December 2005

Telopea 11(2) 141-146

Haloragodendron gibsonii (Haloragaceae),
a new species from the Blue Mountains,
New South Wales

Peter G. Wilson 1 and Michael L. Moody 2

'National Herbarium of New South Wales, Royal Botanic Gardens, Sydney NSW 2000 Australia
2 Dept. Ecology & Evolutionary Biology, University of Connecticut, 75 N. Eaglcvillc Rd„

Storrs, CT 06268-3043 USA

Present address: Dept. Biology, Indiana University, Jordan Hall 142, 1001 E. 3rd St.,
Bloomington, IN 47405 USA

Abstract

A new species, Haloragodendron gibsonii, is described from the Blue Mountains region of
New South Wales. This taxon has always been recognised as being close to H. lucasii, a species
restricted to the Ku-Ring-gai area of Sydney, but its rank has been uncertain. Relationships
within the genus in eastern Australia have now been assessed using molecular data and these
support recognition of the Blue Mountains plants as a distinct species.

Introduction

In October 1982, Sydney naturalist and bushwalker Colin Gibson collected two small
pieces from a plant in the Bungleboori Creek area of the Blue Mountains National
Park (Gibson 1998). Unfortunately, herbarium staff did not recognise the significance
of this specimen when it was brought in and it was overlooked until late 1986 when it
was identified as Haloragodendron lucasii (Maiden & Betche) Orchard. H. lucasii was
originally named from a collection made in the Sydney suburb of Gordon and had
been presumed extinct until its rediscovery in nearby Barra Brui, also in 1986. The Blue
Mountains specimens showed some differences from those from Sydney, particularly in
the unwinged fruit that sets seed {H. lucasii sens. str. has a winged hypanthium and does
not appear to set seed). Orchard (1990), with only a single Blue Mountains specimen
available for examination, said he was ‘inclined to consider it as merely a variant of H.
lucasii’. The differences between the Blue Mountains and Sydney populations, however,
prompted Wilson (1991) to suggest that the former might constitute a distinct subspecies.
Briggs and Leigh (1988) listed this taxon as ’Haloragodendron sp. 1 (Wollongambe Creek)’
then later (Briggs & Leigh 1996) as ‘ Haloragodendron lucasii subsp. 1 (Wollongambe
Ck.; Hind 5981)’. Recent molecular data (Moody 2005) in combination with the
morphological differences support the recognition of this variant taxon at specific rank
and it is here described as H. gibsonii.

© 2006 Royal Botanic Gardens and Domain Trust

ISSN0312-9764

142

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Wilson and Moody

Materials and methods

In a broader study of Haloragaceae, Moody (2005) collected various species of
Haloragodendron and generated sequences of the nuclear (ITS) and chloroplast (trnK
intron, including matK) regions. Vouchers for the eastern Australian species included
in this study are shown in Table 1. Full details of techniques used, and broader, more
detailed, analyses will be presented elsewhere (Moody, in prep.).

Molecular Results

A phylogenetic hypothesis derived from a parsimony analysis of the combined ITS +
trnK-matK DNA sequence dataset for the eastern Australian species of Haloragodendron
is presented in Fig. 1. The eastern species form a monophyletic group with 100%
bootstrap support and a branch length of 15. There are 4 point mutations in the matK
+ trnK cpDNA region that vary between H. lucasii and the new species. There are 8
point mutations and 1 indel (1 bp) in ITS that vary consistently between H. lucasii and
the new species. There is no variation between the Sydney populations in cpDNA while
some autapomorphies are found between ITS sequences from the two samples derived
from the Barra Brui population (Fig. 1). However, the ITS sequence data derived from
one of the ex-Barra Brui plants (Moody 4S2 : cultivated at Mt Annan) and the St Ives
Chase population (Moody 486) did not vary, so the latter was omitted from the analysis.
The DNA variation between H. lucasii and the new species is comparable to that found
between H. monospermum and H. baeuerletiii (Fig. 1).

Key to species of Haloragodendron in eastern Australia

1. Fruits obovoid to obpyramidal, strongly angled and ribbed, or winged

2. Leaves at least 2.5 cm long; hypanthium and fruit distinctly winged

3. Wings conspicuously veined; anthers 2-4 mm long .. 1 .H. baeuerlenii

3* Wings not conspicuously veined; anthers mostly 4-6 mm long . 2. H. lucasii

2* Leaves usually 1-1.5 cm long; fruit angled and ribbed . 3. H. monospermum

1 * Fruits ovoid, obtusely angled when dry, never strongly ribbed . 4. H. gibsonii

Haloragodendron gibsonii Peter G. Wilson & M. Moody, sp. nov.

H. lucasii affinis sed floribus minoribus (petalis antheribusque brevioribus), fructibus
ovoideis nunquam alatis differt.

Type: New South Wales: Cultivated, Royal Botanic Gardens, Mt Annan, Accession No.
NSW498891 ex Blue Mountains National Park ( Hind 5978 & Cherry ), Peter G. Wilson
1684 , 15 Sep 2004 (holo NSW; iso CANB, K, US distribuendi)

Straggling shrub up to 1 m tall; stems 4-angled to narrowly winged. Leaves subsessile,
lanceolate, 15—25(—28) mm long and 2-5.5(—6) mm wide, serrate; teeth reddish, variable
in size, (4—)7—12 each side. Flowering shoots (primary or lateral) terminating in a

Haloragodendron gibsonii (Haloragaceae)

Telopea 11(2): 2006

143

Table 1. Vouchers for samples of eastern Australian species of Haloragodendron used in this
study

Species

Voucher

Locality

Haloragodendron lucasii

Moody 481, CONN

Cultivated, Mt Annan Botanic Garden (ex Australian
National Botanic Gardens—

Original source: Barra Brui)

H. lucasii

Moody 482, CONN

Cultivated, Mt Annan Botanic Garden
(Original source: Barra Brui)

H. lucasii

Moody 486, CONN

St Ives Chase

H. sp.

Moody 480, CONN

Cultivated, Mt Annan Botanic Garden
(Original source: Bungleboori Creek area)

H. monospermum

Moody 475, CONN

NE of Braidwood

H. monospermum

Moody 491, CONN

Cultivated, Australian National Botanic Gardens
(Original source: WNW of Kybean)

H. baeuerlenii

Moody 483, CONN

Cultivated, Mt Annan Botanic Garden
(Original source: Deua National Park)

Fig. 1. Phylogenetic hypothesis of relationships among the eastern species of Haloragodendron
based on combined matK—trnK cpDNA and ITS nrDNA sequence data analysed using parsimony
(taken from Fig. 1.3 in Moody 2005). Numbers above branches are bootstrap support; those
below are branch lengths.

144

Telopea 11(2): 2006

Wilson and Moody

subsessile flower; lateral flowers with tapering bases 1-2.5 mm long. Primary bracts of
the flowering shoot are somewhat reduced forms of normal foliage leaves that become
smaller towards the apex, with the pair subtending the terminal flowers sometimes
bract-like (lacking teeth, browning off and deciduous), 3.2-4.5 mm long, 0.7-0.9
mm wide. Bracteoles 1.8-2.3 mm long, 0.4 mm wide, inserted very close to the node.
Flowers ±sessile, solitary; petals 6-9.5 mm long, 2.5-3 mm wide; stamens 8, filaments

3- 3.5 mm long, anthers 2.5-4.S mm long, apparently dimorphic, with the antesepalous
0.5-1 mm longer than the antepetalous. Ovules 4. Immature fruit ovoid, green, 5-6
mm long and 4-5 mm broad with a fleshy pericarp; mature fruit semipersistent on the
plant, dry, dark brown, 4.5-5 mm long and 2.5-3.5(-4) mm broad, ± ovoid but weakly

4- angled and with a few coarse papillae in the lower half; pericarp woody, 0.8-1.0 mm
thick. Seed solitary. Fig. 2.

Notes: this species is close to H. lucasii but differs by the ± ovoid, unwinged hypanthia
and fruits, and the smaller flowers with petals 6-9.5 mm (compared with 9.5-12) mm
long and anthers 2.5-4.5 mm (compared with 5.5-7 mm) long. The petals are not as
markedly twisted in bud.

Etymology: the species is named for Colin P. Gibson, who made the first collection.

Distribution: seems to be confined to Bungleboori Creek and its tributaries, Blue
Mountains National Park.

Habitat and ecology: in more protected sites, c.g. sheltered gullies and on creek banks,
plants have been recorded in open forest of Eucalyptus oreades with Ceratopetalum
apetalum, Logania albiflora, Todea barbara, Gahtiia sp., and Prostanthera sp. In more
exposed sites, e.g. dissected cliff-lines and pagodas, they have been found in heath
associated with species such as Allocasuarina distyla, Bauksia ericifolia, B. conferta,
Eucalyptus stricta, Leptospermum trmervium, Acacia hamiltoniana, Olax stricta and
Caustis pentandra.

Conservation status: in Briggs and Leigh (1996) this species is coded as 2RCat (i.e.,
rare but adequately conserved with a total known population of 1000+ plants that are
all known to occur in a conservation reserve). Population numbers at individual sites
are mostly high, although at one site the plants were described as ‘either gregarious
or cloning’. There is no evidence available to assess clonality, but the species certainly
regenerates from seed: one herbarium sheet (Keith s.n., 23 Feb 1999) consists of
seedlings 15-35 cm high that had come up subsequent to a wildfire in 1994.

Sc'lected specimens examined: New South Wales: Central Tablelands: Bungleboori Creek,
Gibson s.n., 3 Oct 1982 (NSW 284758); Bungleboori Creek, Jones s.n., 26 Feb 1992 (NSW
262806); Yarramun Creek, Lembits.n., 7 Dec 1986 (NSW 171423, HO); Hole-in-the-Wall Creek,
Gibson 13 ,26 Dec 1988 (NSW); E of Mt Norris towards Bungleboori Creek, Hind5981 & Cherry,
26 Feb 1990 (NSW).

Acknowledgments

We thank Tracey Armstrong and Trish Meagher (Mt Annan Botanic Garden) for
their assistance with cultivated plants, and Gill Towler and Surrey Jacobs for field
companionship. We thank Catherine Wardrop for the fine illustration.

Haloragodendron gibsonii (Haloragaceae)

Telopea 11(2): 2006 145

C WJouvcLt/op

Fig. 2. Haloragodendron gibsonii. a, habit; b, c, leaf; d, bud (just beginning to open); e, flower;
f, developing fruit (all from Wilson 1684). Scale bars: a - 75 mm; b-d = 8 mm; e, f = 10 mm.

146

Telopea 1 1(2): 2006

Wilson and Moody

References

Briggs JD & Leigh JH (1988) Rare or threatened Australian Plants , revised edition. Australian
National Parks and Wildlife Service Special Publication No. 14.

Briggs JD & Leigh JH (1996) Rare or threatened Australian Plants, 1995 revised edition. (CSIRO
Publishing: Collingwood)

Gibson C (1998) Barabruiensis and Bunglebooriensis. The Bushland Bulletin [Newsletter of the
Bankstown Bushland Society] No. 25: 6,10.

Orchard AE (1990) Rediscovery of Haloragodendron lucasii (Haloragaceae). Telopea 3: 593-595.

Moody ML (2005) Systematics of the Angiosperm Family Haloragaceae R. Br. Emphasizing the
Aquatic Genus Myriophyllum: Phytogeny, Hybridization and Character Evolution. (Unpub¬
lished PhD thesis: University of Connecticut)

Wilson PG (1991) Haloragaceae. Pp. 208-219 in Harden GJ (ed.) Flora of New South Wales vol. 2.
(New South Wales University Press: Kensington)

Manuscript received 20 October 2005, accepted 19 December 2005

Telopea 11(2) 147-154

Lysimachia (Myrsinaceae) in New South Wales

P.G. Kodela

National Herbarium of New South Wales, Botanic Gardens Trust, Mrs Macquaries Road,

Sydney NSW2000, Australia

Abstract

This paper describes and provides a key to Lysimachia taxa occurring in New South Wales:
L. fortunei, L. japonica and L. vulgaris var. davurica.

Introduction

Lysimachia was previously placed in the family Primulaceae, but a recent phylogenetic
study analysing molecular and morphological data of primuloid families supports the
transfer of the genus to Myrsinaceae (Kallersjo et al. 2004). In addition to the synopsis
of diagnostic morphological characters of the family provided by Kallersjo et al. (2004),
many Lysimachia species have schizogeneous secretory cavities that appear as lines or
dots on vegetative and reproductive parts, which according to Anderberg and Stahl
(1995) are characteristic of Myrsinaceae.

Lysimachia is a cosmopolitan genus more common in the Northern Hemisphere. There
are about 180 or possibly up to 250 species of Lysimachia worldwide. Some species have
medical and culinary uses, particularly in Asia.

When the treatment for Lysimachia was first published in the Flora of New South Wales
(Makinson 1990), two species were known to occur in New South Wales. Since then
there have been changes to the known distribution of L. japonica (one new and one
historic record from South Coast) and L vulgaris var. davurica (extensions of range
to South Coast and Southern Tablelands), as well as an additional species, L. fortunei,
recorded for the State. In these three perennial species, the flowers are 5-merous with a
deeply dissected, ± campanulate, persistent calyx and a deeply dissected, campanulate
to rotate corolla.

This paper provides an updated brief treatment of the Lysimachia taxa occurring
in New South Wales and was undertaken in order to revise the NSW FloraOnLine
(http://plantnet.rbgsyd.nsw.gov.au/floraonline.htm). There is further scope for detailed
taxonomic assessment of the genus in Australia.

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Key to Lysimachia taxa in New South Wales

1 Stems ± prostrate, to 18 cm long, pilose with septate hairs; leaves ovate or broadly ovate to ±

spathulate, to 2.5 cm long; flowers solitary in leaf axils; pedicels without basal bract; corolla
close to or barely exceeding length of the calyx ... 1 L. japonica

1* Stems erect, to 2 m high, minutely glandular-hairy or ± glabrous; leaves elliptic to narrowly
elliptic, lanceolate or oblanceolate, mostly 3-9 cm long; flowers in panicles or racemes;
pedicels with basal bract; corolla distinctly exceeding calyx.

2 Leaves mostly opposite or in whorls, minutely glandular-hairy particularly along veins (on

both surfaces or mainly on lower surface) and margin; pedicels 5-12 mm long, slender,
distinctly longer than bract; calyx apices acuminate, margins glandular streaked; corolla
yellow, 6-9 mm long . 2 Lysimachia vulgaris var. davurica

2* Leaves mostly alternate, glabrous; pedicels 1-5 mm long, stout, as long as or shorter than
bract; calyx apices obtuse, margins membranous; corolla white, to 5.5 mm long
. 3 Lysimachia fortunei

I. Lysimachia japonica Thunb., FI. Jap.: 83 (1784)

Trailing, stoloniferous herb; stems ± prostrate, rooting at lower nodes, often ascending
towards ends, to 18 cm long or possibly longer, with ± retrorse, often slightly crisped,
hyaline, septate hairs (to c. 0.5 mm long) and scattered minute reddish glandular-
papillae. Leaves mostly opposite, occasionally subopposite or alternate on some stems;
lamina ovate or broadly ovate to ± spathulate, mostly 0.8-1.7 cm long and 5-15 mm
wide, with scattered minute reddish glandular dots (glandular punctate) and hyaline,
septate hairs; petiole 2-10 mm long, narrowly winged. Flowers solitary, axillary, usually
5-8 mm diam.; pedicel 1.5—5 mm long (reported to 10 mm long in overseas treatments),
without a bract; sepals 4—7 mm long, lanceolate with acuminate to ± subulate apex and
scattered septate hairs, glandular-spotted; petals not or scarcely exceeding the sepals,
3.5-5.5 mm long, yellow, apex acute or obtuse; filaments glabrous. Capsule globose or
subglobose, 2-3(-4) mm diam., much shorter than persistent calyx, usually minutely
pilose at apex, longitudinally dehiscing from apex by 5 valves, many-seeded. Flowering
mainly November - January; also recorded flowering in April and possibly later.
Fig. la-c. Common name: Creeping Loosestrife.

Flabitat: moist situations, in and on margins of rainforest, on stream banks and in
swamps, including swampy shallow drainage lines on flats.

Distribution: mainly north from the Williams River (Barrington area), with an historic
record from Tilba Tilba (1881) and a more recent record from near Bega (1996); also
Victoria where reported to be of uncertain status (see Walsh 1996) and widespread
from Japan and Taiwan west to the Himalaya region, India and Sri Lanka and south
through Indochina to Indonesia and Papua New Guinea. Occurrences in Queensland
and perhaps eastern Victoria would not be improbable.

Conservation status: not listed as threatened or endangered in New South Wales where
it is locally frequent to rare in various moist habitats.

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Selected specimens: New South Wales: North Coast: near Williams River, 1802-5, R. Brown
(MEL237010!, NSW155424!); Wiangarie State Forest, 27 Dec 1979, R.G. Coveny 10560 &
P.D. Hind (NSW!), southern end of Gloucester Buckelts, 1 Jan 1994, R.G. Coveny 16710 (BRI,
CANB, K, MEL, MO, NSW!), Bentley, Richmond River, 1893, A.H. Edwards (MEL1517684!);
above Shannon Creek, c. 30 km S of Grafton, 10 Apr 2002, J. & P. Edwards (NSW498015!), upper
Bellinger River, Bishops Creek, 30 Nov 1993, P. Gilmour 7512 (CANB, MEL, NSW!); Tooloom
Range, Dec 1907, J.H. Maiden (NSW155422!); South Coast: Tilba Tilba, Nov 1881, M. Bate 207
(MEL!); Jellat Flat, 5 km SE of Bega, 26 Nov 1996,7. Miles (NSW433736); Northern Tablelands:
N of Mooraback Rest Area, Werrikimbe National Park, 5 Dec 1987, J.R. Hosking (NSW213496!,
TARCH).

2. Lysimachia vulgaris var. davurica (Ledeb.) Knuth, Pflanzenr. IV 237 (Heft 22): 304
(1905)

L. davurica Ledeb., Mem. Acad. Imp. Sci. St Petersbourg Hist. Acad. 5: 523 (1812)

Rhizomatous herb; aerial stems erect, simple or weakly branched, to 1.5 (-2) m high,
with minute, erect, reddish brown-tipped glandular hairs (to 0.1 mm long or slightly
longer) and some times with occasional longer septate hairs intermixed, becoming
glabrous on lower stem. Leaves mostly opposite or in whorls of 3 or 4, elliptic to
narrowly elliptic, lanceolate or oblanceolate, mostly 4-9 cm long and 10-22 (-32) mm
wide (leaves becoming smaller towards stem apex), often long and narrow (to 10.5
cm long, 7-8 mm wide) on young plants, discolorous, green, paler beneath, minutely
glandular-hairy, dotted with orange to blackish glands (glandular punctate), sessile
or with petiole to 3 mm long. Panicles terminal and upper-axillary, many-flowered.
Flowers 9-15 mm diam.; pedicel mostly 5-12 mm long, with a distinctly shorter basal
bract 2-6 mm long; sepals 3-4 mm long, lanceolate-ovate with an acuminate apex
and distinct orange to blackish (on drying) glandular line on the outer surface near
the margins, ciliolate with gland-tipped hairs; petals clearly exceeding the sepals, 6-9
mm long, bright yellow, often with minute orange to reddish glandular lines or dots,
apex broadly acute to obtuse; filaments with minute glandular hairs. Capsule globose
or subglobose, 2-4 mm diam., longer than persistent calyx when mature, with minute
gland-tipped hairs at apex, longitudinally dehiscing from apex by (4—)5 valves, many-
seeded. Flowering December - March. Aerial stems die back after flowering and fruiting.
Fig. ld-f. Common name: Yellow Loosestrife.

Habitat: moist situations, including peatland swamps, stream banks and lagoons.
Grows in damp soils to saturated peat, in sedgeland and wet heath communities,
associated with Phragmttes australis, Carex spp., Juncus spp., Bolboschoenus fluviatilis,
Lycopus australis, Lythrum salicaria, Leptospermum spp. and Melaleuca ericifolia.

Distribution: rare in Australia; recorded in several wetlands in New South Wales,
including Wingecarribee Swamp between Robertson and Moss Vale, a swamp/pond
system in the Lower Boro - Braidwood area, and a lagoon near Bega. These populations
are far disjunct from the main natural range of this taxon in NE Asia where it occurs
from Japan west to Mongolia and eastern Russia. Its occurrence in Australia may provide
evidence supporting long-distance dispersal; in this case the agency is likely to have
been migratory waterbirds, such as Latham’s (Japanese) Snipe, which visit Australian
wetlands in the summer. Both var. davurica and var. vulgaris have been recorded in
Victoria with uncertain status (see Walsh 1996).

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Fig. 1. Lysimachia japonica. a, b, part of stem; c, flower. L. vulgaris var. davurica. d, e, part of stem;
f, flower. Lfortunei. g, h, part of stem; i, flower, a-c,/. & P. Edwards (NSW498015); d-f, /. Miles
(NSW6J5689); g-i, L.M. Copeland3557, I.R. Telford & J. Hodgon (NSW).

Scale bar = 0.24 cm (g), 0.3 cm (a, d), 0.75 cm (i), 1 cm (c), 2 cm (b), 6 cm (f, h) or 8 cm (e).

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Conservation status: listed as Endangered under Schedule 1 of the New South Wales
Threatened Species Conservation Act 1995 and is not located in any protected reserves.
The largest population in New South Wales is under threat from major disturbance
of its wetland habitat since most of the Wingecarribee Swamp peatland collapsed and
fragmented in August 1998.

Selected specimens: [precise localities withheld for conservation purposes] New South Wales:
South Coast: SEofBega, 18 Dec 2002,/. Miles (NSW615689!); ibid. 21 Feb 2003 (NSW702297!);
Central Tablelands: Wingecarribee Swamp, 26 Feb 1969, R.G. Covetiy 906 (CANB, MEL, NSW!),
9 Feb 1988, P.G. Kodela (NSW254143!, UNSW!), 27 Feb 1997, P.G. Kodela (CANB!, K!, MEL!,
MO!, NSW425850!, PE!, UNSW!), 28 Mar 2001, P.G. Kodela 574 cl al. (CANB!, MEL!, MO!,
NSW!), 1 Jun 2001, P.G. Kodela 600a (NSW!); Southern Tablelands: Lower Boro - Braidwood
area, 19 Feb 2001, R.O. Makinson 1787 (CANB).

Notes: A possible taxonomic synonym is L. tnuelleri N.P.Balakr., the replacement name
for L. salicifolia Benth. nom. illeg. (Walsh 1996). Investigations by Bob Makinson (pers.
comm.) indicate likely synonymy, which has implications for the status of L. vulgaris var.
davurica as native based on historic specimens from Victoria labelled as L. salicifolia.

Lysimachia vulgaris L. var. vulgaris and var. davurica are some times treated as separate
species (e.g. Hu & Kelso 1996). The present paper maintains the variety level, with a
molecular phylogenetic study finding these taxa to be very close (Hao et al. 2004) and
there being strong morphological similarities (pers. obs.).

Lysimachia vulgaris var. vulgaris is native mainly to Europe and is most readily
distinguished from var. davurica by having conspicuous, long, septate hairs on the
stems, leaves and other parts (pers. obs.). In var. davurica there may be occasional long
septate hairs amongst predominantly minute glandular hairs. The leaves may also be
broader in var. vulgaris, to 50 mm wide. There are occasional records of L. vulgaris
(variety uncertain) from New Zealand and the species is treated as naturalised there.
In parts of North America L. vulgaris is a problem weed of wetlands that was probably
introduced from Eurasia as an ornamental.

In Chinese folk medicine flavanol glycosides have been extracted from Lysimachia
vulgaris var. davurica for treatment of high blood pressure.

3. Lysimachia fortunei Maxim., Bull. Acad. Imp. Sci. Saint-Petersbourg 12: 68 (1868)

Rhizomatous herb; aerial stems erect (to decumbent), simple or some times branching
near apex, reddish, to c. 70 cm high, with sparse, minute, gland-tipped hairs, appearing
± glabrous. Leaves mostly alternate, narrowly elliptic or oblanceolate, mostly
3-6.5 cm long and 6-12 mm wide, glabrous, dotted with minute reddish glands
(glandular punctate), ± sessile. Racemes terminal, many-flowered, to 15 cm long.
Flowers 5-7 mm diam.; pedicel mostly 1-2 mm long (occasionally to 5 mm long in
basal flowers), with a ± equally long or longer basal bract 2-4(-5) mm long; sepals
1.5-2.5 mm long, ovate-elliptic with obtuse apex, glandular-dotted, margins whitish-
membranous; petals exceeding the sepals, to 5.5 mm long, white, often with minute
purplish glandular dots and lines, apex rounded; filaments with minute glandular
hairs. Fruits not seen. Flowering around February. Fig. lg-i. Common name (Japan):
Swamp Tiger Tail.

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According to Iwatsuki et al. (1993) and Hu and Kelso (1996) the stems can reach 80 cm
high, the leaves can also be lanceolate and to 11 cm long and 25 mm wide, the racemes
to 20 cm long and the capsule is globose, 2-2.5 mm diam., glabrous, dehiscing by valves
and many-seeded.

Habitat: moist situations; recorded in New South Wales from a broad gully swamp,
growing in peaty loam over basalt, on stream bank and Hat, in wetland herbfield with
Carex, Geum and Juncus.

Distribution: recorded in Australia only from a swamp in Werrikimbe National Park
where it occurs possibly as a result of long distance dispersal by a wetland bird; main
natural distribution is in E Asia from China to Japan.

Specimens examined: [precise locality withheld for conservation purposes] New South Wales:
Northern Tablelands: Werrikimbe National Park, 13 Feb 2003, L.M. Copeland 3557,1.R. Telford
& J. Hodgon (BRI, CANB, MEL, NE, NSW!, TARCH) & L.M. Copeland 3566, I.R. Telford & J.
Hodgon (CANB, MEL, NE, NSW!).

Conservation status: rare in Australia, being only recorded from one wetland which
is protected in Werrikimbee National Park; status requires further investigation, but
L.fortnnei is likely to be a ‘natural adventive’ or ‘distance-dispersed native’ based on the
similar distribution patterns of the other species.

Note: in addition to the more distinct differences in leaf arrangement and inflorescence
features (including flower colour), L.fortnnei can be distinguished from L. vulgaris var.
davurica by the indumentum. In L. fortunei the stems appear ± glabrous with only a
few scattered minute hairs while in L. vulgaris var. davurica the minute indumentum is
usually moderately dense, at least in the upper parts, as seen with a lOx lens.

Conclusion

Three Lysimachia taxa are currently known to occur in New South Wales: L. fortunei,
L.japonica and L. vulgaris war. davurica. In Australia they are also known from Victoria,
along with the commonly cultivated species L. nutnmularia L. which has been recorded
as naturalised.

Because Lysimachia is not widespread or common in Australia and some species are
sold for cultivation as ornamentals it has often been assumed that records in the wild
represent naturalised escapes in this part of the world. For example, Lysimachia vulgaris
var. davurica in Wingecarribee Swamp attracted great debate in 1997 at a mining
warden’s inquiry on peat mining as to whether it was a rare native or introduced weed
growing in the wetland. The presence of this taxon and L.fortnnei in particular, in New
South Wales, could well provide examples of chance distributions resulting from long
distance dispersal, which is not uncommon for cosmopolitan wetland plants. There is
no obvious garden escape pattern for the genus in New South Wales and some of the
populations are in remote localities. The question of how long some of the populations
have been in Australia remains unanswered. Palynological and macrofossil analyses of
peat or sediments where Lysimachia grows might be ways of investigating pre-European
occurrence. From what is known about the distribution of the genus and the history
of specimen records, Lysimachia is likely to be an elusive cosmopolitan in Australia.
It is quite possible that new localities for Lysimachia will be found in Australia in the
future.

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Acknowledgments

I am grateful to Bob Makinson, Peter Wilson and Elizabeth Brown for constructive
comments on the manuscript. I thank Catherine Wardrop for her skilful illustration.
MEL kindly allowed access to specimens.

References

Anderberg AA & Stahl B (1995) Phylogenetic interrelationships in the order Primulales, with spe¬
cial emphasis on the family circumscriptions. Canadian Journal of Botany 73:1699-1730.

Iwatsuki K, Yamazaki T, Boufford DE & Ohba H (eds) (1993) Flora of Japan, vol. Ilia. (Kodansha:
Tokyo)

Kallersjo M, Bergqvist G & Anderberg AA (2000) Generic realignment in primuloid families
of the Ericales s.l.: a phylogenetic analysis based on DNA sequences from three chloroplast
genes and morphology. American Journal of Botany 87: 1325-1341.

Makinson RO (1990) Primulaceae. Pp. 504-506 in Harden GJ (ed.) Flora of New South Wales,
vol. 1. (New South Wales University Press: Kensington)

Walsh NG (1996) Primulaceae. Pp. 517—522 in Walsh NG & Entwisle TJ (eds) Flora of Victoria,
vol. 3. (Inkata Press: Melbourne)

Hao G, Yuan Y-M, Hu C-M, Ge X-J & Zhao N-X (2004) Molecular phylogeny of Lysimachia
(Myrsinaceae) based on chloroplast truL-F and nuclear ribosomal ITS sequences. Molecular
Phylogenetics and Evolution 31: 323-339.

Hu C-M & Kelso S (1996) Primulaceae. Pp. 39-189 in Wu Z-Y & Raven PH (eds) Flora of China,
vol. 15. (Science Press: Beijing & Missouri Botanical Garden: St Louis)

Manuscript received 10 October 2005, accepted 7 February 2006

.

Telopea 11(2) 155-160

Three new species of Nymphaea
(Nymphaeaceae) in Australia

Surrey W.L. Jacobs 1 and C. Barre Hellquist 2

'National Herbarium, Royal Botanic Gardens, Sydney, NSW, 2000, Australia
Author for correspondence: surrey.jacobs@rbgsyd.nsw.gov.au
2 Department of Biology, Massachusetts College of Liberal Arts, North Adams,
MA 01247-4100, USA.

Abstract

Nymphaea alexii (aff. N. hastifolia, subgenus ‘Confluentes’), N. carpentariae and N. georginae
(both aff. N. macrosperma, subgenus Anecphya) are described from the Gulf Savannah region
of Queensland and immediately to the south. A key is provided for the native and naturalised
species of Nymphaea in Australia.

Introduction

As part of continuing studies in Nymphaea (Jacobs 1989, 1992, 1994), including
sampling for an extensive DNA study, we have been able to collect good material
sufficient to describe three new species.

Nymphaea alexii S.W.L.Jacobs 8< Hellq. sp. nov.

N. hastifoliae similis sed staminibus cremiis, apice ovarii rubro, cristis seminis
nonnunquam carunculatis differt.

Holotype. Queensland: c. 23 km N of Normanton, Karumba rd, 17° 31.741'S 141°
09.625'E, S. Jacobs 9325 & C.B. Hellquist, 17 Apr 2005 (NSW). Isotypes: NASC, BRI.

Annual or perennial with a globose rhizome c. 2 cm diarn. Blade elliptic, to 15 cm
long, to 10 cm wide; margins slightly sinuate; stipules fused for c. 1 cm, the apical lobe
free, acute c. 1 cm long. Flowers to 30 cm above the water, pleasantly-scented, day¬
flowering. Sepals 4-5, to 6 cm long, green outside; tip acute. Petals (18—)20—25(—40), to
5.5 cm long, 1.5 cm wide, lanceolate, white, grading into the stamens, no gap between
petals and stamens; tip acute. Stamens cream, to c. 150; filaments membranous, to
17 mm long; anthers to 10 mm long; appendage white, much reduced and only visible
on outer stamens. Ovary often red at the apex, with vestigial or obsolete sterile lobes;
carpels 8 to 16; fruit globose, c. 4.5 cm diam. Seeds elongated, c. 1—2 mm long, glabrous,
with longitudinal ridges, parts of the ridges sometimes proliferating irregularly into
linear outgrowths when mature; cells of the testa with a comparatively large lumen (as
judged from surface view) and short arms of more or less equal length, with a single

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margin (as opposed to the apparent double margin in N. hastifolia), ends of the arms of
the epidermal cells neither expanded nor raised, the cell walls covered with regularly-
arranged minute papillae (visible at c. 500X on a SEM).

Selection of specimens examined: Queensland: c. 23 km N of Normanton, Karumba rd,
Jacobs 9325a & Hellquist, 17 Apr 2005 (NSW, NASC, BRI); c. 25 km S of Normanton, Croydon
rd, Jacobs 9326 & Hellquist, 17 Apr 2005 (NSW, NASC, BRI).

N. alexii grows in ephemeral billabongs and the shallow margins of more perennial
lagoons (similar to N. hastifolia) during the end of the Wet and shortly after, and has
often disappeared by May. It can be readily distinguished from N. hastifolia, or indeed
from any other native Nymphaca sp., by its distinctive cream stamens and its small,
ridged seeds. This species belongs in subgenus ‘Confluentes' 1 .

The species is named after Alex James Fussell, grandson of SWLJ.

A minutely papillose seed surface, has not been observed elsewhere in subg. Anccphya
or ‘ Confluentes ’. Interestingly, the seeds of this taxon are similar to those of Ondinea
purpurea, as depicted by Schneider and Ford (1978), both in their longitudinal ridges
and the granulate surface. A similar seed surface is known elsewhere in Nyniphaea only
in a group of species within subg. Hytirocallis.

Nymphaea carpentariae S.W.L.Jacobs 8< Flellq. sp. nov.

N. macrospertnae similis sed floribus albidis plerumque grandioribus, seminibus
minoribus differt.

Holotype. Queensland: Burketown, bore drain of town bore, 17°44.879'S 139°32.899'E,
S. Jacobs 9329 & C.B. Hellquist, 18 Apr 2005 (NSW). Isotypes: NASC, BRI.

Perennial with a globular to elongate rhizome to c. 4 cm long. Blade orbicular to elliptic,
to 45 cm diam.; margins with regularly-spaced triangular teeth to 1.5 mm long; stipules
fused for (1—)4—8 cm, terminal lobe free, acute, 0.5-1 cm long. Flowers to 40 cm above
the water, lightly scented; day-flowering. Sepals 4, to 6.5 cm long, green outside with
purple streaks, streaks occasionally obscuring the green; tip obtuse. Petals mostly
12-22, 4-6 cm long, 1.5-2.5 cm wide, oblanceolate to spathulate, mostly white, rarely
with some blue, with a space of c. 1 cm between petals and stamens; tip obtuse; when
blue, most of the petals coloured, not just the outer petals and fading only slightly with
maturity. Stamens yellow, mostly 150-300; filaments membranous to cylindrical, to
25 mm long; anthers 2.5-5 mm long, appendages vestigial or obsolete. Ovary lobes
vestigial or obsolete; carpels 7—19; fruit globose, c. 4 cm diam. Seeds spherical to
elongate-spherical, 2-3.5 mm long, c. 2 mm wide, with more or less continuous rows
of short hairs usually c. 0.1-0.15 mm long; cells of the testa with a lumen of variable
width and arms of unequal length with the ends of the arms expanded but not raised.

Selection of specimens examined: Queensland: Between Normanton and Maggieville,
Clarkson 2697 , 6 Nov 1979 (NSW681541); c. 4 km N of Normanton, Karumba rd, Jacobs 9324 &
Hellquist, 17 Apr 2005 (NSW);Boogan Lagoon, Jacobs 1280, 26 Apr 1974 (NSW681369); Forked
Lagoon ‘Wernadinga’, Jacobsl368, 30 Apr 1974 (NSW681 368); E of Croydon, Georgetown rd,
Jacobs 8588 cF Lcs, 22 Oct 1999 (NSW440424); Cumberland Chimney, c. 22 km W of Croydon,
Georgetown rd, Jacobs 9320 & Hellquist, 16 Apr 2005 (NSW).

1. Formal description of the subgenus Confluentes is waiting on the publication of the treatment
of this group in the Flora of Australia series.

Three new species of Nymphaea (Nymphaeaceae)

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157

Fig. 1. Nymphaea seeds; the bare patches, or areas where the features are less well defined, are
those areas where the seeds pack tightly next to each other, a, Nymphaea alexii (SJ9325) seed
showing the ridges and the proliferation or caruncles present on most seeds (c. 35X); b, N. alexii
(SJ9325) seed showing minimal proliferation, some proliferation is visible at top right; the ridges
are clearly visible. This seed is from the same collection as Fig. la and all fruits examined have
both types of seed (c. 35X); c, N. alexii (SJ9325) seed surface showing the characteristic papillae
on the surface of the epidermal cells (c. 425X); d, N. carpcntariae (SJ9329) whole seed showing
the more or less continuous rows of hairs and a smaller seed than N. georginae in Fig. le (c.
25X); e, N. georginae (SJ9335) whole seed showing the partly disorganized and incomplete rows
of hairs and a larger seed than N. carpcntariae in Fig. Id (c. 15X); f, N. georginae (SJ9335) seed
surface showing the epidermal cells with a narrow lumen and short irregular arms (c. 225X)

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Jacobs and Hellquist

N. carpentarine grows in perennial or near perennial billabongs and lagoons around the
Gulf of Carpentaria. It can be distinguished from N. macrosperma by the usually larger
white flowers, and from N. georginae by the smaller seeds and by the blue-flowered
forms of the latter fading with age. Although the differences in shape and dimensions
of the seeds in N. carpentariae and N. georginae do not seem very great, when compared
side by side the differences are quite striking. This species belongs in subgenus Anecphya.
Specimens of this species have usually been included in N. immutabilis.

This is the original ‘Albert De L’Estang’ sent to Bailey in Brisbane; there are still some of
the original distinctive seeds in the Herbarium (BRI). Most of what is now grown and
sold as cv. ‘Albert De L’Estang’ appears to be N. immutabilis. The cultivar ‘Andre Leu’ is
N. carpentariae.

The species is named for the Gulf of Carpentaria region in which it grows.

Nympliaea georginae S.W.L.Jacobs & Hellq. sp. now

N. tnacrospermae similis sed floribus in quoque loco plerumque albidis caeruleisque,
floribus caeruleis decolorantibus differt. A N. gigantea seminibus majoribus differt.

Holotype. Queensland: Georgina River, Camooweal, 19° 55.576'S 138° 06.903'E, S.
Jacobs 9332 & C.B. Hellquist, 19 Apr 2005 (NSW). Isotypes: NASC, BRI (blue-flowered
specimen).

Perennial with a globular rhizome c. 4 cm diam. Blade orbicular to elliptic, to 60
cm diam.; margins with regularly-spaced narrow-triangular to triangular teeth to
2-4 mm long; stipules fused for 1-3 cm with free acute terminal lobes 0.3-1 cm long.
Flowers to 30 cm above the water, fragrant, day-flowering Sepals 4, to 6.5 cm long,
green outside with purple streaks, streaks occasionally obscuring the green; tip obtuse.
Petals mostly 12-26,4-7 cm long, 2-3.5 cm wide, oblanceolate to spathulate, white or,
less commonly, blue, rarely pink, and fading with age, with a space of c. 1 cm between
petals and stamens; tip obtuse; when blue, most of the petals coloured, not just the
outer petals. Stamens mostly 150-250; filaments membranous to cylindrical, to 25 mm
long; anthers to 6 mm long, appendages vestigial or obsolete. Ovary often red with
lobes vestigial or obsolete; carpels 7-19; fruit globose, c. 4 cm diam. Seeds globose
to subglobose, (1.5—)2.5—4 mm diam., with often interrupted rows of short hairs
c. 0.1-0.15 mm long; cells of the testa with a comparatively long-narrow lumen and
arms of unequal length with ends slightly expanded but not raised.

Selection of specimens examined: Queensland: Georgina River, Camooweal, Jacobs 5531 &
P. Wilson, 4 May 1988 (NSW280651); Jacobs 9332 6' Hellquist, 19 Apr 2005 (NSW); Flood chan¬
nel of Thomson River, Longreach, Jacobs 9335 & Hellquist, 20 Apr 2005 (NSW).

N.T.: James River, nr ‘Avon Downs’, Chippendale s.n., 20 Jun 1960 (NSW); Jacobs 5300,
5303, 5304, 5306 & K. Wilson, 7 Jun 2005 (NSW); Jacobs 9331 & Hellquist, 20 Apr 2005
(NSW).

Nympliaea georginae grows in the billabongs and flood channels of the upper parts
of northern rivers flowing into the Lake Eyre system. These waterbodies may hold
water for >1 year but also are frequently dry for >1 year. It can be distinguished from
N. carpentariae by the larger seeds and from N. macrosperma by the larger flowers of
blue-flowered plants fading with age. This species belongs in subgenus Anecphya. This
species has the strongest scent of any in subg. Anecphya, but it is still considerably less
scented than species of subg. ‘Confluentes’. Specimens of this species have usually been
included in either N. gigantea or N. immutabilis.

Three new species of Nymphaea (Nymphaeaceae)

Telopea 1 1(2): 2006

159

The species is named for the Georgina River, one of several in which it grows and the

type locality.

Key to native and naturalised species of Nymphaea in Australia.

1. Petals yellow, grading into stamens (with filaments to 38 mm long); leaf blade margin

sinuate; horizontal stolons and vertical rhizomes both present. N. mexicana 2 3

1* Petals white, blue or pink; grading into stamens or a distinct gap present; leaf blade margins
various; stolons absent; rhizomes either horizontal, or vertical and more or less tuberous.
. 2

2. Rhizomes horizontal or suberect, elongated and vigorous; flowers more or less floating on

the water surface. N. alba, N. odorata and hybrids

2* Rhizomes tuberous, erect; flowers standing clear of the water surface. 3

3. Petals grading into stamens; leaf blade with entire, sinuate or dentate margins. 4

3* Distinct gap present between petals and stamens; blade with sinuate or dentate margin. .. 9

4. Filaments flattened, thickened, tough; leaf blade margin dentate; blade undersurface usually

pubescent . JV. pubescens

4* Filaments membranous and either cylindrical, or all flattened, or only some outer filaments
flattened; leaf blade undersurface always glabrous; blade margin sinuate. 5

5. Anthers to 24.5 mm long; apical appendage to 10 mm long. N. caernlea'

5* Anthers 13 mm long or less; apical appendage minute or absent. 6

6 Anthers cream; top of ovary frequently red; seeds with longitudinal ridges and these often
with proliferations. N. alexii

6* Anthers yellow; top of ovary rarely red; seeds without longitudinal ridges or proliferations.
. 7

7. Filaments to 25 mm long; sepals to 11.5 cm long; petals blue to mauve, white, or pink;

widespread; habitats various. n. violacea

7* Filaments to 18 mm long; sepals to 7 cm long; petals white. 8

8. Sepals usually with purple flecks, to 7 cm long; anthers to 8.5 mm long, sometimes apiculatc;

carpels 11-22; fruit to 2.5 cm diam.; seeds glabrous, 1.75-2.5 mm long, 1-1.5 mm wide, cells
of testa without a ‘double edge; growing in more or less permanent water; Cape York
Peninsula, New Guinea. /y. elleniae

8* Sepals without purple flecks, to 6 cm long; anthers to 5.5 mm long; appendage absent; carpels
8-16; fruit to 4.5 cm diam.; seeds glabrous, spherical, c. 1 mm diam., cells of testa with a
‘double edge’; growing in ephemeral water on floodplains; N.T., W.A. N. Iiastifolia

9. Filaments flattened and tough; petals usually <10; stamens usually <25; leaf blade

margin sinuate; mature seed <1.5 mm long. N. noucliali

9* Filaments membranous, slightly flattened to cylindrical; petals usually >10; stamens
usually >50; leaf blade margin toothed; mature seeds >2.5 mm long. 10

10. Sepal margins pink; petals becoming dark pink with age; blade margin with sparse teeth to

2 mm long... N. atrans

10* Sepal margins white or blue, rarely pink; petals not darkening with age, teeth usually more
than 2 mm long. 11

2. I he plant introduced in Australia is a hybrid involving this species, rather than pure
N. mexicana, as in other places where this taxon is said to be introduced.

3. 1 he introduced plant in eastern Australia is not typical of N. caerulea, originally described from
Egypt, but better fits what has traditionally been treated as N. capensis. However, further work on
the interpretation of the type of this latter name is required (J. Wiersema pers. comm.).

160

Telopea 11(2): 2006

Jacobs and Hellquist

11. Anthers to 6 mm long; sepals to 6.5 cm long; petals to 26 on fully developed flowers, white,

deep blue, rarely pink; blade margin with teeth to 4 mm long. 12

11 * Anthers >6 mm long; sepals to 12 cm long; petals to 34 on fully developed flowers, blue,
white, or rarely pink; blade margin with teeth to 5 mm long. 14

12. Flowers mostly blue, rarely pink or white, comparatively small; petals to 5 cm long

and all the same colour; sepals to c. 4.5 cm; leaves comparatively large with teeth on
mature leaves cylindrical from a shallow triangular base; seeds globose to subglobose,
3-4.5 mm diam... N. tnacrosperma

12* Flowers mostly white with some blue, rarely pink, larger; petals to 7 cm long and all
the same colour; sepals to 6.5 cm; leaves not as large with teeth on mature leaves triangular
to narrow-triangular; seeds globose to elongate-spherical, 2-4 mm diam. 13

13. Seeds spherical to elongate-spherical, 2-3.5 mm long, c. 2 mm wide, with more or

less continuous rows of short hairs; flowers mostly white, rarely blue, no obvious fading
with age.. . N. carpentariae

13*Seeds globose to subglobose, 2.5-4 mm diam., with often interrupted rows of short hairs;

(lowers mostly white but blue flowers still common, the blue flowers fading with age.

. N. georginae

14. Anthers to 10 mm long; rarely apiculate; when coloured the petals fading with age; carpels

12-18; seeds ovate, pubescent, the hairs arranged in more or less continuous rows of short
hairs; cells of the testa with a long lumen and short arms of equal length, the ends not raised,
rarely expanded. N.gigantea

14*Anthers to 15 mm long; often apiculate; when coloured the petals usually not fading with
age; carpels 9-20; seeds oblong, pubescent or rarely glabrous, with sparse to dense hairs in
discontinuous or disorganised rows, sometimes almost appearing scattered; cells of the testa
with a long lumen and arms of equal to unequal length, the ends sometimes slightly expanded
but not raised. N. imntutabilis

Acknowledgments

We thank Andre Leu (Wonga Beach, Qld) for showing us cultivated specimens and
sharing his locality information, Karen Wilson (NSW) for the Latin diagnoses, and John
Wiersema (Beltsville, Maryland, USA) for continued discussion and encouragement.

References

Jacobs SWL (1989) Nymphaea in Australia. Water Garden Journal 5: 14-18.

Jacobs SWL (1992) New species, lectotypes and synonyms of Australasian Nymphaea (Nym-
phaeaceae). Telopea 4: 635-641.

Jacobs SWL (1994) Further notes on Nymphaea in Australasia. Telopea 5: 703-706.

Schneider (1978) and Ford EG (1978) Morphological studies of the Nyntphaeaceae. X. The seed
of Ondinea purpurea Den Hartog. Bulletin of the Torrey Botanical Club 105: 192-200.

Manuscript received 20 September 2005, accepted 7 February 2006

Telopea 11(2) 161-170

An anatomo-ecological experiment
in Austrostipa aristiglumis, a lowland
Stipoid species.

Mirta O. Arriaga 1 and Surrey W.L. Jacobs 2

'Museo Argentine de Ciencias Naturalcs, Av. A. Gallardo 470, C1405DJR, Ciudad Autonoma de
Buenos Aires, Argentina. E-mail: marriaga@macn.gov.ar
2 National Herbarium, Royal Botanic Gardens, Sydney, New South Wales, 2000, Australia. Author
for correspondence: surrey.jacobs@rbgsyd.nsw.gov.au

Abstract

Leaf and sheath anatomy characteristic of an aquatic or amphibious plant is reported for
Austrostipa aristiglumis (Gramineae). Austrostipa and the rest of the Stipoid grasses are
normally considered to be classic xeromorphs. The degree of development of the hydromorphic
characteristics appears to be related to the level of soil saturation. The ecological and phylogenetic
significance of these observations is discussed.

Introduction

The Stipeae is a cosmopolitan grass tribe of c. 450 species, widely distributed and not
clearly delimited (Barkworth & Everett 1987). The tribe has major centres of diversity
in South and North America, Australia and Eurasia, with different genera dominating
in each centre. The tribe is characterised by having florets with a single spikelet, the
lemma usually being coriaceous or firmer, having comparatively large unicellular
macrohairs, and a range of other epidermal features that help delimit the genera.

The species grow primarily in temperate or warm-temperate regions, and are dominant
in many of the arid grasslands of southern Australia, South America, and Asia at varying
elevations (0 -5000 m) and the ecological range includes arid, semiarid, semihumid
and humid habitats (Barkworth & Torres 2001).

Within the tribe, there are recurring patterns of morphological and anatomical
variation. Anatomy in the Stipeae has been the subject of numerous studies, with the
majority on leaf anatomy being carried out for taxonomic purposes. Parodi (Parodi
1944; Parodi & Freier 1945) proposed that leaf anatomy was of value in determining
generic limits in the tribe. Later studies in American species, have confirmed the
importance of leaf (as well as lemma and palea) anatomical characters (Brown 1958;
Matthei 1965; Reeder 8< Reeder 1968; Caro & Sanchez 1971,1973; Clifford & Watson

© 2006 Royal Botanic Gardens and Domain Trust

ISSN0312-9764

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1977; Thomasson 1978; Barkworth 1981; Scholz 1982; Watson & Dallwitz 1981, 1992;
Arriaga 1982, 1983; Renvoize 1985; Roig 1964 [a & b], 1965; Torres 1993,1997 [a , b &
c] ; Penailillo 1996; Rojas 1997; Cialdella & Arriaga 1998; Arriaga & Barkworth 2000;
Cialdella & Giussani 2002).

Taxa from south-west Asia were revised by Freitag (1985), and the Australasian stipoid
taxa by Vickery et al. (1986), Jacobs et al. (1989) and Jacobs and Everett (1996).Vazquez
(1992), Vazquez and Devesa (1996, 1997) and Vazquez and Barkworth (2004) treated
the stipoids of Morocco and the Iberian peninsula, and Martinovsky (1976) those of
central Europe. However, anatomical characters were not included in these studies.
This study is part of a larger project being undertaken by the International Stipeae
Working Groups to explore the diversity within the Stipeae with the ultimate goal of
developing a strongly supported taxonomic revision of the tribe. The results could be
used to predict how the distribution of stipoid species, and other grasses with similar
characteristics, would be affected by changes in climate.

Stipoids are known to be species adapted to mesic and xeric environments. With
the exception of Trikeraia (Clayton & Renvoize 1986), there is no mention of any
relationship with water in the descriptions of the species habitat, nor any mention of
hydromorphic characteristics, in the tribe.

In this contribution we describe the leaf and sheath structure of Austrostipa aristiglumis
(E. Muell.) S.W.L. Jacobs & J. Everett, a species of semi-arid to sub-humid habitats
that has hydromorphic characters that change with the level of soil saturation. While
Austrostipa aristiglumis grows in a variety of habitats, it is most common on the
intermittently-flooded, grey self-mulching clays or heavy alluvial soils of the westerly¬
flowing watercourses of the winter-dominant rainfall areas of the Murray-Darling
Basin.

Methods

Six pots 190 mm internal diameter and 190 mm high were planted with diaspores of
Austrostipa aristiglumis (RK24, voucher held at NSW ) in November 2002, at Sydney,
Australia as this is the time of year seedlings are found in the field. The pots were placed
outside in five trays 6, 11, 44, 86, and 136 mm deep, and one pot (0 mm) retained as
a control with no tray. The treatments were arranged linearly in random order from
random number tables (0 mm position 4; 6 mm position 5; 11 mm position 2; 44 mm
position 3; 86 mm position 6; 136 mm position 1). The pots were filled with a mixture
of potting mix and clay. The trays were each maintained filled with water so that each
pot had different levels of saturated soil. Each time the trays were filled the control was
also watered and allowed to freely drain. The pots also received natural rainfall.

One year later, after flowering, all the surviving plants from different treatments and
the control (0mm, 11 mm, 86 mm) were preserved in 70% ethyl alcohol in November
2003, and herbarium voucher specimens prepared from the 11 mm and 86 mm
treatments). This date was selected for harvesting because the plants had flowered and
good vouchers could be collected.

1. For further information on this group contact Dr S. W. L. Jacobs, Royal Botanic Gardens,
Sydney, New South Wales, 2000, Australia.

Austrostipa aristiglumis, a lowland Stipoid species

Telopea 11(2): 2006

163

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Fig. 1. Iransections of leaves and sheaths of Austrostipa aristiglumis. Schemes representing: a,
TS at the middle of the sheath; b, TS of sheath below the ligule region; c, TS of leaf beneath the
ligule region; d, TS at the middle of the leaf. Legend: • Total leaf area expresed in pm 2 ; O Total
cavities area expresed in pm 2 ; * relationship of air cavities to total leaf area expressed in %.

164

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Arriaga and Jacobs

Transverse sections were made from leaves and sheaths of preserved specimens.
Sections, from the middle segment of lamina and sheath and from 5 mm over and
under the Iigule portion, of the second or third lamina of a sterile innovation, were
obtained either freehand or by embedding the material in wax and sectioning with a
rotary microtome. The sections were stained with Aldan Blue/Safranin (Cutler 1978)
and observed and photographied using a Leica DMLB optical microscope with a JVC
digital camera. Drawings of the cross-sections were prepared using a camera lucida.
Measurements of total leaf area and total cavities area were made with the IM50 Leica’
software, averaged from 10 measurements each.

Results

Leaf anatomy in transection

The transection (TS) outline is U-shaped (Fig. 1). Both semilaminae may be symmetric
or asymmetric (differing in one rib). The upper surface has ribs and deep (>50% of
leaf thickness) and narrow furrows. Ribs alternate between large and flat-topped, and
smaller and round-topped; larger ribs are coincident with one 1 st or 2 nd order vascular
bundle (VB); smaller ribs are coincident with 3 rd order vascular bundles (Figs 1,2). The
lower surface varies from flat to slightly angular.

The midrib is scarcely distinguishable with only one I s ' order VB located in the centre of
the transection (Figs 1,2). Neither colourless parenchyma nor transitional parenchyma
arc associated with the midrib. There are subepidermal strands of sclerenchyma under
both epidermes in the midrib (Fig. 1). In specimens growing in more than 10 mm
of water aerenchyma is associated with the midrib, situated in channels, laterally to
the VB in TS made 5mm above the Iigule (Fig. 2); aerenchyma is not present in TS
made in the midrib at the middle level of the leaf. When present, aerenchyma (and the
corresponding lacunae in TS which contains it) represents 20 -40% of the TS.

The semi-laminae have >6 VBs, alternating 1 st , 2 nd and 3 rd order; 1 st and 2 nd order
are situated in the centre of the TS or towards the lower surface; all 3 rd order VBs
are towards the lower surface (Figs 1, 2). No colourless parenchyma is present in the
semilaminae. Sclerenchyma is present under both epidermes, fragmented, in girders,
sub-epidermal transitional parenchyma is also present adaxially. Sclerenchyma and
transitional parenchyma produced over all types of VBs, heavily over 1 st and 2 nd order
(Figs 1, 2). Aerenchyma in the basal portions of the semi-laminae is present in leaves
of specimens growing with >10 mm of water in the tray, channels between VBs are
present in 60% of the semi-laminae.The channels with aerenchyma occupy 20 -40%
of the depth of the TS. The l" 1 order VBs in the semilaminae are elliptic to ovate, with a
strand of phloem adjacent to the xylem. First order VBs have an outer vascular bundle
sheath, and are abaxially and adaxially interrupted by transitional parenchyma or
sclerenchyma girders; and have an inner sheath complete around the VBs (Fig. 2). Walls
of cells of the inner sheath in contact with phloem are more thickened than the other
sheath cell walls, the thickening in “U” or “O” form. Second order vascular bundles are
elliptic, with the outer sheath complete around the VBs or abaxially interrupted only by
sclerenchyma; the inner sheath is complete. I hird order VBs are ciicular, with complete
outer and inner sheaths. No extensions of the bundle sheath were seen.

Austrostipa aristiglumis, a lowland Stipoid species

Telopea 11(2): 2006

165

Leaf margins are acute in TS, with sclerenchyma or transitional parenchyma in the
margins (Figs 1, 2).

The epidermal cells in TS are smaller and thicker-walled over the VBs. Bulliform cells
occur between all VBs, at the same level or slightly above the level of the other epidermal
cells (Fig. 2). Epidermal hairs are absent.

Sheath anatomy in transection

The outline of the transection is permanently folded. The upper surface is flat; the
lower surface is flat or ribs and furrows are slightly developed. There are more than 6
VBs in TS, alternating l sl , 2 nd and 3 rd order, all located at the same level, in the centre of
the TS or towards the lower surface (Figs 1, 2). No colourless parenchyma is present in
the TS. Abaxially there is fragmented sclerenchyma below the epidermis, exceptionally
also adaxially. Sclerenchyma or transitional parenchyma is associated with all types of
VBs. Aerenchyma is present between all VBs, in lacunae, situated in the centre of TS, or
closer to the abaxial surface occupying 40 -70% of the depth of TS, sometimes >70%.

Fig. 2. Photomicrographs of Austrostipa aristiglumis. a, TS below the ligule region, showing
air cavities developed on a leaf under 11 mm saturated soil treatment (35x); b, half of a TS
below the ligule region, with no air cavities developed on a leaf under 0 mm saturated soil
treatment (35x); c, TS of leaf showing bigger inter-cellular air spaces (arrows) in the first stage
development of an air space cavity (70x); d, TS of leaf showing remains of cell walls from cells
that have disintegrated to form an air cavity (lOOx); e, TS of sheath beneath the ligule region of
a leaf under 86 mm saturated soil treatment (35x); f, detail of previous transection (70x).

166

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Arriaga and Jacobs

First order VBs are ovate, with a strand of phloem adjacent to the xylem; the outer
vascular bundle sheath abaxially is interrupted by sclerenchyma girders; inner vascular
bundle sheaths are complete around the bundle. Second order VBs are elliptical, with
an outer VB sheath abaxially interrupted by sclerenchyma girders, the inner sheath
complete around the VB. Third order VBs are circular, with an outer sheath abaxially
interrupted by sclerenchyma girders, and the inner sheath complete around the VBs.

No extensions of the bundle sheaths were seen (Fig. 2). Walls of the outer sheath cells
are not thickened, walls of the inner sheath cells are “O” thickened, cells in contact
with the phloem have thicker walls. Margins are acute in TS, with sclerenchyma or
transitional parenchyma only in the margin.

Epidermal cells over the VBs are smaller and thicker in transverse section than the
epidermal cells between. Epidermal hairs are absent.Tangential external walls of
epidermal cells are thicker than internal walls. Stomata are in both epidermes or only
in the adaxial epidermis.

The air canals have a lysigenous origin in both the leaf and the sheath. Close to the
growing points, some cells separate from adjacents cells, developing larger spaces.
Afterwards some cell walls begin to disintegrate and form cavities resulting in the
building of big air channels along the sheath and up to the middle of the leaf. Remains
of the cell walls can always be seen inside the air channels.

Discussion

Renvoize (1985) described the outstanding feature of the tribe Stipeae as the adaptation
by many genera to dry habitats, reflected in the often xeromorphic nature of the leaf
blade morphology and anatomy. Among the species listed at the end of his work is
Trikerciin hookeri (Stapf) Bor but there is no mention of hydromorphic characters,
despite Trikemia being described as living by water (Clayton & Renvoize 1986), often
on hummocks in bogs or swamps.

Austrostipa aristiglumis, on the other hand, is a species mostly growing on intermittently-
flooded inland floodplains in semi-arid and sub-humid zones. These soils are dry most
of the time.

Sculthorpe (1967) and Font-Quer (1970) described as amphibious those species that
may live in dry soils flooded during a period of the year. One theory suggests that
different groups or clades of angiosperms have retained different levels of vegetative
plasticity that is manifiest in varying abilities to adapt to aquatic habitats (Arberl920;
Sainty & Jacobs 1982; Jacobs & Wilson 1996). Amphibious species provide, in a
structural and an ecological sense, a gradual transition between truly terrestrial and
truly aquatic species (Sculthorpe, l.c.).

One characteristic of hydrophytous or amphibious species is the presence of air cavities
that develop early in the different organs of the plant, while their number and size
varies with the age and nature of the organ (Sculthorpe, l.c.).

The hydromorphic characters, stimulated by increasing soil saturation, presumably are
an adaptation that allows A. aristiglumis to maximise its growth when water is available,
even in “excess”.

Austrostipa aristiglumis, a lowland Stipoid species

Telopea 11(2): 2006

167

Aliscioni (2000) and Fabbri et al. (2006 in press) in their studies of the anatomy of
the genus Paspdlum , concluded that, under wet substrate conditions, species developed
air cavities and aerenchyma and that this ability to develop air cavities represents a
constant character. Aliscioni (2002) considered that this plasticity should be considered
in systematic and phylogenetic analysis. We agree with that assessment, especially as A.
aristiglumis represents the only known Stipoid species with hydromorphic plasticity, a
characteristic that may have value in phylogenetic considerations.

Representatives of the Stipeae have been included in numerous molecular phylogenetic
studies of grasses (Hsiao et al. 1999; Jacobs et al. 2000; Jacobs et al. 2006 in press);
all show it to be a basal lineage within the Pooidcae. Studies of the Stipeae molecular
phylogeny (Jacobs et al. 2006 in press) support recognition of Austrostipa as a well-
differentiated clade that includes A. aristiglumis. All the other species in that clade
are adapted to live in xeric and semi-mesic habitats. Only A. aristiglumis has been
observed to have the genetic adaptability and vegetative plasticity to capitalise on the
intermittently-flooded habitat.

Conclusions

1) In Austrostipa aristiglumis the sheath structure in all cases (soil with normal water
content or saturated soils) always has channels with aerenchyma; the channels increase
in size with increasing saturation of the soil.

2) In Austrostipa aristiglumis the leaves show channels and aerenchyma in the lower
part of the laminae, near the meristem and only in those samples living on the more
saturated soils.

In any case, the channels and aerenchyma decreaeses and are completely lost towards
the middle of the lamina.

3) In Austrostipa aristiglumis leaf and sheath air cavities have a lysigenous origin.

4) Most stipoid grasses have morphological and anatomical adaptations to mesic and
xeric environments but Austrostipa aristiglumis is an unusual case of a stipoid grass
adapted to intermittent flooding in inland south-eastern Australia, representing an
amphibious plant.

Acknowledgments

We wish to thank Lie. Monica Stampacchio and Mrs Maria Dolores Montero for
technical assistance.

References

Aliscioni SS (2000) Anatomia ecologica de algunas especies del genero Paspalutn (Poaceae-
Panicoideae, Paniceae). Darwiniana 38 (3 -4): 187 -207.

Aliscioni SS (2002) Contribution a la filogenia del genero Paspalutn (Paniceae-Poaceae). Annals
of the Missouri Botanical Gardens 89: 504 -523.

Arber A (1920) Water plants, a study of aquatic angiosperms. (Cambridge University Press:
Cambridge)

168

Telopea 11(2): 2006

Arriaga and Jacobs

Arriaga MO (1982) Presencia dc fibras en epidermis de Stipa (Gramineae). Boletin de la Socicdad
Argentina de Botdnica 21 (1-4): 143-151.

Arriaga MO (1983) Anatomia foliar de las especies de Stipa del subgenero Pappostipa (Stipeae-
Gramineae) de Argentina. Revista del Museo Argentina de Ciencias Naturalcs. (Botdnica) VI
(4): 89-141.

Arriaga MO 8c Barkworth ME (2000) Leaf anatomy in Nassella and other South American
stipoid grasses. American Journal of Botany 87 (6): 33.

Barkworth ME (1981) Foliar epidermesand taxonomy of North American Stipeae (Gramineae).
Systematic Botany 6:136-152.

Barkworth ME 8c Everett J (1987) Evolution in the Stipeae: identification and relationships of
its monophyletic taxa. Pp. 251-264 in Soderstrom TR, Hilu KW, Campbell CS, & Barkworth
ME (eds) Grass systematics and evolution. (Smithsonian Institution: Washington, D.C.)

Barkworth ME 8c Torres MA (2001) Distribution and diagnostic characters of Nassella (Poaceae:
Stipeae). Taxon 50: 439-468.

Brown W (1958) Leaf anatomy in grass systematics. Botanical Gazette 119 (3): 170-178.

Caro JA 8c Sanchez E (1971) La identidad de Stipa brachychaeta Godron, S. caudata Tr. y S. ber-
trandii Phil. Darwiniatm 16 (3-4): 637-653.

Caro JA 8c Sanchez E (1973). Las especies de Stipa (Gramineae) del subgenero Jarava. Kurtziana
7:61-116.

Cialdella AM 8c Arriaga MO (1998) Revision de las especies sudamericanas del genero Pipto-
chaetium (Poaceae, Pooideae, Stipeae). Darwiniana 36 (4): 107-157.

Cialdella MA and Giussani LM (2002) Phylogenetic relationships of the genus Piptochaetium
(Poaceae, Pooideae, Stipeae): evidence from morphological data. Annals of the Missouri
Botanic Gardens 89 (3): 305-336.

Clayton WD 8c Renvoize SA (1986) Genera Graminum: grasses of the World. Kew Bulletin
Additional Series XII. (H.M. Stationary Office: London)

Clifford HT 8c Watson L (1977) Identifying grasses: data, methods and illustrations. (Queensland
University Press: Brisbane).

Cutler DF (1978) Applied plant anatomy. (Longmans: London)

Fabbri LT, Rua GH 8c Bortoloni N (2006 in press). Different patterns of aerenchyma formation
in two hygrophytic species of Paspalum (Poaceae) as response to flooding. Flora 200 (4).

Font-Quer P (1970) Diccionario de Botdnica. (Labor: Barcelona)

Freitag LI (1985) The genus Stipa in southwest and south Asia. Notes from the Royal Botanic
Gardens Edinburgh 42:355-489.

Hsiao C, Jacobs SWL, Chatterton NJ 8c Assay KH (1999) A molecular phylogeny of the grass
family (Poaceae) based on the sequences of nuclear ribosontal DNA (ITS). Australian
Systematic Botany 11: 667-688.

Jacobs SWL, Everett J, Connor HE 8c Edgar E (1989) Stipoid grasses in New Zealand. New
Zealand Journal of Botany 27: 569-582.

Jacobs SWL and Everett J (1996) New names for Australasian species formerly included in Stipa.
Telopea 6: 579-595.

Jacobs SWL, Everett J, Barkworth ME 8c Hsiao C (2000). Relationships within the stipoid grasses
(Gramineae). Pp. 75-82 in Jacobs SWL 8c Everett J (eds) Grasses, systematics and evolution.
(CSIRO: Melbourne)

Jacobs SWL 8c Wilson KL (1996) Biogeographical analysis of the freshwater plants of Australasia.
Australian Systematic Botany 9 (2): 169-183.

Jacobs SWL, Bayer R, Everett J, Arriaga MO, Barkworth ME, Sabin-Badereau A, Torres MA,
Vasquez F 8c Bagnall N (2006 in press) Systematics of the tribe stipeae (Gramineae) using
molecular data. 4"’ International Grass Symposium, Rancho Santa Anna.

Martinovsky JO (1976). Neue Stipa- Sippen und einige Erganzungen der fruiter beschriebenen
Stipa- Taxa. Preslia 48: 186-188.

Matthei OR (1965) Estudio critico de las gramineas del genero Stipa en Chile. Gayana Botdnica
(13): 1-137.

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Telopea 11(2): 2006

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Parodi L (1944) Revision de las gramineas australes americanas del genero Piptochaetium. Rc-
vistn del Museo de La Plata, section Botdnica 6: 213-310.

Parodi L & Freier F (1945) Observaciones taxonomicas sobre las gramineas estipeas. Ciencia e
Investigation 1:144-146.

Penailillo P (1996) Anatherostipa un nuevo genero de Poaceae. Guyana Botdnica 53: 277-284.

Reeder JR & Reeder CG (1968) Parodiella a new genus of grasses from the high Andes. Boletin de
la Sociedad Argentina de Botdnica 12: 268-283.

Renvoize SA (1985) A survey of leaf-blade anatomy in grasses: VI. Stipeae. A survey of leaf blade
anatomy in grasses Vll. Pommereulleae, Orcuttieae and Pappophoreae. Kew Bulletin 40:
731-744.

Roig FA (1964a) Las gramineas mcndocinas del genero Stipa I. Taxonomia. Revista de la Facultad
de Ciettcias Agrarias, Mendoza 11 (1-2): 3-110, lams. 1-26.

Roig FA (1964b) Las gramineas mcndocinas del genero Stipa IV. Nova Addenda. Deserta 2:
107-114.

Roig FA (1965) Las gramineas mendocinas del genero Stipa 11. Addenda taxonomica. Revista de
la Facultad de Ciencias Agrarias, Mendoza 12 (1): 79-94.

Rojas PF (1997) [1998] Nuevas especies y nuevas combinaciones para la tribu Stipeae (Poaceae)
en Bolivia. Guyana Botdnica 54: 163-182.

Sainty GR & Jacobs SWL (1982) Waterplants of New South Wales. (Water Resources Commission,
New South Wales: Sydney)

Scholz H (1982) Uber Mikro- und Makrohaare eininger Piptatherutn und Stipa-Arten (Stipeae,
Gramineae). Willdenowia 12: 235-240.

Sculthorpe CD (1967) The Biology of Aquatic Vascular Plants. (Edward Arnold: London)

Thomasson JR (1978) Epidermal patterns of the lemma in some fosil and living grasses and their
phylogenetic significance. Science 199:975-977.

Torres MA (1993) Revision del genero Stipa (Poaceae) en la Provincia de Buenos Aires. Comiswn
de Investigaciones Cientificas de la Provincia de Buenos Aires, Monografta 12.

Torres MA (1997a) Nassella (Gramineae) del noroeste dela Argentina. Comisidn de Investigaciones
Cientificas de la Provincia de Buenos Aires, Monografta 13: 1-42.

Torres MA (1997b) Stipa (Gramineae) del noroeste de la Argentina. Comisidn de Investigaciones
Cientificas de la Provincia de Buenos Aires, Monografta 13: 47-64.

Torres MA (1997c) Nicoraclla (Gramineae) un nuevo genero para America del Sur. Comisidn de
Investigaciones Cientificas de la Provincia de Buenos Aires, Monografta 13: 69-76.

Vazquez FM (1992) Estudio biosistematico del genero Stipa L. en la Peninsula lberica e Islas
Baleares. PhD Thesis. Inedit. Extremadura University. Badajoz, Spain.

Vazquez FM & Barkworth ME (2004) Resurrection and emendation of Macrochloa (Gramineae:
Stipeae). Botanical Journal of the Linnean Society 144: 483-495.

Vazquez FM and Devesa JA (1996) Revision del genero Stipa L. y Nassella Desv. (Poaceae) en la
Peninsula lberica e Islas Baleares. Acta Botanica Malacitana 21:125-189.

Vazquez FM & Devesa JA (1997) Two new species and combination of Stipa L. (Gramineae)
from northwest Africa. Botanical Journal of the Linnean Society 124: 201-209.

Vickery JW, Jacobs SWL 8< Everett J (1986) Taxonomic studies in Stipa (Poaceae) in Australia.
Telopea 3: 1-132.

Watson L 8c Dallwitz MJ (1981) An automated data bank for grass genera. Taxon 30: 424-429.

Watson L & Dallwitz MJ (1992) The grass genera of the World. (CAB International: Wallingford,
UK)

Manuscript received 24 March 2005, accepted 7 February 2006

Telopea 11(2) 171-194

New taxa and combinations for Oedogonium
and Bulbochaete (Oedogoniales,
Chlorophyceae) in Australia

Stephen Skinner and Timothy J. Entwisle

National Herbarium of New South Wales, Mrs Macquaries Road, Sydney NSW 2000 Australia

Abstract

Fifteen new taxa in the Oedogoniaceae, the species Bulbochaete kakaduettsis [ for B. clatior sensu
Scott & Prescott (1958) ] B. rodwayi , Oedogonium Bancroftii, Oe. capricornicutn, Oe.cerasinum ,
Oe. cribbianum, Oc. hardyi [for Oe. tnonile var. victoriense G.S.West] Oe. tnoebiusii
[for Oc. undulation var. nwbiusii Schmidle] Oe. pilbaranum and Oe. utrarium, and the
varieties B. calospora var. murrayana, B. gigantea var. glabra, B. wuhanensis var. spechtii,
Oe. kwangsiense var. playfairii and Oe. starmachii var weetalibahense are described. Two other
taxa are new combinations: Oe. undulatum var. wissntanii for Oe. wissmanii Stephen Skinner
and Oe. uleanum var. continuum for Oe. platygynum var. continuum Nordstedt ex Hirn.

Introduction

Seventeen taxa (8 species and 4 varieties of Oedogonium , 2 species and 3 varieties of
Bulbochaete) require taxonomic review in preparation foran account of the Oedogoniales
(Chlorophyta) for the Algae of Australia series. Most are new taxa and require formal
taxonomic recognition. A few require revision of status, although previously described.
The taxa are arranged in systematic order following Mroziriska (1991,2000).

The earliest mention of Oedogonium from Australia in the literature is Sonder
(1852) but without description or illustration. Next comes Berkeley and Harvey’s
(1860) enthusiastic description of Oe. monile. It was Baron von Mueller who laid
the Australian foundation for freshwater phycology generally and whose enthusiasm
sparked the first local interest in the Oedogoniales. Mueller encouraged people in all
parts of the continent to send him curious and unexpected plants, and so received
quite a few freshwater algal samples including numerous Oedogoniales. F.M. Bailey in
Brisbane made a major contribution in the late 19"' and early 20"' Century. All three
Bailey papers (1893, 1895, 1898) include Oedogonium and Bulbochaete species.

Hint’s (1900) world monograph of the Oedogoniales, the starting point for
nomenclatural priority in this order, used collections from the herbaria of Mueller and
Bailey, sent to V.B. Wittrock, M. Mobius, W. Schmidle and C.F.O. Nordstedt. Although
Hirn did not designate types as such, there are ten taxa based solely on Australian

© 2006 Royal Botanic Gardens and Domain Trust

ISSN0312-9764

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collections. Four of these (and also numerous new desmid taxa) come from a single
collection by T. Gulliver made ‘between the Norman and Gilbert Rivers’ in 1874. We
have been able to find and view this collection, and some others from Australia, in LD,
but not all Hirn ‘types’ could located. In the absence of voucher material, we have relied
on protologues and subsequent monographs and floras to define existing taxa.

Further records of Oedogoniales from parts of Australia can be found in Scott and
Prescott (1958) for Arnhem Land, Skinner (1980) for the New England Tableland of
New South Wales, and the many contributions of Cribb to the Queensland Naturalist
(see Day et al. 1995 for a list). Cribb (1956) provides the only report of Oedocladium
in Australia.

This paper uses material from historical collections in AD, MEL and BRI, as well as
recent collections forwarded to us from collectors across the continent and our own
recent collections.

Methods

Recent collections were fixed in either 4% formaldehyde or 70% ethanol in the field,
and sent to the National Herbarium of New South Wales where part of each sample was
mounted in 40% Karo as semi-permanent slides, while the remainder of the specimen
was stored in 70% ethanol and 5% glycerine. Slides were examined with a Leitz Dialux
compound microscope and drawings made with a drawing tube. Collections from
MEL, AD and BRI were also examined. Data sheets for each collection are also retained
at NSW.

Oedogonium Link

Mrozinska (1991) defined 20 informal groups for Oedogonium, based on phylogenetic
analysis. Later she (Mrozinska 1999) presented a new subgeneric classification and
allocated her informal groups to this new infrageneric system, which we follow here.
Mrozinska (pers. comm.) kindly provided us with a list of how she has allocated taxa
within this new system.

Subgenus Archaeoedogotiium Mrozinska (1999)

Cell walls, possibly excepting antheridia, helically striped with finely granulate or
punctate outer surface; basal cell globose or depressed globose, vertically plicate.
Antheridia either intercalary with single spermatozoids or in globose, single-celled
dwarf males.

Group A

Macrandrous taxa.

Oedogonium cribbianum Stephen Skinner & Entwisle sp. nov.

Oe. punctato-striatum sensu Cribb, Lake Broadwater. The Natural History of an Inland
Lake and its Environs: 39 (1988)

Diagnosis: Oedogonio fasciculato simile, sed macrandro et oosporis globosis depressis
dense verrucosisque.

Oedogonium & Bulbochaete (Oedogoniales, Chlorophyceae) Telopea 11(2): 2006

173

Fig. 1. Oedogonium cribbinnum: a,basal cell; b, intercalary antheridia; c, oogonium with oospore;
d, enlargement of oospore, with sketch of sculpturing (Cribb 1090.4 ); Oe. utmrium: e,f, basal
cells; g, antheridia; h, solitary oogonia, with oospores; i, pair of developing oogonia (Towler 283c
& Wilson s.n.). (Scales = 20 pm)

Type: Queensland: Surveyors Gully, Lake Broadwater (27° 21’S; 151° 06’E), Cribb
1090.4, 21 Feb 1987 (holo BRI).

Illustration: Fig. l,a-d.

Macrandrous, monoecious or dioecious. Vegetative cells capitellate, finely helically
punctate, L/D (3—)5—10, (14-) 16-18 pm diam., chloroplast reticulate, few pyrenoids;
basal cell hemispherical, vertically plicate, 12-15 pm long, 22-23 pm diam.; terminal cell
shallowly domed. Oogonium solitary, globose to depressed globose, punctate, intercalary
or occasionally terminal (and then with a domed top), circumcision median, wide at
one side, 34-41 pm long, 40-44(-46) pm diam.; oospore depressed globose, not filling
oogonium, golden to reddish golden, exospore banded verrucose, 29-32 pm long,
35-38 pm diam.; suffultory cell as vegetative cells, frequently L/D 3-4. Antheridia solitary

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Skinner and Entwisle

to triple, short (L/D c.1.5) same width as surrounding vegetative cells, top opening, in
groups with vegetative cells between, either several cells above or below an oogonium
or on separate filaments, with one spermatozoid.

Etymology: named in honour of Professor Alan B. Cribb who has done so much to
alert people to the fascination of freshwater algae.

Distribution: known from type locality in Queensland, on Vallisneria sp. and northern
New South Wales on Myriophyllum sp.

Very similar to Oe. fasciculum Zhu (1964) from China but differs in not being
nannandrous, and in having slightly more depressed oospores with more regular and
dense cover of verrucae. Cribb’s (1988) determination of Oe. punctato-striatum de Bary
ex Hirn is superficially similar to the type, but that taxon has smooth oospores.

Specimens examined: Queensland: Surveyors Gully, Lake Broadwater, Cribb 1090 subsamples,
21 Feb 1987 (BRI). New South Wales: North Western Slopes: quarry, 1 km W of township, How¬
ell Rd, Tingha, Skinner 0801, McPherson & Towler, 13 Oct 2004 (NSW).

Subgenus Prooedogonium Mrozinska (1999)

Cell walls smooth; basal cells globose to depressed globose, hemispherical or if vase¬
like then a short goblet, L/D no more than 3. Antheridia with single spermatozoid,
macrandrous or nannandrous, and then internal.

Section C

Macrandrous; basal cell hemispherical, occasionally short vase-like; oogonium usually
depressed globose, oospore smooth walled.

Oedogonimn utrarium Stephen Skinner & Entwisle sp. nov.

Diagnosis: species bene distincta, cellula suffultoria reflexa, oogonio asymmetrico
gleboso utriculoide, oospora quoque biconica sulcoque medio.

Type: Western Australia: pool in bed of tributary of Hammersley R., Fitzgerald River
Nat. Pk (33° 46’S; 119° 43’E), Towler 283c & Wilson , 10 Aug 2003 (holo NSW).

Illustration: Fig. l,e—i.

Macrandrous, dioecious. Vegetative cells narrow cylindrical, L/D 4-6,2.5-3.5 pm diam.;
basal cell irregular depressed globose, 4-5.5 pm high, 5-5.5 pm diam.; terminal cell
truncated. Oogonium solitary or paired, irregularly saccate,, 23-28 pm long, 10-12 pm
diam., circumcision median, wide; oospore irregular bi-conical, with median furrow,
wall smooth but lumpy, not filling the chamber but forcing the circumcision open,
14-19 pm long, 10-12 pm diam.; suffultory cell as vegetative cell but reflexed, so that the
filament is curled through 90-180°. Antheridia solitary or paired, terminal on separate
filament, 5.5 pm long, 2-3 pm diam., single spermatozoid.

Etymology: from utrarius (Latin) a water-bag carrier in the Roman army. These water-
bags were frequently made from whole goat or other animal skins, like wineskins, and
the oogonia of this taxon are similar in shape.

Distribution: known only from the type locality, Western Australia, in a deep riverine
pool on granite with aquatic grass, apparently forming a greenish layer close to the
gravel.

Oedogonium & Bulbochaete (Oedogoniales, Chlorophyceae) Telopea 11 (2): 2006

175

Fig. 2. Oedogonium cerasinum. a, individual antheridial, vegetative and oogonial thalli (Cherry
446); Oe. pUbaranum. b, vegetative filament; c, antheridia; d, oogonium; e, oogonium with
mature oospore (Edinger 3516); Oe. uleanum var. continuum-, f, terminal series of oogonia; g, top
view of oogonium; h, intercalary series of oogonia (Cribb 496.9). (Scales = 20 pm)

While sharing vegetative characters with the rest of Group C, this taxon is distinctive in
possessing the bent suffultory cell and the lumpy, irregular, wineskin-shaped oogonium
as well as the bi-conical oospore with a median furrow. There are no other taxa with
similar oogonia.

Oedogonium cerasinum Stephen Skinner & Entwisle sp. nov.

Diagnosis: Oedogonio fretnyi simile sed oogonio pyriformi angulari et cellularibus
vegetativis capitellatibus celluloque basi hemispherico.

Type: Western Australia: Brixton Street wetland, Kenwick, Perth (32° 22’S; 115° 56’E),
Cherry 446, 9 Sept 2002 (holo NSW).

Illustration: Fig. 2, a.

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Skinner and Entwisle

Dioecious, macrandrous. Vegetative cells capitellate, 15-23(-26) pm long, (3—)6-6.5
pm diam., three to seven or eight in a whole plant; basal cell shallow hemispherical,
3-5 pm high, 35-50 pm diam.; both male and female plants terminate in a very long,
(up to three or four times the length of the rest of the thallus) fine seta. Oogonium
pyriform to angular pyriform, 15-21 pm long, 17-20 pm diam., often acropetal, opening
by a supreme hinged pore (or partial circumcision); oospore closefitting oogonium,
pyriform to angular pyriform, wall smooth, thick, laminated, mesospore yellowish to
cherry red, 14-20.5 pm long, c. 17-20 pm diam. Antheridia paired, below seta on male
plant, chambers 6-8.5 pm long, 6 pm diam, single spermatozoid.

Etymology: as the oogonia are cherry-shaped, the mesospore cherry red in colour and
the collector was Wayne Cherry, cerasinum (Latin) for cherry-like seems appropriate.

Distribution: known only from the type locality, Western Australia, where it is epiphytic
on fine, terete stems of water plants in both still and moving water.

Oedigonium cerasinum is one of a number of species with few-celled filaments,
including Oe. mirpurense Islam, Oe.fremyi Gauth.-Lievre, Oe. nanum Wittr. ex Hirn,
Oe. capitellatum Wittr. ex Hirn, Oe. sphaerandrium Wittr. & Lundell ex Hirn, and
Oe./o7tgipi/uwJao.Oe.cerasimmiisdistinctiveinthefollowingcombinationofcharacters:
dioecious, distinctly angular pyriform oogonia, and a shallowly hemispherical basal
cell. It is closest to Oe. longipilum and Oe.fremyi. It shares a terminal seta and supreme
circumcision with Oe. longipilum, but that species has a short, vase-shaped basal cell,
and ovoid oogonia, sometimes in pairs. Oe.fremyi has a supramedian circumcision, the
oogonium is globose, the basal cell is diffuse and the vegetative cells are less consistently
capitellate than Oe. cerasinum.

Group G

Macrandrous; basal cell shortly vase-like; oogonium opening through a median to
superior pore, or pore and slit; oospore smooth.

Oedogotiium pilbaranum Stephen Skinner & Entwisle sp. nov.

Diagnosis: Oedogonio rufescente simile sed in dimensiones minore atque circumcissione
oogonii rostrato.

Type: Western Australia: 28 km SSW of Turee Creek Station (23° 52’S; 118° 32’E),
Edinger 3516, 9 May 2003 (holo NSW).

Illustration: Fig. 2,b-e.

Macrandrous, dioecious. Vegetative cells cylindrical or slightly capitellate, L/D 3-10,
7-8 pm diam.; basal cell depressed globose, slightly undulate, c.10 pm tall, 13-14 pm
diam.; terminal cell ending in a shallow cap. Oogonium solitary or paired, depressed
globose to spindle-like, 20-26 pm long, 20-23 (-25) pm diam., opening a supramedian
pore or short slit; oospore depressed globose, smooth walled, filling chamber, mesospore
brown, 18-19 pm long, 20 pm diam.; suffultory cell as vegetative cells. Antheridia
intercalary or acropetal, in pairs, 7-10 pm long, c.7 pm diam.; single spermatozoid.

Etymology: from Pilbara, the region of Western Australia from which the sole collection
was made.

Oedogonium & Bulbochaete (Oedogoniales, Chlorophyceae) Telopea 11(2): 2006

177

Distribution: known only from the type locality, Western Australia, in a permanent
seepage.

Although very close to the cosmopolitan Oe. rufescens Wittr. ex Hirn, this specimen
shows a globose basal cell, more similar to those of Oc. inversum Wittr. ex Hirn and Oe.
infirmum Tiffany, and the oospore is more nearly globose, not tending to be laterally
ovoid/elliptical. The oospore fills the chamber and conforms to its shape to some extent,
exhibiting a slight dome below the terminal dome of the oogonial chamber.

Subgenus Oedogonium Mrozinska (1999)

Cell walls smooth; basal cell short to elongate vase-shaped, usually with a basal disc
which may be clawed.

Supersection Monospermatozoidiae

Antheridia with single spermatozoid; in nannandrous species either internal in goblet¬
like dwarf male, or extruded from the body cell, with diffuse antheridial walls.

Group E

Macrandrous or nannandrous; dwarf males with internal antheridia; basal cell shortly
vaselike; oogonia with median whorl of projections or ribs.

Oedogonium uleanum Hirn, Acta Soc. Sci. Fenn. 27: 311 (1900).var. continuum
(Nordstedl) Stephen Skinner & Entwisle comb.nov.

Oedogonium platygynum var. continuum Nordstedt, Bot. Not.: 153 (1887).

Oc. platygynum var. continuum Nordstedt ex Hirn (1900).

Type: New Zealand: Ohaeawai, (35° 21’S; 173° 53’E), Berggren(64): Herb Nordstedt,
(LD n.v.).

Illustration: Nordstedt op cit: PI. 1, figs 16, 17; hie Fig. 2, f-h.

Dioecious, macrandrous. Vegetative cells cylindrical to slightly capitcllate, L/D
2.5-3.5, 7-8 pm diam.; basal cell not seen; terminal cell obtusely capped. Oogonia
solitary, in pairs, threes, fours (common) or fives, depressed globose, 13.5-23 pm long,
18-25 pm diam., flanges prominent rounded mamillatc, circumcision submedian;
oospore depressed globose to spherical, not filling the chamber, 14—18 pm diam.;
suffultory cells similar to vegetative cells. Androsporangia , dwarf males or antheridia not
observed.

Distribution: Queensland in seepage on upper shelf near entrance of cave, and in New
Zealand.

The Queensland and New Zealand collections are of the same taxon. Nordstedt
(1887, 1888) described the New Zealand taxon as a variety of Oe. platygynum Wittr.
ex Hirn. However Oe. platygynum var. platygynum is described as nannandrous and
figured with dwarf males, and the flanges are rounded. Nordstedt (1887) contains
the six word protologue for Oe. platygynum var. continuum Nordstedt, while the
illustrations, including the antheridia/androsporangia, are in Nordstedt (1888).This
taxon is clearly a macrandrous taxon, and the flanges are often slightly truncated.
Although the oogonia and oospores are rather smaller than Oc. uleanum as described

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Skinner and Entwisle

in Hirn (1900) (oogonia 28-32 pm diam., 21-26 pm long) the Queensland and New
Zealand collections have oogonia in series and otherwise fit closer to Oe. uleanum
than Oe. platygynum. Interestingly Hirn (1900, p. 279) queried the position of
Oe platygynum var. continuum and expressed the desire for more information. “Fur die
Var. continuum, die leider noch wenig bekannt ist, sind die reihenstandigen Oogonien
und die abgestutzte Form der Oogoniumvorspriinge hervortretende Merkmale.” [As
for the var. continuum, unfortunately we do not know as yet if having oogonia in series
and truncated oogonial wall flanges are characteristic enough to make it distinct.] This
record from Queensland was not included in Cribb (1965).

Specimens examined: Queensland: Paradise Cave, Noosa Heads, Cribb 496.9, 13 Oct 1962
(BRI).

Group P

Nannandrous; dwarf males with a basal cell and internal antheridia later extruded;
vegetative cells may have undulate walls; oogonia with inframedian to supramedian
circumcision; oospore smooth or variously sculptured.

When establishing this group Mrozinska (1991) allowed only Oe. undulatum and
Oe. croasdaleae Jao as examples, and later listed Oe. undulatum var. undulatum
f. undulatum and f. senegalense, Oe. undulatum var. americanum Transeau f. americanum
and f. serriatum (Prescott) Mrozinska, Oe. mirificum Zhu and var. minus Jao,
Oe. croasdaleae and f. goczalkowicensis Mroz. Webb in the group (Mrozinska, pers.
comm.). We have adopted a much wider definition here, based on comparison of the
dwarf male structures in Oe. undulatum with those in Oe. Ilians Nordstcdt & Hirn in
Hirn, Oe. monile and similar taxa, where the antheridial cell appears to be extruded from
the body cell and to contain only a single spermatozoid at a time. The fine structure of
dwarf males of members of this group requires examination similar to Leonardi et al.
(1998) for Oe. pluviale Nordst. ex Hirn.

Oedogonium moebiusii Stephen Skinner & Entwisle sp. nov.

Oedogonium undulatum var mobilis'd Schmidle, Flora 82: 297, fig. 1 (1896). Type:
Burpengary, Bancroft', 1894?: Herb. Schmidle (B probably lost, n.v.).

Diagnosis: a Oedogonio undulato parietium tholis concavis in ambobus cellulibus
vegetabilis androsporangiisque differt.

Type: Queensland: Peregian Beach (26 30’S; 153° 06’E), Peberdy s.n., Sept 1973: (holo
BRI).

Illustration: Fig. 3, a-d.

Nannandrous, gyn- or more frequently idioandrosporous. Vegetative cells undulate
with 4 invaginations, 3 complete, 2 short, flat to concave vaults, L/D 3-4(-5),
15-22 pm diam.; basal cell vase-shaped, smooth walled; terminal cell with domed cap
(sometimes as terminal oogonium). Oogonium spherical, inframedian hinged pore
becoming circumcision, solitary, paired or up to 6 in a row, 46-75(-87) pm long,
44-64(—75) pm diam.; oospore spherical, 44-58(-67) pm diam., thick walled, lamellate
and occasionally with faint denticulations below outer wall, chinese-red mesospore,
almost filling oogonium,; suffultory cell similar to vegetative cells, but at the broad end

Oedogonium & Bulbochaete (Oedogoniales, Chlorophyceae) Telopea 11(2): 2006

179

Fig. 3. Oedogonium tnoebiusii: a, basal cell; b, androsporangium; c, series of oogonia with
oospores and dwarf male (Peberdy s.n. BRI704656); Oe. hardyi: d, new thallus with basal
cell and terminal cap; e, intercalary androsporangia; f, oogonium with oospore and dwarf
male; g, mature oospore (Towlcr 283a & Wilson ); Oe. kwangsiense var playfairii: h, basal cell;
i,j, androsporangia; k, terminal oogonium; 1, series of oogonia with oospores (Cherry 447);
Oe. capriconicum: m, basal cell; n, intercalary antheridia; o, oogonium with oospore (Reid
270103.04). (Scales = 20 pm)

of sizes, 19-23 pm diam.. Androsporangia undulate walled, 1 —3(—4), often below the
groups of oogonia, c. 15 pm long, c. 12 pm diam.; dwarf male on suffultory cell, curved
vase-shaped body cell, 28-44 pm long, 8-12 pm diam., upper internal antheridium,
6-9 pm long, 6 pm diam. (body cell as lower antheridium), single spermatozoid.

Distribution: fertile material from Western Australia, South Australia, New South
Wales and Queensland, with vegetative material noted for Victoria, in still or slow
flowing habitats; Argentina, probably widespread (sometimes reported as f. incision
I-Iansg. or f. senegalense Nordst.).

Schmidle’s iconotype is of a poorly sketched vegetative fragment, and Bancroft’s
specimen appears to have been lost. We have used the Schmidle epithet to describe
a new species rather than make a new combination based on inadequate material.

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Peberdys.n. has all reproductive structures, and is from coastal south-eastern Queensland
like the Bancroft collection. Hirn (1900), in a long and strongly worded discussion of
the variation in Oe. undulatum sens, lat., insists that he found no consistency in the
vaulting of undulations of cells even within the same filament. There is no distinction
made between 'integris’ and ‘ repandis ’ cell kinds in the Hirn (1900) protologue until you
get to the various forms: ‘integris’ referring to those forms with the evenly undulate wall
outline, and ‘repandis (= bent back)’ to those with the concave wavetops. Formae 8 and
£ of Oe. undulatum and the subforma below Oe. undulatum f. senegalense (Nordstedt)
Hirn all show what Hirn called repand waves. Hirn (1900) quotes Hansgirg as being of
the same opinion, even though forma (3 incisum Hansg. is Hansgirg’s (1886) published
interpretation, and he was confident enough to distribute exsiccatae under that name.
Herring (1914) illustrated reproductive material of forma £ and contended that it was
the common forma in Schleswig-Holstein. A clear illustration of both wall forms is
given in Fig. 5c in Velez (1995).

Hirn’s observation has not been the experience with Australian material examined
for this study. Communities of filaments with either smoothly undulate cells or with
concave vaulting have been the norm, all cells in any filament being similar in wall
form. Intergrading forms, if present at all, are extremely rare. There are three known
sites (Brixton St wetland, Ham Common, and Collector) where both forms of filaments
may be found in the same sample, but the cells are consistent within each filament. Velez
(1995) pointed out that culture studies of filaments of various forms of Oe. undulatum
sensu lato should clear up this problem once and for all, a sentiment with which we
concur. Preliminary results indicate that culture strains are morphologically consistent
(Velez, pers. comm.). Culture studies may also demonstrate the worth of Hirn’s other
subgeneric character in Oedogottium undulatum, number of oogonia in sequence,
which in Australian material of both Oe. moebiusii and Oe. undulatum varies widely
within any one sample. With significant vegetative and reproductive differences, there
is strong support for the distinction between these two varieties of Oe. undulatum.
Collections of sterile filaments fitting Oe. moebiusii in Australia can be separated readily
from filaments of Oe. undulatum (4 inflexions and 3 domes with 2 half-domes).

Specimens examined: Queensland: Peregian Beach, Peberdy s.n., Sep 1973 (BRI). New South
Wales: Southern Tablelands: Collector Ck, Collector, Skinner 0543, 4 Jan 2002 (NSW). Central
Coast: Centennial Park, Lucas s.n., -Sep 1915 (NSW); Lake Nadumgamba, Mt Annan Bot. Gdn,
Lings.n., 22 Oct 2001,and Coveny 18980,22 Nov 2001 (NSW). Northern Tablelands: Barleyfields
Lagoon, TSR paddock, near Uralla, Brock s.n., 6 Feb 2002 (NSW). North Western Slopes: quarry,
Howell Rd 1km W of Tingha, Skinner 0801, McPherson & Towler, 13 Oct 2004 (NSW). South
Australia: 6 km NW Beachport, towards Robe, Skinner 0556, 14 Nov 1978 (NSW); transient
pool, main pond, Piccaninnie Ponds Cons. P., Skinner 0446, Arnold & Towler, 1 Oct 2001 (NSW).
Western Australia: Fernbrook Falls, Deep R., Mt Franklin N.P., Entwisle 2392, 5 Jan 1994 (MEL);
Brixton St wetland, Kenwick, Cherry 447, 9 Sep 2002 (NSW); Lake Gillon, Gillon St, Karawarra,
Skinner 0577, Casanova, Garcia & Novis, 3 Oct 2002 (NSW).

Oedogottium undulatum (Breb.) A.Braun ex Hirn, Acta Soc. Sci. Fenti 27: 257 (1900)
var. wissmanii (Stephen Skinner) Stephen Skinner & Entwisle comb, et stat. nov.

Oedogoniutn wissmanii Stephen Skinner, Proc. Linn. Soc. N.S.W. 104: 261 (1980).

Type: New South Wales: Northern Tablelands: Bullock Ck (30° 07’S; 152° 15’E), Skinner
s.n., Dec 1974 (NSW!).

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Illustration: Skinner, op.cit.: fig. 3.3a-c.

Nannandrous, idioandrosporous. Vegetative cells evenly undulate with four grooves
and five domes, L/D 2.5-4, 15—27(—30) pm diam., few large pyrenoids; basal cell tall,
waisted vase shaped with digitoid holdfast,: terminal cell (sometimes oogonium) with
domed cap. Oogonium solitary or in series 2-6, globose, 57-88 pm long, 45-72 pm
diam., inframedian hinged circumcision; oospore globose, 56-67 pm diam., outer wall
thick, opaque, often with a pinch opposite the opening of the circumcision, mesospore
chinese-red; suffultory cell variable, from as vegetative cells to tumid, 18-28 pm diam.
Androsporangia solitary or in series 3-5, two undulations, 16-23 pm long, 11-16 pm
diam.; dwarf male narrow clavate, 45-54 pm long, 9-11 pm diam., antheridia internal,
single spermatozoid.

Distribution: Western Australia, Northern Territory, New South Wales and Queensland,
in both still and running water.

We have submerged the Skinner (1980) taxon in Oe. undulation as var. wissmanii,
for Australian material, because the undulate androsporangia which distinguish this
variety from Oe. undulatam var. undulatum, were previously mistakenly described as
antheridia in Skinner (1980). The type slides of Oe. wissmanii show 1) Oe. undulatum
var. undulatum-like vegetative cells, 2) androsporangia with the undulate walls, and
3) no dwarf males on the oogonium. More recent collections confirm the presence of
dwarf males similar to those of the type variety. The width of cells in the same filament
in wild populations may vary by a factor of two in a distance of ten or fewer cells.

Novis (2003) appears to have a similar taxon, cited as Oedogonium sp. aff. Oe. wissmanii,
from Chatham Island, New Zealand.

Specimens examined: Queensland: Sir John Chandler Park, Brisbane, Peberty s.n., 17 Jul 1964
(BRI). New South Wales: Northern Tableland: Bullock Ck, Skinner s.n., Dec 1974 (NSW). Cen¬
tral Coast: Nepean R., Yarramundi Bridge, Agnes Banks, Skinner 0335 & McPherson, 9 Aug 2001
(NSW); The Driftway, Richmond, watermeadow, Skinner 0337 & McPherson, 9 Aug 2001 (NSW);
Ham Common, Richmond, watermeadow, Skinner 0338 & McPherson, 9 Aug 2001 (NSW).
Northern Territory: Douglas R., u/s Douglas Hot Springs, Padovan & Mecalfe A129, 20 Aug 2003
(NSW). Western Australia: Brixton Street wetland, Kenwick, Cherry 447, 9 Sep 2002 (NSW).

Oedogonium liardyi Stephen Skinner & Entwisle sp. nov.

Oe. monile Berkeley & Harvey ex Hirn f. victoriense G.S.West, /. Linn. Soc. 39:45 (1909).
Type: Victoria: Yan Yean Reservoir, macrophytic benthos, A.D. Hardy s.n., 2 Dec 1905
(BM, reported dried out 2004).

Diagnosis: Aut grande aut grandiore Oedogonio monile var. eminente , sed circumcisione
oogonii medio, strato medio exosporae stellato spiraleque, stipa nannandrii longissima
et gracile.

Type: Western Australia: tributary Hammersley R., Fitzgerald River Nat. Pk (33° 46’S;
119° 42’E), Towler 283a & Wilson 10 Aug 2003 (holo NSW).

Illustration: Fig. 3, e-g.

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Nannandrous, gyn- or more often idioandrosporous. Vegetative cells regular cylindrical,
L/D 4-7(-8), 11—15(—20) pm diam., cf usually finer; basal cell tall vase-like, 70-75 pm
long, 15-16 pm diam.; terminal cell with domed cap. Oogonium solitary or paired (the
upper one slightly larger than the lower one), globose, 58-81 pm long, 52-61 pm diam.,
circumcision median or submedian; oospore globose, 38-45(-51) pm diam., outer
exospore smooth, inner exospore tightly spiraled serrations, mesospore red; suffultory
cell slightly tumid, to 20 pm diam. Androsporangia intercalary, solitary or paired, if
gynandrosporous then well above oogonia, 23-25 pm long, 18 pm diam.; dwarf males
long narrow clavate, 91-104(—113) pm long, 8-11 pm diam., two spermatozoids formed
one above the other in tip, faintly septate.

Etymology: commemorates Alfred Douglas Hardy, the Victorian botanist who sent
numerous specimens to G.S. West in the 1900’s and 1910’s and continued to study
phytoplankton in Victorian reservoirs until the 1950’s (Entwisle 1990). The name
Oe. victoriense West has been used for a different taxon.

Distribution: Western Australia and Victoria, in ponds and river pools.

The description in West (1909) is extremely brief, highlighting the length of the dwarf
males, but the iconotype shows the spore wall with evenly spaced teeth. As the inner
exospore wall of Oe. monile has longitudinal serrated ribs, and that of Oe. hardyi has a
tight spiral of teeth, West’s (1909) figure 6a-b appears to fit the latter more closely.

The suffultory cell develops first, upon which dwarf males mature before the oogonia
are initiated. The form of the dwarf males with internal antheridia, the morphology of
the oospore, with a wide exospore sculptured below the outer layer, and general aspect
similar to Oe. undulatum and Oe. monile suggest a position in Group P. The presence
of a few short rhizoids on the base of the dwarf males may also indicate affinities with
Group Q.

Specimens examined: Victoria: pond opposite Phillip Island Information Centre, New I laven,
Entwisle 3133,29 Dec 2001 (NSW). Western Australia: tributary of Hammersley R., pool in river
bed, Fitzgerald River Nat. Pk, Towler283a & Wilson, 10 Aug 2003 (NSW); Brixton Street wetland,
Kcnwick, Cherry 448, 9 Sep 2002 (NSW).

Oedogonium kwangsiense Jao, Bot. Bull. Acad. Sin. 1: 85 (1947) var .playfairii Stephen
Skinner & Entwisle var. nov.

Diagnosis: Oedogonio kwangsiensi var. kwangsiensi simile sed costis undulatis
levigatisque in oospora.

Type: Western Australia: Brixton Street wetland, Kenwick (32° 01’S; 115° 58’E), Cherry
447 ,9 Sep 2002, (holo NSW).

Illustration: Fig. 3, i-1.

Nannandrous, idioandrosporous. Vegetative cells regular cylindrical, L/D 5-8, 40-74
pm long, (7-) 10-19 pm diam., cT frequently narrow, 6-8 quite large pyrenoids; basal
cell tall, narrow vase-like, 50-55 pm long, LO-12 pm diam., sometimes flattened on
one side, holdfast digitoid; terminal cell (often oogonium) with rounded or shallowly
peaked cap. Oogonium solitary to 3 in series, globose to ovoid, 32-44(-50) pm long,
32—36(—41) pm diam., (pore and) circumcision superior; oospore globose to barrel¬
shaped, top and bottom inset, oblique in the chamber, 35-39 pm long, 33-35 pm
diam., inner exospore with broad, deep, undulate costae, mesospore ruby red; suffultory

Oedogonium & Bulbochaete (Oedogoniales, Chlorophyceae) Telopea 1 1(2): 2006

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cell 48-70 pm long, 25-29 pm diam. Androspomngium intercalary, single or paired, in
groups, I 1-16 pm long, 10-13 pm diam.; dwarf males clavate, on suffultory cell, body
cell 29-35 pm long, 4.5-6 pm diam., antheridium with single spermatozoid extruded
at maturity.

Etymology: commemorates George Israel Playfair, the honorary freshwater phycologist
associated with the National Herbarium of New South Wales from 1902 to 1923. In the
draft manuscript of his Census (Playfair 1917) Playfair notes ‘Oedogonium sp. long
corrugate spore’ from Broughton Island, but did not complete the determination or
include this collection {Bottle 29) in the Census or any other publication.

Distribution: Western Australia and New South Wales, in coastal wetlands.

The Brixton Street wetland sample contains numerous filaments with terminal solitary
oogonia and the general aspect of the cosmopolitan Oe. acrosporiitn de Bary, but the
equally common occurrence of intercalary series of oogonia and the pronounced
depth of the costae favour inclusion in Oc. kwangsiense from China. The Australian
material comes close to the type form of Oe. kwangsiense but has cylindrical cells, the
costae are broad and smooth, and the basal cells are tall vase-shaped with a digitoid
holdfast. This taxon is idioandrosporous, and the antheridium, while the fragile wall
is persistent, appears to be extruded from the body cell of the dwarf male, rather than
being a separate cell of a small plant as in both Oe. acrosporiitn de Bary ex Hirn and
Oe. tentoriale Nordstedt & Hirn in Hirn.

Specimens examined: New South Wales: North Coast: Broughton Island, Dr E. Stokes (Playfair
29, 30), -1903? (NSW). Western Australia: Brixton Street wetland, Kenwick, Cherry 447 , 9 Sep
2002 (NSW).

Supersection Dispermatozoidiae

Antheridia with paired spermatozoids. In macrandrous species the spermatozoids may
be horizontal, that is lying in the antheridium one on top of the other, or vertical,
where the spermatozoids are found side by side. In nannandrous species the dwarf
male plantlet consists of a persistent body cell, sometimes a narrow vegetative cell, and
above one or a series of discrete, external antheridia, each with two spermatozoids.

Group L

Macrandrous; spermatozoids one above the other in intercalary antheridia; oogonium
opening a circumcision.

Oedogonium capricornicum Stephen Skinner & Entwisle sp. nov.

Diagnosis: Oedogonium oogonio collari cylindrico aut terminale aut basale,
circumcisione oogonii medio.

Type: Queensland: Burdekin River, downstream from Highway crossing, near Ayr
(19° 37’S; 147° 24’E), Reid270103.04, 27 Jan 2003 (holo NSW).

Illustration: Fig. 3, m-o.

Dioecious, macrandrous. Vegetative cells cylindrical, chloroplast reticulate, many small
pyrenoids, slight sexual dimorphism, L/D (5—)8—12, (20 cf'-)23-32 pm diam.; basal cell
elongate goblet-shaped, ridged disc, c. 80 pm long, c. 32 pm diam.; terminal cell domed.

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Fig. 4. Oedogonium bancroftii: a, basal cell; b, intercalary antheridium; c, empty antheridium
above series of oogonia and sufFultory cell; d, oospores in intercalary oogonia; e, terminal
oogonium at a distance from suffultory cell; f, oospore in terminal oogonium (Cribb 587.1);
Oe. starmachii var. weetalibahense: g, oogonium with dwarf male and suffultory cell, and
androsporangia above, in same filament; h, oogonium with mature oospore, and multicellular
dwarf male (Dittgley 050603.4a/b). (Scales = 20 pm)

Oogonium globose to slightly depressed globose, (63-)68-82 pm long, (68-)77-84
pm diam., with collar top and/or bottom, stained with tannin, median circumcision,
narrowly open, internal pore; oospore globose to depressed globose, 59-61 pm long,
59-65 pm diam., smooth exospore, red mesospore; suffultory cell as vegetative cells.
Antheridia solitary to groups of 4-8, 8-14 pm long, 16-20 pm diam., with a short
supporting cell at least in initial stages, spermatozoids paired, horizontally separated.

Etymology; from the Tropic of Capricorn which covers the northern third of Australia,
where this taxon is encountered.

Distribution: Tropical Australia.

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A distinctive taxon and difficult to place. It has a lantern-shaped oogonium with a
well defined median circumcision and thickened wall, which would place it close to
Oe. pmtense Transeau (1914) in subgenus Prooedogoniunt, or the nannandrous Oe.
confcrtum Hirn and Oe. perspicum Him in Supersection Monospermatozoidiae. Paired
spermatozoids in antheridia in small belts, a vase-shaped basal cell and dimensions and
general vegetative form that agree with larger members of the subgenus Oedogonium,
suggest its placement there, in Group L.

Specimens examined: Queensland: Type and Burdekin R., downstream from highway crossing,
near Ayr, Reid 270103.05, 21 (an 2003 (NSW). Western Australia: Camp Ck, Mitchell Plateau,
Kemieally 11765a, 18 Jun 2002 (NSW);

Oedogonium bnneroftii Stephen Skinner & Entwisle sp. nov.

Diagnosis: Oedogonio areolato-costato simile sed oosporibus globosis vcl subglobosis et
proportionibus minoribus.

Type: Queensland: Lake Weyba, near Noosa (26° 27’S; 153° 05’E), Cribb 587.1, 26 Jul
1964 (holo BRI 704657).

Illustration: Fig. 4, a-f.

Monoecious/dioecious macrandrous. Vegetative cells narrow cylindrical, at first
slightly capitellate near basal cell, L/D (5—)6—9(—10), (8—)9—11 pm diam.; basal cell
elongate vase-like, basal pad deeply costate; terminal cell usually with blunt peaked cap.
Oogonium spherical to ovoid, (27—)34—45 pm long, (29—)34—36 pm diam., circumcision
superior, solitary, paired, triples and fours common, five in a row occasional; oospore
spherical to subglobose, top and bottom inset, 31-34 pm diam., smooth on very
outside, longitudinally denticulate-costate from basal and apical rings, some forking
and anastomosis, inside, mesospore red?; suffultory cell terminally widely tumid, 27-32
pm upper diam. Antheridium solitary, either subepigynous or intercalary, 12-14 pm
long, c. 9 pm diam.; spermatozoids 2, horizontally displaced.

Etymology: commemorated Dr Thomas Lane Bancroft, prominent epidemiologist
and assiduous collector of natural history specimens throughout Queensland from the
late 19 th Century to WWI.

Distribution: known only from Queensland, associated with Oe. varians among
stoneworts in coastal tidal lake (Cribb’s notes state ‘highly saline’).

As the oogonia appear to develop sequentially, one or two short cylindrical cells may
be found between the tumid suffultory cell and the developed oogonia above. It is close
to the description of North African specimens attributed to Oe. palocnse Britton in
Gauthier-Lievre (1964), but Britton’s (1949) protologuc based on Philippine material
has solitary ovoid to ellipsoid oospores in similar shaped oogonia, a suffultory cell
similar to vegetative cells, and antheridia in long series. Oe. bancroftii also resembles
Oe. areolato-costatum Jao in Mrozinska (1985), but the dimensions of the Australian
specimen are all half the size of the Chinese taxon, and the oospores are spherical rather
than ovoid.

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Group T

Nannandrous; dwarf males multicellular, antheridia external; oogonia opening through
a pore.

Oedogoniumstarmachii Mrozinska, Frag. Flor. Geobot. 4:252 (1958) var. weetalibaliense
Stephen Skinner & Entwisle var. nov.

Diagnosis: Oedogonio starmarchio var. starmarchio simile sed cellulo suffultorio
inflate.

Type: New South Wales: North Western Plains: Weetalibah Waterholes (29°19’S,
147°50’E) Dingley 050603.4, 5 Jun 2003 (holo NSW).

Illustration: Fig. 4, g, h.

Nannandrous, gynandrosporous. Vegetative cells regular cylindrical, 40-120 pm
long, 7-9 pm diam.; basal cell short vase, 20-23 pm long, c. 8 pm diam.; terminal cell
truncated. Oogonium intercalary, solitary, obpyriform, 32-34 pm long, 24-30 pm diam.,
pore inframedian to inferior; oospore globose, filling chamber, covered in fine spines,
mesospore red, 23-25 pm without spines (spines add c. 1 pm to radius); suffultory
cell usually inflated, 39-50 pm long, 9-12 pm diam. Androsporangia intercalary,
in upper filament, 23-27 pm long, 8 pm diam.; dwarf males on suffultory cell, with
slightly inflated body cell (18-20 pm long, 8-9 pm diam.), one or two vegetative cells
(18—20 pm long, 4.5 pm diam.) and one or more antheridial cells (7 pm long, 4.5-5 pm
diam.), solitary? spermatozoid.

Etymology: for Weetalibah Waterholes in north-western New South Wales, from where
the taxon was first recognized.

Distribution: known from South Australia, in a transient roadside swamp in limestone
country, and New South Wales, on Najas tenuifolia in caustic (pH 9.5) artesian spring,
and on Chara sp. in coastal creek.

There are a number of closely similar taxa with spiniferous spores and inferior pores
to the oogonium. The Weetalibah specimen shows some characters in common with
both Oe. inframediale Jao and Oe. starmachii Mroz., both of which occur in China (Jao
1979). While Oe. inframediale has an inflated suffultory cell, it has regular dwarf males,
and the cells are 13-21 pm in diameter. Oe. starmachii as described by Mrozinska
(1958) and Jao (1979) lacks the inflated suffultory cell but docs have the multicellular
dwarf males, and vegetative cells 9-14 pm in diameter. Oe. excentriporum (Jao) Tiffany
is similar in dimensions, and has an inflated suffultory cell, but has a distinctly superior
pore. Australian collections are the same as Oe. starmachii as described by Mrozinska
(1985) in all aspects except the suffultory cell. We therefore recognize this taxon as a
new variety of Oe. starmachii.

Specimens examined: New South Wales: North Western Plains: Weetalibah Waterholes,
between Lightning Ridge and New Angledool, Dinghy 050603.4 a & b, 5 Jun 2003
(NSW). South Coast: Stony Ck, 3 km S of Bodalla, Skinner 0497, 27 Dec 2001 (NSW).
South Australia: Beachport-Robe road, Skinner 556, 14 Nov 1978 (NSW).

Oedogonium & Bulbochaete (Oedogoniales, Chlorophyceae) Telopea 11(2): 2006

187

Fig. 5. Bulbochaete rodwayi : a, androsporangia and zooidangium; b, immature oogonium
and dwarf male; c, mature oospore in oogonium; d, basal cell ( Entwisle 2497)-, B. wtihattertse
var. spechtii: e, oogonium with androsporangium, dwarf males and oospore, with sketch of
sculpturing ( Specht A30); B. calospora var. murrayana : f, basal cell; g, branch with androsporangia
and mature oospore, with sketch of sculpturing; h, immature oogonium with two dwarf males
( Skinner 0401, Arnold & Towler). (Scales = 20 pm)

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Bulbochaete C. Agardh

Bulbochciete rodwayi Stephen Skinner & Entwisle sp. nov.

Diagnosis: Bulbochaete nannandriata oosporis quadriglobosis et glabris, nannandriis
camelliformibus antheridiisque interne.

Type: Tasmania: pool in Melaleuca Ck, below Alexandra Ck junction, Port Davey
Track, South West National Park (43° 25’S; 146° 08’E), Entwisle2497, 3 May 1996 (holo
MEL).

Illustration: Fig. 5, a-d.

Nannandrous, idioandrosporous? Thallus a densely intertwined cushion, visible to the
naked eye. Vegetative cells subcylindrical, 48-56 pm long, 16—18 pm diant.: basal cell
vase shaped, 18-20 pm diam. Oogonium squared globose, c.52 pm long and diam.,
median circumcision; oospore filling chamber, smooth, laminated thick exospore,
mesospore red; suture in suffultory cell median, faint. Androsporangia in short lateral
or terminal series, subtending a seta, c.16 pm long, 10-12 pm diam.; dwarf male goblet
shaped, 23-25 pm long, c.10 pm diam., upper half antheridium, lower half rhizoidal,
on suffultory cell.

Etymology: commemorates Leonard Rodway, botanist and natural history collector in
Tasmania in the first half of the 20 ,h Century.

Distribution: Tasmania, known only from the type locality, as free living cushions
among mosses and leafy liverworts in high latitude stream.

A distinctive nannandrous Bulbochaete with squared globose smooth-walled oospores,
and goblet-shaped dwarf males with internal antheridia. Unlike any reported
Bulbochaete species both B. rodwayi and at least one population of 11. wuhanensis var.
wuhanensis, from Litchfield National Park in the Northern Territory, form dense masses
of filaments and appear to be free living rather than epiphytes.

Bulbochaete wuhanensis Jao, Monograpica Oedogoniales Sinicae : 377 (1979)
var. spechtii Stephen Skinner 8c Entwisle var. nov.

Diagnosis: Bulbochaetae wuhanense var. wuhanense similis, sed gynandrospora,
antheridiis epigynis atque lateralibus, oosporis verrucosis.

Type: Northern Territory: freshwater pool, South Bay, Bickerton Island (13° 46’S; 136°
13’E), Specht A30, 7 Jun 1948 (holo AD).

Illustration: Fig. 5, e.

Gynandrosporous. Oogonium 48-56 pm long, 59-88 pm diam.; oospore sculpturing
very fine warts. Androsporangia epigynous or lateral, 11-14 pm long, 12-14 pm diam.

Etymology: commemorates Raymond Louis Specht, Australian ecologist and member
of the American-Australian Expedition to Arnhem Land in 1948.

Distribution: Northern Territory, on aquatic vegetation in tropical seasonal
waterways.

Oedogonium & Bulbochaete (Oedogoniales, Chlorophyceae) Telopea 11(2): 2006

189

Fig. 6. Bulbochaete Gigantea var. glabra: a, androsporangia; b, oogonium with dwarf male,
sketch of sculpturing ( Divola s.n.(Skinner 0039))- B. kakaduensis: c, basal cell; d, androsporangia
and zooidandium; e, mature oogonium with dwarf males, oospore, and sketch of sculpturing
(. Dostine , Metcalfe & Padovan A104).{ Scales = 20 pm)

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Differs from B. wuhanensis var. wuhanensis from China in being gynandrosporous, and
the sculpturing of the oospore being very finely warty.

Specimens examined: Northern Territory: freshwater pool, South Bay, Bickerton
Island, Specht A30 , 7 Jun 1948 (AD); East Charlotte R„ Dingley 110903.2, 11 Sept 2003
(NSW).

Bulbochaete calospora Jao, Sinetisia 10: 147 (1939) var. murrayana Stephen Skinner
& Entwisle var. nov.

Diagnosis: Bulbochaetae calosporae var. calosporae similis sed in dimensionibus
brevior.

Type: New South Wales: South West Slopes: Corowa-Mulwala road (36° 00’S; 146°
16’E), Skinner 0401, Arnold & Towler, 27 Sep 2001 (holo NSW).

Illustration: Fig. 5, f-h.

Dioecious gynandrosporous.Thallusmoderatelylarge,open but infrequently branching.
Vegetative cells subcylindrical, 36-45(-48) pm long, 11—14(—18) pm diam., finely
spirally spinose; basal cell vase-like, 48 pm long, 20 pm diam., base ridged, spreading.
Oogonium depressed globo-pyriform, 36-39 pm long, 43—45 pm diam., cingulum,
opening supramedian (closed at maturity); oospore filling chamber, exospore with
large, sharp edged scrobiculae, mesospore yellow; suture in suffultory cell supramedian
to superior. Androsporangia epigynous or lateral, solitary or in series 2-4,9-11 pm long,
c. 9 pm diam.; dwarf males shortly clavate, 25-27 pm long, 7-9 pm diam., antheridia
internal, two chambers, single spermatozoid each.

Etymology: named for the Murray River valley, in which floodplain the collection was
made.

Distribution: New South Wales/Victoria Border: riverine system, on filamentous algae
and stoneworts.

This taxon is close to cosmopolitan B. intermedia de Bary in size of features and
has similar position of suffultory suture, but, with the finely spinose cells, and large
scrobiculae, appears to be a small variety of B. calospora from China.

Bulbochaete gigantea Pringsh. ex Hirn, Acta Soc. Sci. Fenn 27: 347 (1900) var. glabra
Stephen Skinner & Entwisle var. nov.

Diagnosis: Bulbochaetae giganteae var. giganteae similis sed parietibus glabris, oosporis
areolatis scrobiculis.

Type: New South Wales: Northern Tablelands: Barleyfields Lagoon, near Uralla (30°
37’S; 151° 30’E), Divola s.n. [Skinner 0039], 30 May 1996 (holo NSW).

Illustration: Fig. 6, a, b.

Vegetative cells, 60-75(-80) pm long, 19-25 pm diam., smooth walled; basal cell vase
like. Oogonium depressed globose, 43-45(-48) pm long, 58-68 pm diam.; oospore
sculpturing of large, areolate scrobiculae, mesospore yellow; suture in suffultory cell

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191

median to supramedian .Androsporangia in series 1-4, c. 12 pm long, c. 14 pm diam.,
subtending seta; dwarf males clavate to vermiform, 57-60 pm long, c. 12 pm diam.,
antheridia internal.

Etymology: from glaber, ra, rum (Latin) smooth.

Distribution: New South Wales: transient swamp and creek pondages.

Difters from the widespread type variety in having cell walls smooth rather than finely
helically spinose, and large open areolate scrobiculations on oospores. The report of
B. gigantea in Skinner (1980) may refer to this taxon, as spinose cell walls were not
reported. The specimen is missing, although other slides from the collection (Bullock
Ck, Skinner NED002 , Dec 1974(NE)) contain sterile fragments of a large Bulbochaete
with smooth cell walls.

Specimen examined: New South Wales: Northern Tablelands: on stoneworts, Barleyfields
Lagoon, near Uralla, Skinner 0036, Cooper & Divola, 29 Feb 1996, and Divola (Skinner 0039),
30 May 1996 (NSW); Central Coast: on rushes in pond of Porters Ck, Wyong, Gartenstein s.n.,
7 Dec 2001 (NSW).

Bulbochaete kakaduensis Stephen Skinner & Entwisle sp. nov.

Bulbochaete clatior sensu Scott & Prescott, Record of the Arnerican-Australian expedition
to Arnhem Land, 3: 19 (1958).

Diagnosis: Bulbochaetae scrobiculatae similis sed idioandrospora nannandriisque
clavatis, stipa 22-23 pm, antheridio 9-16 pm longo.

Type: Northern Territory: Howard R., upstream of Pioneer Dr. (12° 33’S; 131° 08’E),
Dostine, Metcalfe & Radovan A104, 30 May 2003 (holo NSW).

Illustration: Fig. 6, c-e.

Dioecious, idioandrosporous. Thallus moderate, open branching. Vegetative cells
subcylindrical, L/D 3.5-6, 12.5-14 pm diam.; basal cell short vase-like, 16-18 pm
diam., rhizoidal base. Oogonium depressed globose, 37-42 pm long, 42-50 pm diam.,
cingulum, pore superior; oospore filling chamber, exospore very finely scrobiculate,
mesospore yellow; suture in suffultory cell basal. Androsporangia lateral in series
1-4, 9-14 pm long, 11 pm diam.; dwarf male body cell stalked goblet, 22-23 pm long,
10 pm diam., antheridium single, 9-16 pm long, 7-8 pm diam.

Etymology: named for the Kakadu region of the Northern Territory over which the
Arnerican-Australian expedition to Arnhem Land travelled and collected.

Distribution: Northern Territory, in still and flowing water.

Scott and Prescott (1958) described and figured the Bickerton Island specimen
as Bulbochaete elatior Brings., illustrating clearly the fine scrobiculae. B. elatior, a
cosmopolitan species, has smooth exospores. Scott and Prescott failed to mention
if androsporangia were present. The Arnhem Land collections, and the Howard R.
collection are idioandrosporous, and have dwarf males with almost clavate rather than
goblet shaped body cells (as in B. elatior), and along with the fine scrobiculae on the
oospore, make this a distinctive species.

192

Telopea 11(2): 2006

Skinner and Entwisle

Specimens examined: Northern Territory: freshwater pool, South Bay, Bickerton Is., SpechtA30,
7 iun 1948 (AD); freshwater marsh, behind dunes, Yirrkalla, Specht A80, 27 Aug 1948 (AD);
Howard R., upstream of Pioneer Dr., Dostine, Metcalfe & Padovan A104, 30 May 2003 (NSW).

Acknowledgments

Thanks are due to the many co-workers and interested citizens who sent samples to the
National Herbarium of New South Wales in the Botanic Gardens Trust, Sydney, Western
Australia: Dorothy Edinger, Alan Lowrie, and Kevin Kenneally (CALM);Northern
Territory: Mike Dostine, Peter Metcalfe, Arrnand Padovan and Simon Townsend (Water
Labs, NP&WS); Queensland: David Reid, Townsville; New South Wales: Margaret Brock,
Chris Cooper and Jill Divola in Armidale, and Simon Gartenstein at Sydney University,
N.S.W. and our many colleagues here who included ‘green slime’ in their collections,
and especially to the late Marion Wi§cek for translations from Polish, and Ms Alice
Cheung for translations from Mandarin. All major herbaria in Australia checked their
collections and loaned specimens, as did the British Museum and LD. ABRS provided
the funding. Dr Teresa Mroziriska, Wl. Szafer Institute of Botany, Polish Academy of
Sciences, Krakow, provided encouragement in the early stages of this project.

References

Bailey FM (1893) Contributions to the Queensland Flora. Queensland Freshwater Algae. Botany
Bulletin of the Department of Agriculture, Queensland No. 6.

Bailey FM (1895) Contributions to the Queensland Flora. Queensland Freshwater Algae. Botany
Bulletin of the Department of Agriculture, Queensland No. 11.

Bailey FM (1898) Contributions to the Queensland Flora. Queensland Freshwater Algae. Botany
Bulletin of the Department of Agriculture, Queensland No. 15.

Britton ME (1949) New species of Oedogonium from Leyte, the Philippine Islands. American
Journal of Botany 35: 715-719.

Cribb AB (1956) A new terrestrial alga from Australia. Proceedings of the Royal Society of Queens¬
land 67: 25-26.

Cribb AB (1965) An ecological and taxonomic account of the algae of a semi-marine cavern,
Paradise Cave, Queensland. Papers of the Department of Botany University of Queensland
4(16): 259-282.

Cribb AB (1988) Filamentous Algae. Pp. 39-43 in Scott G (ed.) Lake Broadwater. The Natural
History of an Inland Lake and its Environs (Darling Downs Institute Press: Toowoomba)

Day SA, Wickham RP, Entwisle TJ & Tyler PA (1995) Bibliographic Checklist of Non-Marine Algae
in Australia. (Australian Biological Resources Study: Canberra)

Entwisle TJ (1990) The lean legacy of freshwater phycology in Victoria. Pp. 239-246 in Short
PS (ed.) History of Systematic Botany in Australia. (Australian Systematic Botany Society:
Melbourne)

Gauthier-Lievre L (1964) Ocdogoniacees Africaines. Nova Hedwigia 7: 153-272, 273-481,545-
558.

Hansgirg A (1886) Prodromus der Algenflora von Bohmen. Erster Theil. Arcltiv fiir die
naturwissenschaftliche Landesdurchforshung von Bohmen 5(6): 1-96.

Herring W (1914) Chlorophyceae III Ulothricales, Microsporales, Oedogoniales. DieSiisswasser-
Flora Deutschlands, Osterreichs und der Schweiz 6 (Gustav Fischer Verlag: Jena)

Berkeley MJ & Harvey W1I (1860) Fungi and algae. Pp. 241-343, plate CXCVI, B in Hooker JD
The botany of the Antarctic voyage of H.M. discovery ships Erebus and Terror, 1839-43. III.
Flora Tasmaniae. Part 2. Monocotyledones and Acotyledones.

Oedogonium & Bulbochaete (Oedogoniales, Chlorophyceae) Telopea 1 1(2): 2006

193

Hirn KE (1900) Monographic und Iconographie der Oedogoniaceen. Ada Societatis Scientiarum
Fennicae 29:395 pp, 64 plates.

Jao Chin—Chih (1979) Motiographia Oedogoniales Sinicae (Science Press: Beijing)

Leonardi PI, Caceres EJ & Velez, CG (1998) Fine structure of dwarf males in Oedogonium plu-
viale (Chlorophyceae). Journal of Phycology 34: 250-256.

Mroziriska T (1985) Chlorophyta VI Oedogoniophyceae: Oedogoniales. Siisswasserflora von
Mitteleuropa 14 (Gustav Fischer Verlag: Jena)

Mrozinska T (1991) A preliminary investigation of the taxonomical classification of the genus
Oedogonium Link (Oedogoniales) based on the phylogenetic relationship. Archiv fiir
Protistenkunde 139: 85-101.

Mrozinska T (1999) New proposals for classification of Oedogonium genus. Pp. 31-33 in XVIII
Symposium Sekcji Fykologicznej PTB, Kielce-Wdlka Milanowska 6-9 maja 1999.

Mrozinska T (2000) Oedogonium szymanskae sp. nov. (Oedogoniales, Chlorophyceae, Chloro¬
phyta). Archiv fiir Hydrobiolica, Supplementband 113 (Algological Studies 98): 1-5.

Nordstedt CFO (1887) Utdrag ur ett arbete ofver de af Dr S. Berggren pa Nya Seland och I
Australien samlade sotvattensalgerna. Botaniska Notiser 1887: 153-164.

Nordstedt CFO (1888) Freshwater algae collected by Dr S Berggren in New Zealand and Aus¬
tralia. Bihang til Kongliga Svenska Vetenskaps-Akademiens Handlingar 22: 1-98.

Novis PM (2003) A taxonomic survey of Oedogonium (Oedogoniales, Chlorophyta) in the South
Island and Chatham Islands, New Zealand. New Zealand Journal of Botany 4 1: 335-358.

Playfair GI (1917) Supplement 1 Fresh-water Algae. Pp. 219-263 in Maiden JH & Betche E A
Census of New South Wales Plants. (Government Printer: Sydney)

Scott AM 8c Prescott GW (1958) Some Freshwater Algae from Arnhem Land in the Northern
Territory. Pp. 9-136 in Specht RL & Mountford CP (eds) Records of the American-Austral-
ian Scientific Expedition to Arnhem Land 3 Botany and Plant Ecology. (Melbourne University
Press: Melbourne)

Skinner S (1980) New Records of Zygnemaphyceae and Oedogoniophyceae (Chlorophyta) from
northern New South Wales. Proceedings of the Linnaean Society of New South Wales 104:
245-263.

Sonder O (1852) Plantae Muellerianae, Algae. Linnaea 25: 657-709.

Transeau EN (1914) New species of green algae. American Journal of Botany 1: 289-301.

Velez CG (1995) Oedogoniales (Chlorophyta) de la Republica Argentina. IV. Bolotin de la Socie-
dad Argentina de Botanica. 31:113-123.

West GS (1909) The Algae of Yan Yean Reservoir, Victoria: a biological and oecolgical study.
Journal of the Linnaean Society, Botany 39: 1-88.

Zhu Hui-Zhong (1964) Materials for the Oedogoniaceae of China (1). Acta Phytotaxonomica
Sinica 9: 203-211, 3 plates.

Manuscript received 2 September 2005, accepted 10 February 2006

.

Telopea 11(2) 195-232

A review of the taxonomy and relationships
of the Dendrobium speciosum complex
(Orchidaceae), and recognition of two new taxa

Peter B. Adams, Jacinta M. Burke and Sheryl D. Lawson

The School of Botany, University of Melbourne, Victoria, Australia 3010

Abstract

Dendrobium speciosum is one of Australia’s most prominent orchids. Two hundred years after
the type description, there are several contrasting taxonomic treatments of the D. speciosum
complex. We review the taxonomy from first descriptions and incorporate recent numerical
approaches. Our explorations extend the plant range to an almost continuous distribution along
the east coast of Australia, with disjunct populations along the Tropic of Capricorn. Data from
field observations, and studies of 453 plants over the entire distribution range were analysed
using morphological and multivariate techniques. Results indicate a species complex forming
a continuum of variation. Within the continuum from north to south the varieties boreale,
pedunculatum, curvicatdc, capricornicum, grandiflorum, hillii and speciosum occupy geographical
areas which are defined approximately, and they exhibit significant infra-varietal variation. At
interfaces of described varieties, populations show intermediate characteristics. There are no
distinct qualitative differences in characters between any of the varieties or groups. We describe
two new varieties, var. blackdownense and var. carnarvonense from central Queensland and
propose that var. capricornicum Clemesha be retained for plants of east Capricorn.

Introduction

Dendrobium speciosum Sm. is one of Australia’s most outstanding, widespread, and
variable orchids. The complexity of variation has been highlighted by Burke (1998)
and Burke and Adams (2002) using numerical analytical methods. They interpreted
the complex as consisting of intergrading varieties, with some forms that could not
be ascribed to any distinct groups. Nonetheless, taxa can be identified and treated as
varieties. In very confusing populations along the Tropic of Capricorn, a detailed study
of north Queensland plants resulted in the description of a new variety, var. boreale ,
and establishment of a neotype (and narrowed re-circumscription) for plants of var.
curvicaule , which occurs between St Lawrence and Mt Dryander (Adams et al. 2006a).

Detailed field work from eastern Victoria to far north Queensland and west to Carnarvon
Gorge has led to a reappraisal of the species and its taxonomy, defining the limits of
taxa in relation to habitat and latitude. In this study we analyse a data set from all areas,

© 2006 Royal Botanic Gardens and Domain Trust

ISSN0312-9764

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Adams, Burke and Lawson

including plants from recently explored remote areas not previously known to contain
D. speciosum. We review taxonomic approaches and the currently defined varieties and
geographically defined groups, which are compared in relation to the complex.

Historical review

Detidrobium speciosum was described by Smith (1804), and the type specimen is held
in the Linnaean herbarium at the Linnaean Society in London (LINN). 'Port Jackson
New South Wales’ is written on the type sheet and the following is a transcription of
the original description.

“DENDROBIUM speciosum Great Dendrobium

GYNANDRIA Diandria

“Nectary a lip without a spur. Petals 5; the 2 lower external ones forming a pouch with
their base. Lid vertical.

“Flowers often reversed. Cluster solitary, many-flowered. Lip three-lobed. Leaves radical,
coriaceous.

Dendrobium speciosum var. hillii was described by Masters in 1877 (having been
described previously at species rank as D. hillii by Sir William Hooker in 1861), followed
by var. curvicaule and var. grandiflorum (Bailey 1896). The original descriptions are not
sufficiently diagnostic to distinguish the varieties in question from other members of
this complex. The varieties established in the nineteenth century were based on a few
specimens, and as new areas of distribution were examined the range of morphological
variation was found to be far greater than initially appreciated, blurring distinctions
between some varieties. This has led to questions about the taxonomic rank of these
variants (Clemesha 1982). Since DockrilPs first book in 1969 there have been reviews
by several authors (Clemesha 1981a, 1981b, 1986, Clements 1989, Banks & Clemesha
1990, Adams 1991), with limited new evidence offered.

Dockrill (1969) initially recognised five varieties: var. speciosum, var. hillii Mast.,
var. grandiflorum F.M.Bailey, var. curvicaule F.M.Bailey and var. nitidum F.M.Bailey.
Clemesha (1981a, 1986) reviewed the described varieties, reproduced original
descriptions, photographs of type specimens, photocopies of plants typical of varieties,
and commented on atypical forms and variation in central Queensland, without
making categorical statements about taxonomic rank. He also described a new variety,
var. pedunculatum, from the Atherton area of north Queensland (Clemesha 1981a). He
regarded the central Queensland plants, and plants at Byfield and west at Carnarvon
Gorge, as dwarf races of var .grandiflorum. In 1982, Clemesha described a new variety,
var. capricornicum, from the volcanic plug of Mt Jim Crow, near Rockhampton in Central
Queensland, and considered that plants at Blackdown Tableland and Carnarvon Gorge
represented different colour forms of this variety. Plants from different habitats close to
the plugs were noted to differ from typical var. capricornicum in some characters.

The six varieties of D. speciosum recognised by Dockrill (1992) were D. speciosum
var. speciosum (Smith 1804), D. speciosum var. hillii (Masters 1877), D. speciosum
var. grandiflorum (Bailey 1896), D. speciosum var. curvicaule (Bailey 1896), D. speciosum
var. pedunculatum (Clemesha 1981), and D. speciosum var. capricornicum (Clemesha
1982). Dendrobium speciosum var. nitidum F.M.Bailey, D. x nitidum (F.M.Bailey)

Dendrobium speciosum

Telopea 11(2): 2006

197

M.A.Clem. & D.L.Jones, D. speciosum var. bancroftianum Rchb.f., and D.jonesii Rendle
subsp. bancroftianum (Rchb.f.) M.A.Clem. & D.L.Jones are now considered to represent
D. xgracillimum, a natural hybrid between D. speciosum and D. gracilicaule (Clemesha
1981a, Banks & Clemesha 1990).

Jones (1988) treated D. speciosum as having six varieties. Upton (1989) recognised ‘an
extremely variable species’ of six varieties: var. speciosum, var. hillii, var. grandiflorum,
var. capricornicum, var. curvicaule and var. pedunculatum. Clements (1989) raised these
varieties to species rank, with the exception of var. capricornicum, which was subsumed
under D. curvicaule. Clements defined his species concept as one or more significant
and consistent differences in characters, and described differences for each proposed
species that have not been supported by any published studies. The changes have not
been accepted by subsequent taxonomic reviewers (Banks & Clemesha 1990; Grundon
et al. 1990; Adams 1991; Lavarack 1991; Dockrill 1992, 1995).

Banks and Clemesha (1990), in rejecting Clements’ taxonomic treatment, regarded
var. speciosum, var. hillii, var. grandiflorum, var. curvicaule, var. pedunculatum and var.
capricornicum as subspecies, noting that some shared the same habitat, and intermediate
forms were present. Reasons for proposing the rank of subspecies were not offered. In
his revised Australian Indigenous Orchids in 1992, Dockrill recognised six varieties,
and a degree of overlap, stating that his attempt at developing a key could not separate
all plants into distinct varieties. Lavarack and Gray (1985) also accorded the taxa
varietal rank. Adams (1991) provided a review of variation, pollination and breeding
systems in D. speciosum, and recommended the retention of six varieties that were not
completely distinct, with overlapping habitat, and variable, inconsistent features.

All reviews suggested separate distributions for the varieties, with large gaps, especially
in dry areas between Rockhampton and Mackay, and around Townsville. Since then
the range of known forms, localities and distribution has progressively extended. The
following deficiencies are evident in past treatments: insufficient sampling of regions
using living material, errors in distributions, assumptions of gaps in distribution in dry
and inaccessible habitat, lack of published data to support conclusions and taxonomic
proposals, and a strong tendency to place all plants into distinct taxonomic groups,
whether at species, subspecies or varietal rank, even when their characteristics do
not correspond with the circumscription of taxa. Clements (1989) used character
differences including aerial roots, pseudobulb shape, floral segments, labellar notches,
and breeding biology to propose separate species. The use of these characters for
diagnosis has been refuted (Banks & Clemesha 1990, Adams 1991, Dockrill 1992, Burke
& Adams 2002). In view of the large distribution range and variability, these problems
are understandable, but have led to considerable confusion.

Subsequent numerical approaches (Burke 1998, Burke & Adams 2002) have supported
the concept of a species complex, which required further survey and analysis. We
have published new distributional data, undertaken numerical analyses in all regions,
and described a new variety, var. boreale which applies to rainforest plants north of
Townsville that had previously been included in var. curvicaule. We have also respecified
the range of occurrence of neotypified var. curvicaule (Adams et al. 2006a) (Table 1).

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Adams, Burke and Lawson

Table 1. Recent classifications of Dendrobium speciosum.

Clements 1989

D. speciosum
D. tarberi
D. rex

D. curvicaule
D. curvicaule

D. pedunculatum

Banks and
Clemesha 1990

subsp. speciosum

subsp. hillii

subsp. grandiflorum

subsp. capricornicum

subsp. curvicaule

subsp. pedunculatum

Adams 1991

var. speciosum
var. hillii

var. grandiflorum
var. capricornicum
var. curvicaule

var. pedunculatum

Dockrill 1992

var. speciosum
var. hillii

var. grandiflorum
var. capricornicum
var. curvicaule

var. pedunculatum

Adams et al.
2006a

var. speciosum
var. hillii

var. grandiflorum

var. capricornicum

var. curvicaule,
var. boreale

var. pedunculatum

We have now analysed our entire data set for D. speciosum to extend our understanding
of the species and to test previous taxonomic recommendations (Burke & Adams
2002 ).

Materials and methods
Sampling

Specimens were collected during field trips conducted in all regions during the flowering
season (May to September) between 1981 and 2005. Populations studied represented
the variation throughout each geographic area. In each study population, 6-12 plants
were sampled to encompass the major variation in plant and floral characteristics. The
following numbers of individuals were used in the analyses for the regions depicted
in Fig. 1: Region la-39 plants, Region lb-37 plants, Region 2-35 plants, Region 3-13
plants, Region 4—25 plants, Region 5-43 plants, Region 6-46 plants, Regions 7 and
8-52 plants. Wherever possible measurements were made in the field, otherwise samples
were taken by removing a small number of pseudobulbs, together with roots, from the
side of the plant to minimise damage and prevent local depletion. All plants were then
grown in aged pine park in raised beds or pots under greenhouse conditions.

A reference collection of the sampled plants, photographic library and pressed
floral parts have been maintained, as fully representative herbarium specimens
of such large plants are not practical. Four hundred and fifty three plants were
measured. Voucher details (including locations) of these plants can be obtained
by contacting the Queensland Herbarium and the National Herbarium of Victoria.
Dendrobium speciosum is under heavy pressure from illegal collecting, and thus no
specific locality information is provided here.

Seventy-two characters were used to assess variation in the D. speciosum complex.
Thirty-three characters were subsequently considered to be of taxonomic value (Table
2) and used in univariate and multivariate analyses.

Dendrobium speciosum

Telopea 11(2): 2006

199

Table 2. Morphological characters used in analyses for variation within and between
populations of Dendrobium speciosum.

Vegetative characters

1. Pseudobulb length (cm)

2. Pseudobulb width at base (cm)

3. Pseudobulb width at midpoint (cm)

4. Pseudobulb width at top (cm)

5. Number of internodes in pseudobulb

6. Coilum (narrowing of pseudobulb above swollen base) 0 = absent,

1 = 0.5- < 1.0 cm long, 2 = 1.0 -2.0 cm long, 3 = > 2.0 cm long

7. Number of leaves

8. Leaf length (cm)

9. Leaf width (cm)

10. Leaf thickness (mm)

11. Aerial roots 0 = nil, 1 = 1-3 per 10 pseudobulbs, 2 = 4-25 per 10 pseudobulbs
(roots usually > 10 cm in height), 3 = > 25 per 10 pseudobulbs

(roots usually erect and > 30 cm in height)

Floral characters

12. Number of flowers

13. Rachis length (cm)

14. Peduncle length (cm)

15. Peduncle width (mm)

16. Flower length (dorsal sepal tip-lateral sepal tips) (cm)

17. Flower length (dorsal sepal tip-mentum) (cm)

18. Flower width across petals (flattened) (cm)

19. Flower width at mentum (flattened) (cm)

20. Dorsal sepal length (cm)

21. Dorsal sepal width at base (cm)

22. Petal length (cm)

23. Petal width at base (mm)

24. Individual lateral sepal width at base (cm)

25. Width of lateral sepals at widest point (flattened) (cm)

26. Distance between lateral sepal tips (flattened) (cm)

27. Inner width of lateral sepals at widest point (flattened) (cm)

28. Distance from base of lateral sepals to their tips (cm)

29. Pedicel length (cm)

30. Midlobe length (cm)

31. Midlobe width (cm)

32. Forelobe A length (cm)

33. Forelobe width (cm)

A Fore!obe = lateral lobes plus lamina

Morphological data were collected from the 453 specimens. Specimens were coded
according to their region (Fig. 1) to facilitate discussion and the formation of hypotheses.
Individuals were allocated into regions on the basis of latitude, distribution ranges of
taxa previously circumscribed, and additional distribution from recent exploration.
A total of 281 individuals were used for the multivariate analyses in this study using
random sampling from the larger data set of 453 plants, so that all regions were
adequately represented in relation to their geography and extent of variation.

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Telopea 11(2): 2006

Adams, Burke and Lawson

An attempt was made to use pickled and prepared specimens already available or
prepared from our initial surveys. This method was abandoned as the material did not
preserve well and many of the characters of Table 2 could not be accurately assessed.

Therefore fresh living material was used for all measurements. For each specimen,
measurements were made on three representative organs where possible, and the
mean value used for analyses. The first leaf from the base of the pseudobulb was used
for all leaf measurements. In the few cases where it was not possible to obtain three
measurements, due to damaged or missing parts, mean values were calculated from the
available material.

Phenetic analysis

Morphological data were range standardised, and a Manhattan Metric (MM) distance
matrix (Sneath and Sokal 1973, Milligan and Cooper 1988) was calculated. The data
set was subjected to numerical pattern analysis using the PATN package (Belbin 1988).
For cluster analysis, the unweighted pair-group arithmetic averaging (UPGMA) was
used. A cluster intensity co-efficient ((3=-0.25) was chosen because this setting is space-
dilating, and affords maximum opportunity to separate subgroups. It sharpens any
discontinuities in the system so that clustering is intensified (Williams 1975a). It is
particularly useful when it is suspected that no sharp discontinuities exist in the data,
and it may reveal unsuspected groupings. Cramer values were calculated as previously
(Burke & Adams 2002) to determine which characters best discriminated the final
groups identified.

Fig. 1. Distribution range of D. speciosum species complex showing regions defined for this
study.

Dendrobium speciosum

Teiopea 11(2): 2006

201

The distance matrix was used for ordination by principal coordinates analysis (PCoA)
(Gower 1966). Ordination techniques arc considered more appropriate for analysis at
the population level because, unlike cluster analysis, they do not assume a hierarchical
pattern of relationships, which may not be present at the infra-specific level (Gould
& Johnston 1972, Crisp & Weston 1993). Character vectors and their correlation
coefficients were determined to allow examination of trends of characters across the
ordination space (Williams 1975b, Belbin 1988).

Variation of characters within and between taxa

Maximum, minimum and mean values, and first to third interquartile ranges were
calculated for pseudobulb, leaf and flower length for plants in Regions 1-8 of Fig. 1.

Results

Cluster analysis combining vegetative and floral characters

Nine groups were identified in the UPGMA classification, seven containing a mix of
plants from different regions or representing different varieties (Fig. 2, Table 3).

>s

'£Z

_CU

I

‘t/)
c n

b

Group

Regional Composition

A

la

B

la (Paluma, Tinaroo, Tully), 1b, 3, 4

C

1b, la (Tinaroo), 4

D

IP, 2, 3, 4, 5, 6, 7, 8

E

5

F

2, 4, 5, 7, 8

G

2, 4, 7, 8

H

2, 3, 5, 7, 8

1

6

Fig. 2. Classification (UPGMA, (3 = -0.25) ofD. speciosum individuals throughout the distribution
range using vegetative and floral morphological characters truncated at nine groups.

202

Telopea 11(2): 2006

Adams, Burke and Lawson

Table 3. Cramer values for the most important characters of each dichotomy in the classification
of D. speciosum individuals from throughout the distribution range based on vegetative and
floral morphological characters (UPGMA p=-0.25).

Dichotomy

Character Number and Character

Cramer Value

1

33, For'elobe width

0.6926

31, Midlobe width

0.6694

2

18. Flower width across petals (flattened)

0.7793

16. Flower length (dorsal sepal tip-lateral sepal tips)

0.7666

20. Dorsal sepal length

0.7658

22. Petal length

0.7600

28. Distance from base of lateral sepals to their tips

0.6961

3

6. Collum

0.8331

1. Pseudobulb length

0.6746

9. Leaf width

0.6584

4

24. Individual lateral sepal width at base

0.7591

25. Width of the lateral sepals at widest point (flattened)

0.7328

21. Dorsal sepal width at base

0.7132

25. Width of lateral sepals at widest point (flattened)

0.6702

5

28. Distance from base of lateral sepals to their tips

0.7621

14. Peduncle length

0.7393

5. Number internodes in pseudobulb

0.6841

15. Peduncle width

0.6713

8. Leaf length

0.6501

6

1. Pseudobulb length

0.7801

12, Number of flowers

0.7514

7. Number of leaves

0.6601

7

22. Petal length

0.8327

24. Individual lateral sepal width at base

0.8134

28. Distance from base of lateral sepals to their tips

0.7935

21. Dorsal sepal width at base

0.7628

23. Petal width at base

0.7429

8

18. Flower width across petals (flattened)

0.8744

16. Flower length (dorsal sepal tip-lateral sepal tips)

0.8720

22. Petal length

0.8612

20, Dorsal sepal length

0.8665

The first dichotomy separated Group I, representing most, but not all, plants of var.
hillii (or plants from Region 6). The second dichotomy separated the mixed Groups E,
F, G, and H from mixed Groups B, C, D, and Group A, which represents rainforest plants
north of Townsville. Some north Queensland plants of shorter stature were clustered in
Groups B and C. While Cramer values for most floral and the collum characters were
moderately high (>0.8), their values indicate that there is still considerable overlap
between the groups formed at the eighth dichotomy. The remaining characters had
low to moderate Cramer values, indicating considerable overlap in character values
between the groups formed at this level.

Dendrobium speciosum

Telopea 11(2): 2006

203

Axis 1

var. grandiflorum
: Central Qld west
V Central Qld east

Fig. 3. Ordination from PCoA in three dimensions of D. speciosum individuals throughout the
distribution range using vegetative and floral morphological characters, (a) Axis 1 v. 2. (b) Axis
2 v. 3. Character vectors (correlations > 0.7) showing angles in ordination space are shown. See
Table 2 for character codes.

204

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Adams, Burke and Lawson

Cluster analysis using vegetative or floral characters alone

These analyses produced seven sub groups that were mixtures of plants from several
different regions, similar to that produced by using all morphological characters.

Principal coordinates analysis combining vegetative
and floral morphological characters

Rainforest plants north of Townsville partially separated from the main complex
with minimal overlap. These plants form a group largely abutting the main complex
in ordination space (Fig. 3). They partially separate on the basis of the presence of a
collum, longer pedicels, and larger width at the top of the pseudobulbs. There are no
absolute differences in characters of this group when compared to others. Presence
of a collum also contributed to partial separation of var. hillii plants of rainforest
in northern New South Wales and southern Queensland. Compared with all other
D. speciosum, these plants had some of the longest pseudobulbs, which were wider at
the top, with more internodes and longer leaves.

Plants of the dry country north of Townsville, typical habitat for var. pediinculatum,
separate from var. curvicaule on the basis of longer peduncles. Longer peduncles are also
found in some plants of var. boreale and not in all war. pediinculatum so the character is
not diagnostic for var. pediinculatum. In the total analysis the other varieties all overlap
without significant separation.

Regional principal coordinates analysis

The ordination of all 281 plants (Fig. 3) indicates a species complex. An analysis of
regional subsets of plants was required to examine further for putative geographic
varieties.

Regions 1 and 2: The northern rainforest group (var. boreale) has minimal overlap in
ordination space with the southern rainforest group (var. curvicaule) ot Region 2 (Fig.
4). There is considerable overlap between plants of var. pediinculatum in open forested
dry country on and west of the divide, and var. boreale, and little overlap between var.
pediinculatum and var. curvicaule.

Regions 3 and 4: The analysis indicates three groupings along the Tropic of Capricorn.
The Carnarvon area group and Blackdown Tableland group abut one another with
minimal overlap in the ordination space, and var. capricornicum of east Capricorn
overlaps with the Blackdown Tableland group to some extent (Fig. 5). Variation in these
regions is very extensive, especially at Blackdown Tableland, on the volcanic plugs near
Rockhampton, and at Byfield and Shoalwater Bay. In the ordination space, plants from
Mt Jim Crow, the type locality for var. capricornicum, lie at a considerable distance from
those of other volcanic plugs.

Blackdown Tableland and Carnarvon Gorge plants are separated due to the higher
character values in Carnarvon Gorge plants for forelobe length, flower length, dorsal
sepal length, forelobe width, petal length and individual lateral sepal width. Plants on
the plugs are separated from Blackdown Tableland and Carnarvon Gorge plants by
lower values for pseudobulb length and leaf length.

Dendrobium speciosum

Telopea 11(2): 2006

205

Axis 1

<1 Paluma
v Tinaroo

Fig. 4. Ordination from PCoA in three dimensions of D. speciosum individuals from north
Queensland using vegetative and floral morphological characters, (a) Axis 1 v. 2. (b) Axis 2 v. 3.
Character vectors (correlations > 0.7) showing angles in ordination space are shown. See Table
2 for character codes.

206

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Adams, Burke and Lawson

Regions 5 and 6: Ordination of Region 5 plants (var. grcmdiflorum) and plants along
the Tropic of Capricorn (Fig. 6) indicate a small degree of overlap between Region 5
and plants of Blackdown Tableland and Carnarvon Gorge. Shorter plants from isolated
populations of the southern part of Region 2 (Sarina, Clairview), and northern part of
Region 5 (Mt Morgan area) (Fig. 6), overlap in ordination space with the three groups
of Regions 3 and 4. There is no overlap between plants of Region 5 and east Capricorn.
Ordination of Regions 5 and 6 indicated no significant sub groupings in each of these
areas (Figs 7, 8). Intermediates are found at the junction between Regions 5 and 6,
where the distribution of D. speciosum is continuous.

Regions 7 and 8: In the habitat of var. speciosum, there is a cohesive group with no
significant sub groupings (Fig. 9).

Two new taxa from central Queensland

On the basis of field studies, regional ordination and cluster analysis, we consider that
var. capricornicutn Clemesha does not encompass the variation seen in west Capricorn,
at Blackdown Tableland and the Carnarvon area, and recommend it be used only
for plants from the volcanic plugs and forests of east Capricorn near Rockhampton,
Yeppoon and Byfield.

Field observations and the numerical analyses are used to describe two new taxa. For all
taxa, in the descriptions that follow, the dimensions used for flower vertical height and
horizontal width refer to flowers pressed flat, as presentation dimensions have been
shown to vary widely under different environmental conditions and at different times
during the day (Burke 1998).

Dettdrobium speciosum Sm. var. blackdownense R B.Adams var. nov.

Plantae variabiles sed saepe urceolatae; axes breves vel mediocres (9-28 cm longi)
e basi saepe divergentes; folia ovata vel subelliptica; racemi mediocres vel longi ex
comparatione altitudinis plantae (23-58 cm longi) pauciflori vel multiflori; flores
subdistantes vel aggregati (14-113 per racemum), parvi vel mediocres (3.5-5.4 cm alti)
late aperientes; tepala albida vel ochroleuca.

Type: Queensland, Leichhardt District, Blackdown Tableland, P. B. Adams & S.D.
Lawson, 4 Sepl994 (holo: QRS; iso: BRI, MEL).

Plants very variable in size and shape, occasional plants showing a tendency to produce
aerial roots; axes and leaves exhibiting purple pigmentation in parts exposed to direct
light. Leaf bearing axes 9-28 cm long, 2.1-3.8 cm wide at mid point, round to oval in
cross section, variably erect or curved, 1.9-4.6 cm wide at base, 1.5-3.2 cm wide at
apex. Leaves terminal, 2-4, rigid, concave, 9-20 cm long, 4.3-7 cm wide, mainly ovate
to sub-elliptic, broadly ovate on small plants, with a sheathing base, variably sub-elliptic
or sub-oblong, concave, rigid, 1-2 mm thick. Racemes 1-3, 23-58 cm long; peduncle
8-15.5 cm long, 4-6 mm wide, with 1-4 medium sized bracts; rachis 15-42 cm long,
14-113-flowered: flower density varying from openly spaced to densely clustered,
forming a brush. Pedicel including ovary 2.3-3.7 cm long. Flowers usually opening
widely, variable in size and substance, vertical height 3.5-5.4 cm, horizontal width
3.9-5.4 cm, off white to deep gold, the labellum variously and diffusely marked with
purple spots or short bars over forelobe and midlobe. Dorsal sepal 1.8-3 cm long,

Dendrobium spedosum

Telopea 11(2): 2006

207

b

Axis 1

Locality
■ Carnarvon G.
A Blackdown T.
O Volcanic plugs

Fig. 5. Ordination from PCoA in three dimensions of D. spedosum individuals from central
Queensland with vegetative and floral morphological characters, (a) Axis 1 v. 2. (b) Axis 1 v. 3.
Character vectors (correlations > 0.8) showing angles in ordination space are shown. See Table
2 for character codes.

208

Telopea 11(2): 2006

Adams, Burke and Lawson

0.5-0.8 cm wide at the base, tapering to an obtuse-rounded apex. Lateral sepals
1.6-2.4 cm long, 0.7-1 cm wide at base, falcate-oblong, obtuse at apex. Petals 1.8-2.7
cm long, 3-4 mm wide, slightly falcate, with an acute apex. Labellum 0.8-1.3 cm long,
0.9-1.2 cm wide; forelobe 0.5-0.7 cm long, 0.9-1.2 cm wide when flattened, incurved,
sub triangular; midlobe shortly clawed, 0.3-0.6 cm long, 0.7—1.0 cm wide when flattened,
presenting as a curved channel with a short acute apex; callus slightly raised, 2-ridged,
yellow to orange. Column c. 5 mm long with a foot c. 6 mm long at right angles to it.
Mentum bifid, rounded, c. 7 mm from ovary to apex.

Variation: the most variable in vegetative and floral form of all the described varieties,
with flowers densely or openly spaced, flowers white to deep gold, and plant height and
shape varying greatly over the wide range of habitats in this region.

Flowering time: August-September.

Distribution: from the northern limit of Blackdown Tableland southwards along the
Expedition Range, southern limit not determined.

Etymology: the epithet refers to Blackdown Tableland.

Illustration: see Fig. 10.

Notes: the population is geographically separated from plants of the Great Dividing
Range on the east coast and Carnarvon Gorge to the west, by flood plains that are
unsuitable habitat for D. speciosum. Plants of D. speciostim var. blackdownense are
usually lithophytic, urn or basket shaped, occasionally epiphytic in sclerophyll forests,
dry rainforest and exposed rocky cliff faces, often forming dense populations with
Drynaria (Bory) J.Sm. fern on huge boulders out of reach of fire. Larger plants may
have a few aerial roots, which are not usually well developed. Some plants are extremely
short and compact at maturity. Although very variable, they can usually be distinguished
from var. carnarvonense, which tend to be more robust, with large flowers of very heavy
substance. There is some overlap in floral and vegetative characters with plants of var.
capricornicum , which can usually be distinguished by vegetative features of shorter axes
and leaves, more upright plant shape, and an earlier flowering time (May-June).

Dendrobium speciosum Sm. var. carnarvonense P. B.Adams var. nov.

Planta robusta, plerumque urceolata vel crateriforma; axes breves vel mediocres
(15-33 cm longi) e basi divergentes, basi lati, apicem versus contracti; racemi breves
vel mediocres (21-47 cm longi); flores 25-87 per racemum, subdistantes, grandes
(5.1-6.6 cm alti), cupulati; segmenta lata crassiuscula; lobus frontalis 1-1.4 cm latus et
c. 0.7 cm longus; tepala crernea vel aurea.

Type: Queensland Leichhardt District. Carnarvon Gorge, P.B. Adams & S.D. Lawson,
11 Sep 1994 (holo: QRS; iso: BRI, MEL).

Plants robust, urn or basked shaped, with wide based, tapering pseudobulbs. Occasional
plants produce aerial roots; axes and leaves may exhibit purple pigment in areas
exposed to direct light. Leaf bearing axes 15-33 cm long, 2.1-3.1 cm wide at mid point,
round to oval in cross section, variably curved, 2-5.2 cm wide at base, 1.3-3.2 cm wide
at apex. Leaves terminal, 2-3, rigid to very rigid, concave, sub-elliptic to oblanceolate
or ovate, with a sheathing base, 14.8-24 cm long, 4.6-7.4 cm wide, concave, rigid,
1.2-2.0 mm thick. Racemes stout, 1-2, 21-47 cm long; peduncle 7-14.8 cm long,

Dendrobium speciosum

Telopea 11(2): 2006

209

o

Axis 1

O var. grandiflorum
■ Carnarvon Gorge

n

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A

o

Axis 1

Fig. 6. Ordination from PCoA in three dimensions of D. speciosum var. grandiflorum individuals
and individuals from along the Tropic of Capricorn using vegetative and floral morphological
characters, (a) Axis 1 v. 2. (b) Axis 1 v. 3. Character vectors (correlations>0.8) showing angles in
ordination space are shown. See Table 2 for character codes.

210

Telopea 1 1(2): 2006

Adams, Burke and Lawson

4.5-7 mm wide, with 1-4 medium bracts; rachis 14.5-31.5 cm long, 25-87-fIowered:
flowers well spaced. Pedicel including ovary 2.7-4.5 cm long. Flowers usually cupped
and of heavy substance, medium to large flowered, vertical height 5.1-6.6 cm, horizontal
width 5.5-6.3 cm, cream to gold, the labellum variously purple barred or spotted over
forelobe and midlobe, markings diffuse and variable. Dorsal sepal 2.8-3.9 cm long,
0.5-0.8 cm wide at base, tapering to an obtuse-rounded apex. Lateral sepals 2.1-2.5 cm
long, 0.8-1.1 cm wide at base, falcate-oblong, obtuse at apex. Petals 2.4-3.4 cm long,
2.8-4 mm wide, almost linear with an acute apex. Labellum 1.1-1.4 cm long, 1-1.3 cm
wide when flattened; forelobe 0.7-0.8 cm long, 1.1 -1.4 cm wide, incurved, subtriangular;
midlobe shortly clawed, 0.4-0.6 cm long, 0.9-1.0 cm wide when flattened, presenting
as a curved channel with a short acute apex; callus raised, 2-ridged, low profiled, yellow
to orange. Column c. 5 mm long with a foot c. 6 mm long at right angles to it. Mentum
bifid, rounded, c. 7 mm long from ovary to apex.

Flowering time: August-September.

Distribution: gorges of the Carnarvon region; geographically separate from other
varieties.

Etymology: the epithet refers to the Carnarvon Region.

Illustration: see Fig. 11.

Notes: plants of D. speciosum var. carnarvonense are usually lithophytic, in protected
sites along escarpments and rocky creeks. They are usually distinguishable from var.
blackdownense by more robust, tapering pseudobulbs and large flowers of heavier
substance. They are larger and more robust plants compared to var. capricornicum,
and their flowers, which open later, usually have wider segments. They most closely
resemble var. speciosum in plant form, flower shape and substance, as also indicated in
numerical analyses. Like var. capricornicum and var. blackdownense , new growths show
a strong development of purple anthocyanin pigment, which is not usually evident in
the mature leaves.

Descriptions of other varieties

These are based on measurements of 453 plants combined with field observations.
Occasional plants may be encountered with measurements outside of the ranges of
values presented here. Details of the type collection for recently described var. boreale
and neotypified var. curvicaule are provided.

Dendrobium speciosum Sm. var. speciosum, Exotic Bot. 1:17,1.10 (1804)

Plants variable, basket or urn shaped, with curved axes or more erect if axes less
curved. Usually lacking aerial roots, but occasional plants produce them; axes and
leaves exhibiting purple pigmentation in parts exposed to direct light. Leaf bearing
axes usually wide based and tapering towards the apex, 13-51 cm long, 2.1-4.5 cm
wide at mid point, round to oval in cross section, variably erect or curved, 2.8-6.0 cm
wide at base, 1.4-3.2 cm wide at apex. Leaves terminal, 2-5, rigid to very rigid, concave,
sub-elliptic to ovate, occasionally oblanceolate, relatively broad at the sheathing base,
9-20 cm long and 4-12 cm wide, 1-2 mm thick. Racemes 1-4, 14.5-57 cm long;
peduncle 5-16 cm long, 4.6-9.5 mm wide, with 1-4 medium sized bracts; rachis

Dendrobium speciosum

Telopea 11(2): 2006

211

CM

(/3

'x

<

Axis 1

Locality

Fairlies Knob
l> Kalpowar
■ Kroombit Tops
V Miriam Vale
A Yarwun
O Ulam
<1 Mt Morgan

Fig. 7. Ordination from PCoA of D. speciosum var. grandiflorum individuals using vegetative and
floral morphological characters. Axis 1 v. 2.

CN

c/)

$

Axis 1

Locality
■ Telegerry
• Gloucester
x BennysTops
Yarras

i> Hastings River
<1 Kempsey
V Urunga
A Dorrigo
O Tenterfield
O Rocky River

Fig. 8. Ordination from PCoA of D. speciosum var. hillii individuals using vegetative and floral
morphological characters. Axis 1 v. 2.

212

Telopea 1 1(2): 2006

Adams, Burke and Lawson

15-41 cm long, 18-115-flowcred: flower density varying from openly spaced to fairly
densely clustered. Pedicel including ovary 2.7-5.6 cm long. Flowers usually opening
moderately or widely, variable in substance, vertical height 4.2-8 cm, horizontal width
4.3-7.S cm, off white to deep gold, labellum variously and diffusely marked with purple
spots or short bars over forelobe and mid lobe. Dorsal sepal 2.5-4.6 cm long, 0.4-1.0 cm
wide at the base, tapering to an obtuse-rounded apex. Lateral sepals 1.8-3.9 cm long,
0.7-1.2 cm wide at base, falcate-oblong, obtuse at apex. Petals 2.2-4.1 cm long, 2-5 mm
wide, slightly falcate, with an acute apex. Labellum 1.1-1.7 cm long, 0.8-1.5 cm wide
when flattened; forelobe 0.6-0.9 cm long, 0.8-1.5 cm wide, incurved, subtriangular;
midlobe shortly clawed, 0.3-0.8 cm long, 0.6-1.2 cm wide when flattened, presenting
as a curved channel with a short acute apex; callus slightly raised, 2-ridged, yellow to
orange. Column 4-5 mm long with a foot 5-6 mm long at right angles to it. Mentum
bifid, rounded, 6-7 mm from ovary to apex.

Flowering time: August-October, or November in eastern Victoria.

Distribution: from Genoa, eastern Victoria (formerly at Cann River, but reportedly
extinct there now), north to Bulahdelah, Barrington Tops, New South Wales. Inland,
west 200 km to Munghorn Gap near Mudgee, New South Wales.

Notes: large robust plants which may form dense colonies covering whole rock faces on
the eastern escarpments of the Great Dividing Range. Occur from sea level to the tops
ol ranges in open and closed sclerophyll forest and rainforest. In rainforest it is also
lithophytic and less frequently an epiphyte, occasionally with aerial roots. Pseudobulbs
wide at base, often curved and tapering towards the apex. Flowers relatively large and
well spaced, including superior horticultural forms. Larger flowered forms occur in the

o

Locality

North-east
Victorian coast

t> Southern New
South Wales

<1 MacDonald River

* Wattagan Mountains

* Corabare Mountains

O Brokenback Range

' Northern New
South Wales

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W

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o

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A

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Axis 1

Fig. 9. Ordination from PCoA of D. speciosum var. speciosum individuals using vegetative and
floral morphological characters. Axis 1 v. 2.

Dendrobium speciosum

Telopea 11(2): 2006

213

Fig. 10. Dendrobium speciosum var. blackdownense: (a) whole flower, front view; (b) whole
flower, side view; (c) ovary, column and labellum, side view; (d) ovary and column, side view;
(e) labellum, top view; (f) flattened labellum, top view; (g) column, front view; (h) pollinium;
(i) anther, side view; (j) plant habit.

214

Telopea 11(2): 2006

Adams, Burke and Lawson

northern half of the range, from pure white to golden yellow. The variety is variable
in size, plant shape and floral dimensions, but less so than most of the more northerly
varieties, and intergrades with var. hillii primarily north of the Hunter River, New
South Wales.

Dendrobium speciosum Sm. var. hillii Mast., Gard. Chron. (new ser.) 7: 112, f. 18
(1877)

Plants variable, tall and upright in habit, commonly producing aerial roots, especially
prominent and erect in large rainforest plants. Leaf bearing axes 16-68 cm long,
1.8-5.2 cm wide at mid point, oval in cross section, may be considerably flattened,
variably erect or gently curved, 1.4-4.4 cm wide at base, 1.5-4.2 cm wide at apex. Leaves
terminal, 2-5, often very large, rigid, concave, sub-elliptic to oblanceolate, occasionally
ovate, relatively broad at the sheathing base, 13-28 cm long, 6-12 cm wide, 1-2 mm
thick. Racemes 1-4, 26-65 cm long; peduncle 6-13.6 cm long, 4-9.5 mm wide, with
1-4 medium sized bracts; rachis 20-51 cm long, 44-221-flowered: flower density
varying from openly spaced to densely clustered. Pedicel including ovary 2.3-4.6 cm
long. Flowers small to medium, opening variably, may remain cupped, usually light in
substance, vertical height 3.4-5.2 cm, horizontal width 2.9-5.3 cm, usually off white
to pale yellow, the labellum variously and diffusely marked with purple spots or short
bars over forelobe and midlobe. Dorsal sepal 1.9-3.7 cm long, 0.4-0.6 cm wide at the
base, tapering to an obtuse-rounded apex. Lateral sepals 1.4-2.7 cm long, 0.6-0.9 cm
wide at base, falcate-oblong, obtuse at apex. Petals 1.8-3.2 cm long, 2-3 mm wide,
slightly falcate, with an acute apex. Labellum 0.8-1.3 cm long, 0.7-1.0 cm wide when
flattened; forelobe 0.5-0.8 cm long, 0.7—1.0 cm wide, incurved, subtriangular; midlobe
shortly clawed, 0.3—0.5 cm long, 0.5-0.8 cm wide when flattened, presenting as a curved
channel with a short acute apex; callus slightly raised, 2-ridged, yellow to orange. Column

4- 5 mm long with a foot 4-6 mm long at right angles to it. Mentum bifid, rounded,

5- 6 mm from ovary to apex.

Flowering time: August-October.

Distribution: occurs, or formerly occurred, just south of the Hawkesbury River in
central eastern New South Wales, to Mt Mee-Crows Nest in southern Queensland,
where it begins to intergrade with var. grandiflorum.

Notes: in optimal moist rainforest habitat these plants are very large epiphytes with
pseudobulbs more than a metre in height, and forming masses in the canopy several
metres in diameter. They are much more abundant and larger in upland forest. In
more exposed and drier habitat, rainforest or open forests, they are smaller, and occur
as lithophytes or epiphytes. Recognizable by erect, relatively slender, virtually non¬
tapering pseudobulbs, relatively large leaves and long racemes crowded with many
small, white to cream, or occasionally pale yellow flowers. Aerial roots are less common
in more exposed sites. At the western limit and at the intergrade with D. speciosum,
plants may be squat, atypical, and with widely spaced large flowers which are difficult
to identify as this variety.

Dendrobium spedosum Telopea 1 1(2): 2006 215

Fig. 11. Dendrobium spedosum var. carnarvoncnsc : (a) whole flower, front view; (b) whole
flower, side view; (c) ovary, column and labellum, side view; (d) ovary and column, side view;
(e) labellum, top view; (f) flattened labellum, top view; (g) column, front view; (h) pollinium;
(i) anther, side view; (j) plant habit.

216

Telopea 11(2): 2006

Adams, Burke and Lawson

Dendrobiurn speciosum Sm. var. grandifloruttt F.M.Bailey, Bot. Bull., Dept Agric 14:12
(1896)

Plants variable, often epiphytic, with very large rainforest forms, commonly with well
developed aerial roots. Leaf bearing axes 17-95 cm long, 2.6-5.5 cm wide at mid point,
oval or flattened oval in cross section, variably erect or curved, 1.8-3.3 cm wide at base,
1.5-3.2 cm wide at apex. Leaves terminal, 2-6, often large, rigid, concave, sub-elliptic
to oblanceolate, with a short sheathing base, 12-37 cm long, 5-10 cm wide, 1-2 mm
thick. Racemes 1-4, short to very long, 25-70 cm long; peduncle 9-16 cm long, 4.5-
9.5 mm wide, with 1-5 medium sized bracts; rachis 16-54 cm long, 41-123-flowered:
flower density varying from openly spaced to densely clustered, small to large sized
including some of the largest in the species. Pedicel including ovary 3.2—5.7 cm long.
Flowers very variable in substance, usually opening widely, pale yellow to deep gold,
occasionally bicolour with a cream-white centre, and rarely pure white, vertical height
4.7-8.1 cm, horizontal width 4.8-8.0 cm, the labellum variously and diffusely marked
with purple spots or short bars over forelobe and midlobe. Dorsal sepal tends to be much
larger in relation to other segments, 2.7-4.7 cm long, 0.6-0.9 cm wide at base, tapering
to an obtuse-rounded apex. Lateral sepals 2.3-4.5 cm long, 0.8-1.1 cm wide at base,
falcate-oblong, obtuse at apex. Petals 2.4-4.2 cm long, 3-5 mm wide, slightly falcate,
with an acute apex. Labellum 1-1.6 cm long, 1.7-2.5 cm wide when flattened; forelobe
0.6-0.9 cm long, 1-1.4 cm wide, incurved, subtriangular; midlobe shortly clawed,
0.4-0.7 cm long, 0.7-1.1 cm wide when flattened, presenting as a curved channel with
a short acute apex; callus slightly raised, 2-ridgcd, yellow to orange. Column 5-6 mm
long with a foot 5-7 mm long at right angles to it. Mentum bifid, rounded, 6-8 mm
from ovary to apex.

Flowering time: August - October.

Distribution: including the interface with var. hillii, it occurs from Mt Mee-Crow’s
Nest, to the Mt Morgan area.

Notes: found in suitable rainforest habitat where it forms dense populations in the
canopy on many rainforest trees, and hoop pines (Araucaria cunninghamii Aiton ex
D. Don) and along rocky creeks and escarpments. Pseudobulbs may be almost one
metre long in the rainforest, and similar to, but often thicker than those seen in var.
hillii, with very large leaves in shady habitat. In more exposed situations as epiphytes or
lithophytes, plants may be only 20 cm high, with short, straight or curved pseudobulbs.
In the south of the range there is a gradual merging with var. hilli, where many plants
are short, smaller, or with pale yellow flowers. Deep yellow colour and large flowers with
prominent dorsal sepals help to distinguish only some plants of the variety. Unusual
coloured forms of yellow flowers with white centres are reported between Kroombit
Tops and Miriam Vale.

Dendrobiurn speciosum Sm. var. capricoruicum Clemesha, Orchadian 7: 103 (1982)

Plants variable, relatively short and compact, small to medium sized, usually upright
in shape and lithophytic, without aerial roots; axes and leaves exhibiting purple
pigmentation in parts exposed to direct light. Leaf bearing axes usually cylindrical,
7-19 cm long, 2-3.8 cm wide at mid point, round to oval in cross section, variably
erect or curved, 2-3.9 cm wide at base, 1.6-2.5 cm wide at apex. Leaves terminal, 2-5,

Dendrobium speciosum

Telopea 11(2): 2006

217

80 -i

T

Maximum

a

Mean

I

f to 3'°

interquartile range

Minimum

N = Number of individuals

-20

N= 39 37 35 13 17 8 43 46 52

Variety

Fig. 12. Variation in pseudobulb length of Dendrobium speciosum varieties. N = number of
individuals measured.

Variety

T

Mean

Maximum

I s ' to 3

interquartile range

Y

—Ii— Mini

Minimum
N = Number of individuals

Fig. 13. Variation in leaf length of Dendrobium speciosum varieties. N = number of individuals
measured.

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Adams, Burke and Lawson

very rigid and coriaceous, often channelled, sub erect, ovate to oblanceolate, with a
short sheathing base, 5-16 cm long, 4.5-6 cm wide, 1-3 mm thick. Racemes arching or
erect, 1-2, 17-49 cm long; peduncle short to long, 9-22 cm long, 5-7 mm wide, with
1-3 medium sized bracts; rachis 8-27 cm long, 11-68-flowered; flower density varying
from openly spaced to clustered. Pedicel including ovary 2.7-4.4 cm long. Flowers
usually opening widely, very variable in substance, vertical height 3.4-5.9 cm, horizontal
width 3.9-5.6 cm, off white to deep gold, the labellum variously and diffusely marked
with purple spots or short bars over forelobe and midlobe. Dorsal sepal 2.1-3.4 cm
long, 0.5-0.8 cm wide at the base, tapering to an obtuse-rounded apex. Lateral sepals
1.6-2.9 cm long, 0.7-1.1 cm wide at base, falcate-oblong, obtuse at apex. Petals
1.9-3.0 cm long, 3-5 mm wide, slightly falcate, with an acute apex. Labellum
1-1.6 cm long, 1-1.2 cm wide when flattened; forelobe 0.6-0.9 cm long, 1-1.2 cm wide,
incurved, subtriangular; midlobe shortly clawed, 0.4-0.7 cm long, 0.8-1.1 cm wide
when flattened, presenting as a curved channel with a short acute apex; callus slightly
raised, 2-ridged, yellow to orange. Column 4—5 mm long with a foot 5-6 mm long at
right angles to it. Mentum bifid, rounded, 6-7 mm from ovary to apex.

Flowering time: May-August.

Distribution: Mt Morgan area to Byfield, and west to Berserker Range.

Notes: in east Capricorn, localised colonies with short and very coriaceous leaves are
restricted to suitable habitat on volcanic plugs, and in forest and creek side locations
at Byfield. On volcanic plugs, plants are exposed to full sun, and only their leaves may
be evident between rock crevices. The bulk of the pseudobulbs arc protected between
layers of stone. In the northern part of the range occasional colonies occur with a wide
variety of plant form, and well spaced yellow to gold flowers.

Dendrobium speciosum Sm. var. curvicaule F.M.Bailey, Bot. Bull, Dept Agric. 14:12
(1896)

Neotype (Adams et al. 2006a): Queensland. Kennedy North District: Cathu State
Forest, P.B. Adams 6' S.D. Lawson, 27 Jun 1993 (holo: QRS; isoneo: BRI, MEL).

Brief Neotype Diagnosis: Leaf-bearing axes short to medium (10.7-52 cm long), collum
usually inconspicuous, peduncle short (7.5-16 cm long), flowers moderately spaced to
crowded, off white to bright yellow, tepals variably long, lateral sepals broad (0.8-1.2
cm wide), variably incurving.

Plants very variable in shape and size, aerial roots prominent in occasional plants.
Leaf bearing axes curved, fusiform or linear, 10.7-52 cm long, 1.7-5.1 cm wide at
midpoint, with an indistinct or absent collum at base, if collum present then widening
distally over only 1-2 cm, axis at base 1.3-4.0 cm wide, round to oval in cross section.
Apex of axes usually narrowing to 1.4-3.4 cm wide, variably sharply edged only in
the distal few centimetres. Leaves terminal, 2-4, 15-26.6 cm long, 4-9.2 cm wide, leaf
shape variable, sub-elliptic to oblanceolate with a short sheathing base, concave, rigid,
1-2.7 mm thick, thinner, larger and more flexible leaves in dense shade, smaller and
thicker when exposed to sun. Racemes 1-3, 20-65 cm long, peduncle 7.5-16 cm
long, 3.7-7.9 mm wide, shorter than the rachis with prominent bracts; rachis 12.5—
45.5 cm long, bearing 20-135 flowers, open or very closely spaced and overlapping,
pedicel including ovary medium to very long, 2.2-5.2 cm long. Flowers small to

Dendrobium speciosum

Telopea 11(2): 2006

219

U) 6 '
c
0)

i— 5
tu 3
5
o

LL 4'

3-

II

T

Pi

T

38 37

35 13

%

\ X 4

\ \ '%
\

**» V ^

Mean

I

X

Maximum

I s 'to 3 rd

interquartile range

Minimum

N = Number of individuals

43 46 52

* \ %

Variety

Fig. 14. Variation in flower length of Dendrobium speciosum varieties. N = number of individuals
measured.

Fig. 15. Distribution of taxa of Dendrobium speciosum Smith.

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Adams, Burke and Lawson

large, usually opening widely but sometimes cupped, 4—7.1 cm long vertically,
4.1-7 cm wide horizontally, off white to deep yellow segments with heavy substance,
labellum white to cream, variously marked with purple spots or bars on forelobe
and midlobe. Dorsal sepal 2.2-4.1 cm long, 0.6-1.0 cm wide, tapering from base to
an obtuse-rounded apex. Lateral sepals 1.4-2.5 cm long, 0.8-1.2 cm wide, falcato-
oblong and obtuse at apex. Petals relatively long 2.1-3.6 cm long, 3-5 mm wide,
almost linear and ending acutely. Labellum 0.9-1.5 cm long, 1-1.3 cm wide, forelobe
0.6-0.9 cm long, 1-1.3 cm wide when flattened, incurved and subtriangular, midlobe
on a short claw, midlobe 0.3-0.6 cm long, 0.7-1.6 cm wide when flattened, presenting
as a curved channel with short acute apex. Column c. 5 mm long with a foot 5-6 mm
long at right angles to it, mentum bifid, rounded, 6-7 mm from ovary to apex. Callus
raised, 2-ridged, low profiled, orange coloured.

Distribution: Mt Dryander north east of Proserpine, to St Lawrence, and the
Whitsunday Islands.

Flowering time: August-September.

Notes: previously more broadly circumscribed to include plants treated here as var.
boreale (Adams et al. 2006a). Predominantly a rainforest epiphyte from sea level to
1200 metres, on large trees such as Tulip Oak (Argyrodetidron actinophyllum (Bailey)
Edlin) and Red Cedar ( Toona australis (F. Muell.) Harms), often as very large plants
over two metres in diameter, usually high in the canopy, where populations may be
very dense. Smaller, shorter plants are found in more exposed open forest as epiphytes
or lithophytes, and along watercourses on Casuarina L. ex Adans. Short diminutive
plants are found at sea level on the Whitsunday Coast. They have been discussed and
illustrated by Dockrill (1992, pp. 462—463), who considered it a possible sub coastal
variety. These forms represent atypical ecotypes comparable to those that are often seen
at the eastern and western habitat limits for most other varieties (see also var. hillii ).
South of Sarina, colonies are isolated, and form an intergrade with var. capricornicum
along dry watercourses. Flowers are usually cream, but may be deep yellow or gold.
They may be small or large, with broad segments, and represent some of the finest
horticultural forms of the species.

Dendrobium speciosum Sm. var. boreale P.B.Adams, J.M.Burke & S.D.Lawson [ms.]
Aust. Syst. Bot. (2006a)

Type: Queensland. Cook District: Windsor Tableland, B. Gray, P.B. Adams, J.M. Burke
dr S.D. Lawson, 3 Aug 2004, (holo: QRS; iso: BR1, MEL).

Brief Diagnosis: Leaf-bearing axes short to long (12.6-71 cm), rarely curved; collum
usually prominent; peduncle moderately long (8-29 cm) but not longer than rachis;
flowers well spaced; tepals off white to pale yellow, broad in relation to length; lateral
sepals moderately broad (0.7-1.0 cm) with little incurving.

Plants very variable in shape and habit, occasionally producing aerial roots. Leaf¬
bearing axes 12.6-71.0 cm long, 1.4-4.6 cm wide at mid point, with little tapering
or shorter and fusiform, usually with a prominent collum at base 1.3-4.4 cm long,
axis at base 1.3-2.7 cm wide, oval or flattened in cross section, erect or curved or
gently angled once or twice in distal half, sharply edged over distal half (especially
notable in axes of taller plants 1-2 years old), tapering or broadening towards apex

Dendrobium speciosum

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221

Fig. 16. Dendrobium speciosum var. capricornicum: (a) Rockhampton area; (b) north of Byfield.
Dendrobium speciosum var. blackdownense: (c) densely flowered gold form; (e) open raceme
form; (f) off white form, (d) Dendrobium speciosum var. carnarvonense, Carnarvon Gorge. Scale
bars: (a), (c) = 10 cm; (b), (f) = 5 cm; (d), (e) = I cm.

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Adams, Burke and Lawson

1.2-4.2 cm wide. Leaves terminal, erect and narrower in shorter forms, 2-4, 9-34 cm
long, 5-11 cm wide, sub-elliptic to oblanceolate with a sheathing base, concave, rigid,
1-2.6 mm thick, relatively thin, large, and flexible in dense shade, relatively small
and thick when exposed. Axes and leaves sometimes showing purple pigment in
areas exposed to strong light. Racemes 1-3, 18-76 cm long; peduncle 8-29 cm long,
3-8 mm wide, shorter or no longer than the rachis, with 1-5 prominent bracts; rachis
10-48 cm long, with 9-125 star-like flowers openly spaced and displayed. Pedicel
including ovary 2.5-5.3 cm long. Flowers usually opening widely, 3-5 cm long
vertically, 3.3-5.5 cm wide horizontally, off white to cream, occasionally pale yellow,
with purple spots or short bars marking the forelobe and midlobe of the labcllum;
tepals presenting with wide bases and heavy substance. Dorsal sepal 1.8-3.0 cm long,
0.55-0.9 cm wide at base, tapering from base to an obtuse-rounded apex. Lateral
sepals 1.3-2.1 cm long, 0.7-1.0 cm wide at base, falcato-oblong, slightly incurved,
obtuse at apex. Petals 1.6-2.7 cm long, 2-5 mm wide, almost linear, ending acutely
at apex. Labellum 1.1-1.7 cm long, 1-1.3 cm wide; forelobe 0.7-1.0 cm long,
1-1.3 cm wide when flattened, incurved, subtriangular; midlobe on a short claw,
0.4-0.7 cm long, 0.6-1.0 cm wide when flattened, presenting as a curved channel with
a short acute apex. Column c. 5 mm long, with a foot c. 6 mm long and at right angles
to it. Mentum bifid, rounded, c. 7 mm long from ovary to apex. Callus raised, 2-ridged,
low profiled, yellow to orange.

Flowering time: July-September, usually later than var. pedunculatum.

Distribution: Annan River, south to Mt Elliot (south of Townsville).

Etymology: the epithet alludes to the most northerly distribution of D. speciosum
(Latin: boreale = northern).

Notes: in Adams et al. (2006a) we reviewed D. speciosum var. curvicaule and showed it
to be a name of uncertain application, and described var. boreale for plants north of Mt
Elliot, which were previously attributed to var. curvicaule. Lithophytic or epiphytic in
and around rainforest, very variable in plant size but commonly tall in densely shaded
rainforest gullies, shorter plants lithophytic in more exposed sites and along rocky
creeks with similar floral characters, often associated with small shrubs and moss beds.
Occurs east of the Great Dividing Range from almost sea level to over 1200 metres.
Plants may form dense colonics or be scattered. Till rainforest forms and short forms
are often only a few metres apart at the forest edges. Flowers are similar in both forms,
and not greatly variable compared with other varieties.

There is a small degree of overlap with var. curvicaule in morphology and numerical
characteristics. Dendrobium speciosum var. boreale is distinguished from typical var.
pedunculatum in drier habitats, the latter having smaller pseudobulbs, shorter racemes
and usually smaller flowers with more rounded segments.

In drier rainforest and sclerophyll forests e.g.Tinaroo Dam, Paluma, Bluewater, Mt Elliot,
Hidden Valley and Tully areas, smaller plants show floral and vegetative characteristics
that are intermediate in morphology between rainforest forms and var. pedunculatum.
These forms are interpreted as shorter variants of var. boreale. The PCoA ordination
indicates overlap between var. boreale and var. pedunculatum in a continuum (Fig.
4). The intermediate forms have variable peduncle length, from shorter to longer in
relation to rachis. Axes of var. boreale are not significantly curved unless growing on
vertical surfaces or in dense shade.

Dendrobium speciosum

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223

Fig. 17. Dendrobium speciosum: (a) var. boreale, cream flowers, Mt Finnigan, Queensland; (b)
var. boreale , white flowers, Paluma, Queensland, (c) var. grandiflorum, gold form, Miriam Vale,
Queensland; (d) var. curvicaule, Crediton, Queensland; (e) intermediate form from the boundary
of var. grandiflorum and var. hillii territory, Blackbutt, south Queensland; (f) var. hillii , west of
Wauchope, New South Wales; (g) var. pedunculatum, west of Mareeba, Queensland; (h) var.
speciosum , Kangaroo Valley, New South Wales. Scale bars: (a)-(f), (h) = 5 cm; (g) = 1 cm.

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Adams, Burke and Lawson

At Paluma and Hidden Valley some plants are shorter, 20-40 cm, with narrower
cylindrical axes and small rounded white flowers up to 90 per raceme. To the north there
is a gradation towards the typical rainforest form of var. boreale, as the habitat becomes
wetter and the rainforest canopy denser. Southwards to Townsville and beyond, where
populations are more isolated and close to sea level, they may be diminutive with a less
distinct or absent collum.

Dendrobiutn speciosum Sm. var. peduiiciilatum Clemesha, Orchadiati 6: 261 (1981)

Plants variable, usually short, compact lithophytes without aerial roots. Leaf¬
bearing axes 4.1-36 cm long, 1.8-4.4 cm wide at midpoint, round to oval in cross
section, collum usually absent, 1.2-3.7 cm wide at base, erect or mildly curving.
Apex of axes usually narrowing to 1.1-2.7 cm wide. Leaves terminal, tending
erect, 2-4, 6.3-19.3 cm long, 3.2-8.6 cm wide, leaf shape variable, sub-elliptic to
ovate, relatively broad at the sheathing base, very coriaceous and rigid, concave,
1.1-2.5 mm thick. Axes and leaves exhibiting purple pigmentation in parts exposed to
direct light. Racemes 1-2,15.9-56.9 cm long, peduncle longer or shorter than the rachis,
8.4-30.7 cm long, 2.2-5.7 mm wide with small bracts; rachis 7.5-26.2 cm long, with
9—72 flowers, open or very closely spaced and overlapping forming a brush. Pedicel
including ovary medium to very long, 1.9-4 cm long. Flowers usually opening widely
or cupped, 2.8-4.5 cm long vertically, 3.1-4.7 cm wide horizontally, off-white to yellow
segments of heavy substance, labellum white to cream, variously marked with purple
spots or bars on forelobe and midlobe. Dorsal sepal 1.8-2.7 cm long, 0.45-0.78 cm wide
at base, tapering from base to an obtuse-rounded apex. Lateral sepals 1.3-2.1 cm long,
0.7-1.1 cm wide, falcato-oblong and obtuse at apex. Petals 1.5-2.5 cm long,
2-3.8 mm wide at base, almost linear and ending acutely. Labellum 0.9-1.6 cm long,
0.9-1.3 cm wide, forelobe 0.6-0.9 cm long, 0.9-1.3 cm wide when flattened, incurved
and subtriangular; midlobe on a short claw, 0.3-0.6 cm long, 0.65-1.0 cm wide when
flattened, presenting as a curved channel with short acute apex. Column c. 5 mm long
with a foot c. 6 mm long at right angles to it, mentum bifid, rounded, 6-7 mm long
from ovary to apex. Callus raised, 2-ridged, low profiled, orange colour.

Flowering time: July-September.

Distribution: Parker River headwaters to Lumholz National Park, south of Atherton
Tableland, representing a narrow strip of open forest. The distribution is smaller than
previously considered, and habitat has been reduced by altered land use.

Notes: occurs on and to the west of the Great Dividing Range in northern Queensland.
It includes some plants informally described as “var. compaction". Plants are scattered
diffusely along rocky exposed watercourses, cliff faces and escarpments, and occasionally
found on the base of small trees. Leaves tend to be very coriaceous, and may exhibit
purple anthocyanin pigment. There is usually only one raceme per pseudobulb, which
is apical, with well spaced (occasionally closely) pale cream to yellow flowers of heavy
substance. The peduncle is often as long as or longer than the rachis, but forms are
found with much shorter peduncles.

Dendrobium spetiosum

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225

Variation of characters within and between varieties

The variation in pseudobulb, leaf and flower length for plants assigned to varieties
across the geographic range is presented in Figs 12-14, which provide the median, first
to third interquartile range, and extremes in the data set. Other characters show similar
patterns of continuous variation within and between described varieties.

Distribution of Dendrobium speciosum in major regions of eastern Australia

Our explorations indicate an almost continuous north to south distribution between
the Annan River south of Cooktown (Queensland) and Genoa (eastern Victoria), with
the qualification that small populations in marginal habitat may be up to 50 km apart.
Small populations occur in locations previously considered to be unsuitable habitat.
These include areas between Rockhampton and Gladstone, between Rockhampton
and Sarina, and dry forests 100 km north and south of Townsville. We indicate the
approximate limits of the varieties confirmed in this paper in Fig. 15.

Field surveys concentrated near the geographic limits of varieties revealed areas where
populations exhibited characteristics intermediate between the two adjacent varieties.
Within these populations we could not identify typical individuals of the varieties
admixed with intermediate forms, or the presence of sporadic hybrids. Each population
subtly varied from the next, forming an intergrade or continuum.

Between adjacent varieties along the Great Dividing Range there are plants with
intermediate characters occupying territory generally less than 150 km apart. The
following areas of intermediate forms were identified in relation to Fig. 1. There is
a narrow zone between Regions la and lb, between rainforest and dry, exposed
cliff faces (var. pedunculatum-var. boreale interface). Between Regions 2 and 3, the
interface between var. curvicaule and var. capricomicum, scattered populations occur
in dry country. A narrow zone of populations with intermediate characteristics is
recognized between Regions 3 and 5 (Mt Morgan area) at the var. capricomicum and
var. grandiflorum interface. There is a broad zone of around 150 km between Regions 5
and 6, which extends north from a line between Mt Mee and Crows Nest, and includes
Blackbutt, Gallangowan, Conondale and Kilkivan. This represents the var. grandiflonim
and var. hillii interface. Between Regions 6 and 7, the interface between var. hillii and
var. speciosum forms a band between Bulahdelah and Barrington Tops, which extends
south-west to Munghorn Gap near Mudgee.

Discussion

Interpretation of analyses

This extensive study of variation indicates that D. speciosum is a single variable species
differing mainly in size characteristics, occurring in widely varied habitats. There are
intergrading groups best interpreted as varieties, and no discrete qualitative differences
between any populations or regions. Some plants do not fit exactly with any varietal
description and are most satisfactorily designated by geographical notation. Based on
UPGMA cluster analysis, the most distinct varieties are var. boreale (Region la) and var.
hillii (Region 6).

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The variation within this species complex indicates morphological forms with a
geographic pattern that is largely allopatric. Multivariate analyses of D. speciosum
could not demonstrate a phenetic gap between any identified groups, all of which
overlapped in ordination space to varying degrees. Plants described as varieties may
overlap in ordination space, but in the areas of interface between two varieties they are
not identifiable as admixtures with intervarietal hybrids. These populations consist of
plants which all show intermediate characteristics.

The addition of plants from newly surveyed areas, and interpretations of groups as
three new varieties, one in northern Queensland and two in central Queensland, has
expanded the recognised level of variation. The areas of greatest complexity and most
difficult interpretation occur in north Queensland, and central Queensland along the
Tropic of Capricorn, as demonstrated in the regional ordinations (Figs 4-6).

Distribution of Taxa

The distributional range of taxa within D. speciosum has been difficult to define due
to wide inherent variation, zones of intermediate forms, and significant loss of habitat,
particularly at the var. speciosum/ var. hillii and var. hillii/ var. grcmdiflorum interfaces.
For the six most northerly varieties our surveys have clarified the distributions. Habitat
loss is not confounding, except at the interface of var. pedunculatum and var. boreale on
the Atherton Tableland.

The distribution is much more continuous than previously considered, especially
north of Rockhampton; The gradation between var. curvicaule and var. capricornicum
between Clairviewand Byfield represents the largest north-south zone of intermediates.
The more southerly intermediate zones between var. grandiflorum and var. hillii, and
between var. hillii and var. speciosum have been noted by previous reviewers, who
have highlighted the difficulty of assigning these plants to varieties. Gregory (1988)
noted that varieties “gradually merge into one another up and down the Great
Dividing Range”. Jones (1988 p. 478) stated that “confusing intermediate forms may
occur where ranges (of varieties) overlap”. Dockrill (1992, vol. 1 p.457) noted that “all
varieties within themselves are quite variable, and merging of intermediates.... made
it virtually impossible to give a satisfactory determination of a particular clone.” Banks
and Clemesha (1990) noted intermediate areas where it was “difficult to say where one
stops and another starts”.

For the var. hillii and var. speciosum interface, the intermediate zone requires more
exploration at the western limit. There has been considerable habitat loss and plant
collection, especially south of the Hunter River. Reports of plants attributed to var hillii
from Peat’s Ridge (Clemesha 1986), and from the Wattagan area, cannot be conclusively
evaluated. Our recent surveys on the coastal ranges south of Newcastle, including the
Wattagan Mountains, indicated only plants attributable to var. speciosum.

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227

Biogeography

In north Queensland, medium to large plants of var. boreale have the largest distribution,
in wet tropical rainforest (Region la). In Region la there are also shorter forms of var.
boreale, usually in intermediate sclerophyll forest habitat on margins of rainforest. In
the dry zone north and south of Townsville, shorter plants attributed to var. boreale
are found in small isolated populations, and a similar plant form is found in isolated
colonies between Sarina and Rockhampton.

Variety pedunculatum is a short plant form occupying a very narrow strip of dry
country on or west of the Great Divide, on the fringe of var. boreale territory, from west
of the Windsor Tableland to Lumholz National Park (Region lb). The distribution
area of var. pedunculatum is extremely small in relation to that of var. boreale (Fig.
15). The type form of var. pedunculatum from Bone’s Knob is in this narrow strip, and
one of the major clarifications of this study is the designation of shorter plants east
of the divide in and around rainforest as variants of var. boreale. Many of these were
previously regarded as forms of var. pedunculatum. They are usually differentiated by
long upright racemes with many small to medium sized white or pale cream flowers
which have wide segments, and their peduncles are variable, often short in relation to
rachis length.

On volcanic plugs near Rockhampton, short plants meeting the description of var.
capricornicum (Clemesha 1982) intergrade with forms similar to those in the southern
area of Region 2 (var. curvicaule). West to Blackdown Tableland and Carnarvon Gorge,
two new taxa are described on the basis of morphological characteristics and regional
PCoA analyses, var. blackdownense and var. carnarvonense. These groups abut in
ordination space and are geographically distinct, with a small amount of overlap in
numerical analyses with plants of Regions 2,3 and 5 (Figs 3,5 and 6). Clemesha (1982)
and Walsh (2000) commented that some plants of var. capricornicum may resemble var.
grandiflorum. We consider from repeated observations that var. carnarvonense and var.
blackdownense are clearly distinguishable from var. gratidiflorum.

In Region 5, primarily consisting of var. gratidiflorum, there is a fairly abrupt narrow
interface between var. capricornicum and var. gratidiflorum in the north, determined by
a limit of moister rainforest south of Rockhampton. There is a considerable gradation
between var. grandiflorum and var. hillii at the boundary of Regions 5 and 6, the var.
gratidiflorum at the southern limit being usually shorter in stature, with smaller flowers.
Around Gladstone, tall forms in dry rainforest and shorter forms on volcanic plugs are
found with large gold flowers.

Plants of Regions 7 and 8 (var. speciosum) are easily identified by their locality, but
overlap considerably with populations from other regions in the ordination (Fig. 3).
Skilled observers are usually able to identify them from their vegetative and floral
form, except for the area of intermediate forms at the var. speciosum and var. hillii
interface. The associations of populations with habitat assist in designation of varieties
in Region 1 and on the volcanic plugs of Region 3 (Table 4), and suggests that much of
the variation is ecophenetic.

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Table 4. Associations of varieties with habitat in regions.

Region

Variety

Rainforest

Sclerophyll Forest, or Riparian

Rocky Plugs

la

boreale

+

+

-

1b

pedunculatum

-

+

-

2

curvicaule

+

+

-

3

capricornicum

-

+

+

4

blackdownense

-

+

-

4

carnarvonense

-

+

-

5

grandiflorum

+

+

+

6

hillii

+

+

-

7, 8

speciosum

+ +

Recommended taxonomy

There are

no generally

accepted criteria

or definitions for grouping

or ranking

botanical taxonomy, especially at specific and infraspecific level. Subspecies, varieties,
subvarieties and forms have been used in many ways to interpret degrees of difference.
More recent practice has tended to treat intergrading taxa, and even subgroupings
within some dines, as subspecies. Stuessy (1990, p. 193) attempted to define levels of
difference for subspecies, varieties and forms, and recommended that existing ranks be
retained in the interest of nomenclatural stability for groups with a history of taxonomic
study, and that subspecies could be preferred for newly studied groups. Choice of rank
remains an individual decision of taxonomists based on their understanding of a
group.

In taxonomic studies of Dendrobium, Schlechter (1982) arranged individual species as
naturally as possible within the genus, and did not provide a species definition. Dockrill
(1969,1992) discussed the difficulties of interpretation of sub specific taxa and species,
questioning the proposals of Clements (1989) to upgrade varieties to species, and
disputing the proposal to elevate varieties of D. speciosum. He considered it important
to take into account similarities as well as differences (Dockrill 1995). We have found no
clear-cut qualitative differences between regional subgroups of D. speciosum, with the
possible exception of the earlier but still variable flowering time for D. speciosum var.
capricornicum, and thus have not used specific rank, as proposed by Clements (1989)
and Clements and Jones (2002). In this study we experienced difficulty in defining taxa
in a large data set encompassing all major variation, even with numerical techniques
that maximise separation.

The following reasons are advanced for the decision to interpret the variation as
representing varieties in a species complex.

1. We understand the species as having a clinal or continuum of variation, with
features running north-south along the Great Dividing Range, and east-west along the
Tropic of Capricorn. We describe nine sub taxa within the species, but within each
there are atypical forms and considerable infra taxa variation, all characters varying
continuously over the geographic range. We could find no convincing evidence to
support the interpretation of taxa at specific level.

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Telopea 11(2): 2006

229

2. Analyses and observations did not demonstrate any qualitative differences for any
sub group. This interpretation is in accord with the view of Stuessy (1990) that varieties
may have one to few conspicuous differences, and subspecies several differences. Bayly
and Kellow (2006) have made similar distinctions in their use of subspecies and varieties
for Hebe Jussieu.

3. All subgroups overlap to a significant extent in ordinations.

4. Near the geographic limits of varieties there are populations of plants identifiable as
intermediates.

5. Despite disjunct distributions we found that var. carnarvonetise and var.
blackdownense have a similar level of differences from other varieties, and consider
they are best regarded as varieties.

6. A satisfactory key to groups cannot be constructed, even for 70% (estimated) of
plants, similar to the opinion of Dockrill (1992).

7. The rank of variety maintains nomenclatural stability for a species with a long
history of description and classification. Dendrobium speciosum is one of the most
spectacular horticultural plants and a species widely discussed by orchid taxonomists
and enthusiasts. There is a need for infraspecific classification to aid communication
concerning variation over a very large distribution range. The rank ot variety has been
effective for communication, and we concur with the guidelines proposed by Entwisle
and Weston (2005) of minimal taxonomic change, especially in plant groups of high
interest.

A summary of the proposed taxonomy for D. speciosum is presented in Table 7. Examples
of varieties and forms illustrated in Figs 16 and 17 represent a small part of the total
variation in plant and floral characteristics in the respective regions.

Table 7. Taxonomic nomenclature for Dendrobium speciosum.

Region No.

1 .

2 .

3.

4.

5.

6 .

7.

Region

North Queensland
Whitsunday

Taxa

var. boreale
var. pedunculatum

var. curvicaule

East Capricorn
(central Queensland)

West Capricorn
(central Queensland)

var. capricomicum
atypical individuals and populations
intermediate between var. capricomicum and
var. curvicaule, and var. capricomicum and
var. grandiflorum specified by locality

var. blackdownense
var. camarvonense

Southern Queensland

var. grandiflorum

intermediates with var. hillli specified by locality

Southern Queensland and var. hillii

Northern New South Wales intermediates with var. speciosum specified
by locality

Southern New South Wales var. speciosum

8.

Far south New South Wales var. speciosum
to Victoria

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Telopea 11(2): 2006

Adams, Burke and Lawson

Most of the plants of var. hillii fit Masters’ (1877) description, except at the distribution
limits. Plants from eastern Victoria to the Hunter River form a cohesive group that can
be ascribed to var. speciosum. Dendrobium speciosum var. grandiflorum is useful to refer
to plants between southern Queensland and the Gladstone area, but at the southern
boundary with var. hillii, intermediates are shorter with smaller (lowers, and require a
locality designation.

Plants of central Queensland may meet descriptions of var. ccipricormcum in the
east on some volcanic plugs, but others in forest and on other plugs form a variable
group. This finding extends the field observations made by Walsh (2000). He noted a
resemblance of floral characteristics to var. grandiflorum, in particular the gold flower
colour. Clemesha (1981a) described some west Capricorn plants as a dwarf race of var.
grandiflorum. Our analyses do not support a close relationship to var. grandiflorum.
The Carnarvon plants have shorter, curving robust pseudobulbs like var. speciosum, but
clearly differ in floral characteristics consistent with a separate entity.

Taxonomic approaches in Australian Orchidaceae

The purpose of all classificatory systems is to advance the understanding of individuals,
their variation and relationships, and to promote ease of communication when
discussing them. Orchids are a difficult taxonomic group, and Australian orchids with
large distributions over diverse habitat pose particular problems in achieving adequate
sampling for assessment of their variation, and consideration of taxonomic rank. We
contend that several dcndrobiums, including D. speciosum and D. kingianum, are
species complexes with various levels of continuous and sometimes discontinuous
variation (Adams et al. 2006b). Within these complexes there may be sub groupings
which intergrade and cannot be easily differentiated. If a complex is not optimally
sampled, sub groups may appear to represent distinct taxa.

Subjective assessment of gross morphology is difficult in these complexes, as noted in
the early phases of classificatory systems of Locke, Ray, du Buffon, Linnaeus (Huxley
2003). Non-morphological methods such as biochemistry and DNA analyses could
be used to form a data set for analysis and interpretation to recognise taxa. At the
infraspecific level, these may be limited in usefulness or practicality. For D. speciosum
we have found that all varieties are compatible in cross pollination, giving rise to inter
varietal seedlings; bee pollination was not variety specific (Adams 1991, Adams &
Lawson 1993), and fragrance biochemistry did not provide useful taxonomic characters
(Adams 1991). An alternative approach for users is to specify plants by their original
location, or to use ordination and other numerical representations as has been proposed
for infraspecific taxa by Prentice (1986).

Species complexes like D. speciosum are best categorised by considering the full range
of variation and accurately describing similarities and partial differences between
populations. Intermediates and atypical variants must be described as such, and
not ignored or categorised arbitrarily. The value of naming systems, given their
inherent limitations, is appreciated in how effectively they establish conventions for
communication and development of ideas (Pearson 2001).

Dendrobium speciosum

Telopea 11(2): 2006

231

Acknowledgments

State National Parks and Wildlife Services allowed access to plants for this study.
We thank Prof. Pauline Ladiges for constructive comments on earlier drafts of the
manuscript, and Dr Jim Itoss and Neville Walsh (National Herbarium, Royal Botanic
Gardens, Melbourne) for assistance with taxonomy and Latin diagnosis. Stuart
Gardner provided technical assistance. Peter Weston contributed helpful discussions
and comments.

References

Adams PB (1991) Variation, multiple pollinators and breeding system in Dendrobium speciosum
Smith : A biological review. Orchadian 10: 124-140.

Adams PB, Burke )M & Lawson SD (2006a) Dendrobium speciosum Sm., Sect. Dendrocoryne:
Orchidaceae in North Queensland, numerical taxonomy and descriptions of two taxa. Aus¬
tralian Systematic Botany (2006 in press).

Adams PB, Burke JM 8c Lawson SD (2006b) Systematic analysis of Dendrobium Schwartz section
Dendrocoryne in the Australian region. Plant Systematics and Evolution (2006 in press).

Adams PB 8c Lawson SD (1993) Pollination in Australian orchids: A critical assessment of pub¬
lished reports 1882-1992 .Australian Journal of Botany 41: 553-75.

Bailey FM (1896) Contributions to the Queensland Flora. Botany Bulletin, Department of Agri¬
culture, Queensland 14:12.

Banks DP & Clemesha SC (1990) Dendrobium speciosum : a review of the species. Australian
Orchid Review 55: 4-14.

Bayly M) & Kellow AV (2006) An Illustrated Guide to New Zealand Hebes. (Te Papa Press: Wel¬
lington)

Belbin L (1988) ‘PATN Analysis Package’, CS1RO Division of Wildlife and Rangeland Research,
Canberra, Australia.

Burke JM (1998) A numerical analysis of Dendrobium speciosum Smith and section Dendro¬
coryne (Orchidaceae). (Unpublished MSc Thesis: James Cook University of North Queens¬
land)

Burke JM 8c Adams PB (2002) Variation in the Dendrobium speciosum (Orchidaceae) complex: a
numerical approach to the species problem. Australian Systematic Botany 15:63-80.

Clements MA (1989) Catalogue of Australian Orchidaceae. Australian Orchid Research 1: 1-62.

Clements MA 8c Jones DL (2002) Nomenclatural changes in the Dendrobieae (Orchidaceae) 1:
The Australasian region. Orchadian 13:485-497.

Clemesha SC (1981a) A review of Dendrobium speciosum Sm. Orchadian 6: 247-262.

Clemesha SC (1981b) Dendrobium speciosum var. bancroftianum. Orchadian 6: 271-272.

Clemesha SC (1982) A new variety of Dendrobium speciosum Smith from Central Queensland.
Orchadian 7: 103-106.

Clemesha SC (1986) Dendrobium speciosum - an update. Orchadian 8:173-178.

Crisp MD 8c Weston PH (1993) Geographic and ontogenetic variation in morphology of Aus¬
tralian Waratahs ( Telopea: Proteaceae). Systematic Biology 42:49-76.

Dockrill AW (1969) Australian Indigenous Orchids, vol. 1. (Society for Growing Australian Plants:
Sydney)

Dockrill, AW (1992) Australian Indigenous Orchids, vol. 1,2nd edition. (Surrey Beatty and Sons
Pty. Ltd., Chipping Norton)

Dockrill AW (1995) Some thoughts on taxa and taxonomic categories (particularly species) of
orchids. Orchadian 11: 292-294.

Entwisle TJ 8c Weston P (2005) Majority rules, when systematists disagree. Australian Systematic
Botany 18: 1-6.

Gould SJ 8c Johnston RF (1972) Geographic variation. Annual Review oj Ecology and Systematics
3: 457-98.

232

Telopea 11(2): 2006

Adams, Burke and Lawson

Gower JC (1966) Some distance properties of latent root and vector methods used in multivari¬
ate analysis. Biomctrika 53: 325-338.

Gregory T. (1988) Some thoughts on Dendrobium speciosum. Australian Orchid Review 53: 17-
26.

Grundon NJ, Wallace BJ & Harrison M (1990) Review: M. A. Clements Catalogue of Australian
Orchidaceae. Orchadicm 9: 287-92.

Hooker W (1861) Dendrobium hillii. Curtis’s Botanical Magazine 87: t. 5261.

Huxley R (2003) Challenging the dogma: classifying and describing the natural world. Pp. 70-79
in Sloan K (ed.) Enlightenment. (The British Museum Press: London)

Jones DL (1988) Native Orchids of Australia. (Reed Books: Frenchs Forest)

Lavarack PS (1991) An appraisal of the species concept in the taxonomy of Australian orchids.
Orchadian 10: 141-43.

Lavarack PS & Gray B (1985) Tropical Orchids of Australia. (Thomas Nelson: Melbourne)
Masters MT (1877) Dendrobium speciosum var. hillii. Gardener’s Chronicle (new ser.) 7: 112, f.
18.

Milligan GW & Cooper MC (1988) A study of standardisation of variables in cluster analysis.

Journal of Classification 5:181-204.

Pearson I I (2001) Biology’s name game. Nature 411: 631-632.

Prentice HC (1986) Continuous variation and classification. Pp. 21-32 in Styles BT (ed.) Infra¬
specific Classification of Wild and Cultivated Plants. (Oxford University Press: New York)
Schlechler R (1982) The Orchidaceae of German New Guinea. (The Australian Orchid Founda¬
tion: Melbourne)

Smith JE (1804) Exotic Botany vol. 1: 17, t.10 (R. Taylor & Co.: London)

Sneath PHA & Sokal RH (1973) Numerical Taxonomy: The Principles and Practice of Numerical
Classification. (WH Freeman and Co.: San Francisco)

Stuessy TF (1990) Plant Taxonomy. (Columbia University Press: New York)

Upton WT (1989) Dendrobium Orchids of Australia. (Houghton Mifflin: Australia)

Walsh G (2000) A bit about the rock orchids ( Dendrobium speciosum complex) of Capricornia.
Orchadian 13: 210-215.

Williams WT (1975a) Hierarchical agglomerative strategies. Pp. 84-90 in Williams WT (ed.)
Pattern Analysis in Agricultural Science. (CSIRO & Elsevier Scientific Publishing Company:
Amsterdam)

Williams WT (1975b) Other Ordination Procedures. Pp. 59-75 in Williams WT (ed.) Pattern
Analysis in Agricultural Science. (CSIRO 8c Elsevier Scientific Publishing Company: Amster¬
dam)

Manuscript received October 2005, accepted 17 February 2006

Telopea 11(2) 233-244

New combinations in the genus Vachellia
(Fabaceae: Mimosoideae) from Australia

Phillip G. Kodela and Peter G. Wilson

National Herbarium of New South Wales, Botanic Gardens Trust, Mrs Macquaries Road,

Sydney NSW 2000, Australia

Abstract

Based on the evidence that the genus Acacia sens. lat. is polyphyletic and the acceptance that
the major groups warrant generic status, and because the name Acacia has been conserved
with a new type, the following new combinations in Vachellia Wight & Arn. are necessary for
Australian taxa formerly included in Acacia subg. Acacia: Vachellia bidwillii (Benth.) Kodela,
Vi clarksoniana (Pedley) Kodela, Vi ditricha (Pedley) Kodela, Vi dotiglasica (Pedley) Kodela,
Vi pachypldoia (W.Fitzg.) Kodela and Vi pachyphloia subsp. brevipinnula (Tindale & Kodela)
Kodela, Vi pallidifolia (Tindale) Kodela, Vi suberosa (A.Cunn. ex Benth.) Kodela, Vi sutherlandii
(F.Muell.) Kodela, and Vi valida (Tindale & Kodela) Kodela. Vachellia farnesiana (L.) Wight &
Am., a pre-European introduction to Australia, has previously been transferred to the genus.
Acacia nilotica (I..) Delile has been left for the attention of other researchers more familiar with
this taxon, its synonymy and its various subspecies.

Introduction

Synopses of recent morphological and molecular studies showing that the genus Acacia
sens. lat. is polyphyletic and requires division into at least five genera are provided by
Maslin et al. (2003,2003a). At the July 2005 International Botanical Congress in Vienna
a conservation proposal by Orchard and Maslin (2003) was endorsed, resulting in the
retypification of the genus Acacia (McNeill et al. 2005). This decision results in the type
of the genus being changed from the African species Acacia nilotica (L.) Delile to the
Australian species Acacia penninervis Sieber ex DC. As a result of this, if the genus is
split, Vachellia Wight & Arn. is the earliest available name applicable to species formerly
included in Acacia subg. Acacia, while the name Acacia is retained for species of the
former Acacia subg. Phyllodineae , the majority of which occur in Australia.

In Australia there are eleven species of the former Acacia subg. Acacia (Pedley 2002);
nine of these are endemic to tropical Australia. Acacia farnesiana (L.) Willd., which is
widespread through the tropics in central America, Africa and Asia, probably arrived
in Australia prior to European settlement (Kodela & Tindale 2001, Pedley 2002). The
more recent introduction, Acacia nilotica subsp. indica (Benth.) Brenan (Fig. 1), which

© 2006 Royal Botanic Gardens and Domain Trust

1SSN0312-9764

234

Telopea 11(2): 2006

Kodela and Wilson

is naturalised in Queensland and the Northern Territory, is now known to show genetic
diversity in Australia indicating that more than one infraspecific taxon may be present
here (Wardill et al. 2006).

Acacia farnesiana is the type of the genus Vachellia, so the combination
Vachellia farnesiana (L.) Wight & Arn. {Prodr. FI. Ind. Orient. 272, 1834) already exists.
Plants of this taxon in Australia seem all to be referable to var . farnesiana (Pedley 2002,
Kodela pers. obs., Seigler pers. comm.) (Fig. 2). Combinations in Vachellia for New
World taxa of the former Acacia subg. Acacia have recently been made by Seigler and
Ebinger (2006).

New combinations in Vachellia are made here for the endemic Australian taxa so that
these names are available for use in floras and checklists. No combination, however, is
made for Acacia nilotica (L.) Delile, which presents more difficulties (see Wardill et al.
2006) and has been left for the attention of other researchers more familiar with this
taxon, its synonymy and its various subspecies.

New Combinations

1. Vachellia bidwillii (Benth.) Kodela, comb. nov. (Fig. 3)

Basionym: Acacia bidwillii Benth., Linnaea 26: 629 ( 1855).

Holotype: Queensland: ‘Ad Wide-Bay, Australiae boreali-orientalis (Bidwill)’ (K).

A. leptoclada var. (?) polyphylla Benth., FI. Austral. 2: 416 ( 1864).

Holotype: ‘East Coast’ [Queensland], R. Brown [Bennett sheet No. 4331], 1802-5
(K; photo NSW).

A. bidwillii var. bidwillii , FI. Austral. 2: 420 (1864), autonym.

A. bidwillii var. polytricha Domin, Biblioth. Bot. 89: 273 (1926).

Type citation: ‘Nord-Queensland: Savannenwalder bei Mungana (Domin II. 1910)’
Holotype: Queensland: in xerodrymio apud opp. Mungana, K. Domin ‘5148’, Feb 1910
(PR 527948).

2. Vachellia clarksoniana (Pedley) Kodela, comb. nov.

Basionym: Acacia clarksoniana Pedley , Austrobaileya 6: 184, fig. 1 A, B (2002).

Holotype: Queensland: 3.9 km from ‘New Dixie’ homestead, J.R. Clarkson 3585, 15 Oct
1980 (BRI). Isotypes: ILL, K, MEL, MO, NSW, PERTH, PR, QRS.

3. Vachellia ditricha (Pedley) Kodela, comb. nov. (Fig. 4)

Basionym: Acacia ditricha Pedley, Austrobaileya 1: 307 (1980).

Holotype: Queensland: Kowanyama Aboriginal Reserve on the Mitchell River,

B. Alpher, Dec 1977 (BRI).

Vachellia (Fabaceae: Mimosoideae) from Australia

Telopea 11(2): 2006

235

Fig. 1 . ‘Acacia’ nilotica subsp. Hndica. a, branchlet with leaves and inflorescences; b, leaf base and
stipular spines; c, pinnule; d, petiolar gland; e, gland at apex of leaf rachis; f, bark; g, inflorescence
in bud; h, flower bud; i, flower; j, gynoecium; k, bracteoles; 1, pod; m, seeds, a-e, i-k, Newman
565 (NSW); f, Coveny 6925 & Hind (NSW); g, h, 1, m, C.E.F. Allen (NSW271068). Scale bar = 0.6
cm (d, e, h-k), 1 cm (c), 1.2 cm (b, g), 3 cm (m), 6 cm (a, f, 1).

236

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Kodela and Wilson

4. Vachellia douglasica (Pedley) Kodela, comb. nov.

Basionym: Acacia douglasica Pedley , Austrobaileya 6: 181, fig. 1 C, D (2002).

Holotype: Northern Territory: 0.5 km W of Douglas River, M. Parker 508, 23 Oct 1974
(BR1). Isotypes: CANB, DNA, NSW.

Acacia ‘Douglas R.’, C.R. Dunlop et al ., FI. Darwin Region 2: 20, fig. 4 (1995).

Note: Pedley (2002) suggested that Tindale and Kodela (1996) included A. douglasica as
part of A. ditricha but this was not the case. Although specimens of the then undescribed
species A. douglasica were known at the time and were considered as a possible variant
of A. ditricha they were neither included in the description of A. ditricha nor cited in
the Tindale and Kodela (1996) paper. The Douglas River taxon was later addressed in
the Flora of Australia (Kodela & Tindale 2001: 202) under Acacia bidwillii.

5. Vachellia pachyphloia (W.Fitzg.) Kodela, comb. nov.

Basionym: Acacia pachyphloia W.Fitzg. in ).H. Maiden, /. & Proc. Roy. Soc. New South
Wales 51: 116 (1917).

Type citation: ‘Slopes of Bold Bluff; hills near C. 92, in proximity to the Synnott Range;
hills by the Charnley and Calder Rivers (W.V.F.)’

Lectotype (fide Tindale & Kodela 1996: 311): Western Australia: near Synnott [Synnot]
Range, W.V. Fitzgerald 1267, July 1905 (NSW). Isolectotype: PERTH.

Residual syntypes [paralectotypes]: slopes of Bold Bluff, W.V. Fitzgerald-, hills by the
Charnley and Calder Rivers, W.V.Fitzgeraid (n.v., location unknown).

5a. Vachellia pachyphloia (W.Fitzg.) Kodela subsp. pachyphloia

5b. Vachellia pachyphloia subsp. brevipinmda (Tindale & Kodela) Kodela, comb,
nov.

Basionym: Acacia pachyphloia subsp. brevipinmda Tindale & Kodela, Austral. Syst. Bot.
9: 314, fig. 3 (1996).

Holotype: Western Australia: 26 km N of turnoff to Mitchell Falls along road to Port
Warrender, M. McDonald 336, 14 June 1987 (NSW). Isotype: PERTH.

6. Vachellia pallidifolia (Tindale) Kodela, comb. nov. (Fig. 5)

Basionym: Acacia pallidifolia Tindale, Telopea 1: 82 (1975), based on Acacia pallida
F. Muell., /. Linn. Soc., Bot. 3: 147 (1859), partial, non Humb. et Bonpl. ex Willd.
(1806).

Lectotype (fide Tindale, loc. cit .): Northern Territory: Fitzmaurice River, ‘arbuscula in
campis apricis’, F. Mueller 76, 75, Oct 1855 (MEL). Isolectotype: NSW.

Residual syntypes [paralectotypes]: McAdam Range, F. Mueller 75, Oct 1855 (K,
PERTH).

Acacia turbata Pedley , Austrobaileya 6: 180 (2002).

Holotype: Northern Territory: E of Mary River, S.T. Blake 17095, 29 Sep 1946 (BRI).
Isotype: NSW.

Vachellia (Fabaceae: Mimosoideae) from Australia

Telopea 11(2): 2006

237

Fig. 2. Vachellia farnesiana var . farnesiana. a, branchlet with leaves, inflorescences and immature
pods; b, leaves, inflorescences and stipular spines; c, stipular spines; d, petiole showing gland;
e, petiolar gland; f, inflorescence in bud; g, involucel of united bracts at apex of peduncle
(usually hidden by flowers); h, flower bud; i, flower; j, gynoecium; k, anther; 1, bracteoles; m,
n, pods; o, seeds, a, f, h-j, 1, Coveny 6572 & Powell (NSW); b, d, Milvain (NSW108509); c, m,
K. Moore (NSW108510); e, G.M. Cunningham 704 (NSW); g, Milthorpe 3821 (NSW); k, Rodd
s.n. (NSW70825), n, o, Coveny 6522 & Powell (NSW). Scale bar = 0.075 cm (k), 0.4 cm (e), 0.6
cm (h-j, 1), 1.2 cm (d, f, g), 2 cm (c), 3 cm (o), 6 cm (a, b, m, n).

238 Telopea 11(2): 2006

Kodela and Wilson

Fig. 3. Vachellici bidwillii. a, branchlet with leaves; b, petiole and stipular spines; c, petiolar gland;
d, apex of leaf rachis; e, inflorescences; f, inflorescence in bud; g, flower bud; h, bracteoles;
i, flower; j, gynoecium; k, anther; 1, pod; m, seed, a—1, Tindale (NSW52690); m, Coveny 6876
&Hind (NSW). Scale bar = 0.1 cm (k), 0.4 cm (c), 0.6 cm (g— j ), 1 cm (b, d), 1.2 cm (f),
4 cm (m), 6 cm (a, e, 1).

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239

Fig. 4. Vachellia ditriclm. a, apex of branchlet with leaf and pod; b, base of leaf showing petiolar
gland; c, apex of leaf; d, apex of pinna; e, bark; f, pod; g, section of pod showing indumentum on
outer surface; h, seed in pod; i, j, seeds; k, seedling, a, Maconochie 206 (DNA); b-d, j, Perry 2833
& Lazarides (CANB); e-i, Jacobs 1732 (NSW); k, AZRI (NT sheet), seedling grown from seed
of N. McK. Henry 7 (NT31554). Scale bar = 0.5 cm (g), 0.6 cm (d), 1.2 cm (b, c), 3 cm (i), 4 cm
(a, j), 6 cm (e, f, h,k).

240

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Kodela and Wilson

Note: The possible type material of Acacia pallida F.Muell., nom. illeg., includes two
different taxa. However, there have been two different interpretations of this material
that have resulted in the replacement name A. pallidifolia Tindale being applied to
each of these two different taxa. We have followed the concept adopted by Tindale
(1975) while Pedley (2002) applies this name to the taxon named A. valida by Tindale
and Kodela (1996). To further add to the confusion, Pedley (2002) reverses Tindalc’s
application of the name Acacia pallidifolia, renaming her species concept as A. turbata
Pedley. The species concept originally adopted by Tindale (1975), and which was in
use for almost 30 years, has now been challenged by Pedley’s (2002) rejection of her
lectotypiiication. While Pedley’s view has merit, we suggest that nomenclatural stability
is not well served by his approach because it would lead to a confusing reversal of
names. A case for conservation of the name A. pallidifolia in the sense of Tindale
(1975) is in preparation and will be submitted for consideration by the Committee for
Spermatophyta.

7. Vachellia suberosa (A.Cunn. ex Benth.) Kodela, comb. nov. (Fig. 6)

Basionym: Acacia suberosa A.Cunn. ex Benth., London /. Bot. 1: 499 (1842).

Lectotype ( fide Kodela, FI. Australia 11 A: 642, 2001): Western Australia: Careening
Bay, North West Coast, 3rd Voyage of‘Mermaid’, A. Cunningham 300, Sep 1820 [larger,
right-hand element] (K). Isolectotype: BM.

Residual syntypes [paralectotypes]: Western Australia: Encounter Cove, Vansittart Bay,
North West Coast, 2nd Voyage of‘Mermaid’, A. Cunningham 488, Oct 1819 (BM, K).

8. Vachellia sutherlandii (F.Muell.) Kodela, comb. nov. (Fig. 7)

Basionym: Albizia sutherlandi F.Muell., Fragm. 6: 22 (1867). Acacia sutherlandii
(F.Muell.) F.Muell. (as ‘Sutherlandi’), Iconogr. Austral. Acacia Dec. 12: [pi. 8] (June
1888).

Holotype: Queensland: Flinders River,/. Sutherland 114 (MEL).

Acacia tnelaleucoides F.M.Bailey, Proc. Roy. Soc. Queensland 5: 121 (Sep 1888).
Holotype: Queensland: South-eastern slope of Newcastle Range, between Georgetown
and Junction Ck, R.C. Burton (?BRI, «.v.)

9. Vachellia valida (Tindale & Kodela) Kodela, comb. nov.

Basionym: Acacia valida Tindale St Kodela, Austral. Syst. Bot. 9: 307, fig. 1 (1996).

Holotype: Northern Territory: Mathison Ck, c. 91.5 km W of Katherine on the Victoria
Hwy, M.D. Tindale 10101, P. Munns & R. Turley, 2 Aug 1989 (NSW). Isotypes: AD, BRI,
CANB, DNA, K, MEL, MO, PERTH.

A. pallida F.Muell., /. Linn. Soc., Bot. 3: 147 (1859), partim, non Humb. et Bonpl. ex
Willd. (1806).

A. bidwillii Benth. var. (?) major Benth. (as‘Bidwilli’),F/. Austral. 2: 421 (1864).
Holotype: (Dry) ridges of the Victoria River, F. Mueller 76, Nov 1855 (MEL).
Isotype: K.

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241

Fig. 5. Vachellia pallidifolia. a, branchlet with leaves and inflorescences; b, pinnules at apex of
pinna; c, leaf scar showing small stipular spines; d, leaf glands; e, inflorescence in bud; f, flower
bud; g, flower; h, gynoecium; i, bracteoles; j, pod; k, seed, a, b, d-i, M. White MR13(CANB);
c, j, k, Blake 17095 (NSW). Scale bar = 0.4 cm (f), 0.6 cm (g-i), 0.66 cm (d), 1.33 cm (e),
2 cm (b, c),4cm (k),6cm (a, j)

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Fig. 6. Vaclwllia suberosa. a, branchlet with leaves and inflorescences; b, leaf and stipular spines;

c, gland at apex of leaf rachis; d, bark; e, inflorescence in bud; f, inner surface of bracteole; g,
outer surface of bracteole; h, flower; i, gynoecium; j, anther; k, pod; 1, section of pod showing
indumentum on outer surface; m, seed, a, c, e, j, Lazarides 7744 (NSW); b, Lazarides 3104 (NSW);

d, Macotiochie 1183 (NSW); f-i Byrnes2228 (NSW); k-m, Lazarides 6515 (NSW). Scale bar = 0.1
cm (j), 0.3 cm (1), 0.4 cm (c), 0.6 cm (f-i), 1.5 cm (e), 2 cm (b), 4 cm (m), 6 cm (a, d, k).

Vachellia (Fabaceae: Mimosoideae) from Australia

Telopea 11(2): 2006

243

Fig. 7. Vachellia sutherlandii. a, part of branchlet with leaf and inflorescence; b, part of leaf
rachis showing petiolar gland; c, apex of pinna; d, bark; e, llower bud; f, flower; g, gynoecium;
h, bracteoles; i, anther; j, pod; k, seed, a-c, e-i, Waterhouse s.n. (NSW108060); d, Jacobs 1374
(NSW); j, k, Hind 1069 & Ingram (NSW). Scale bar = 0.1 cm (i), 0.6 cm (c, e-h), 1 cm (b), 4 cm
(k), 6 cm (a, d, j).

244

Telopea 11(2): 2006

Kodela and Wilson

Note: Pedley (2002) treated A. valida as a synonym of Acacia pallidifolia. Correct
application of the latter name is contingent upon resolution of the issue of its
typifkation, as discussed above.

Acknowledgments

We are grateful for comments and advice on the Acacia pallida typifkation issue provided
in 2000 by Drs Mary Tindale and Jim Ross (MEL), as well as earlier advice provided to
Dr Tindale by the late Dr L.A.S. Johnson. Many thanks also to Bruce Maslin and Paul
Wilson for constructive comments on this manuscript. Dave Seigler kindly provided
details of his recently published paper and other information. Access to collections
was generously provided by BM, K and MEL. Illustrations were by Robin Griffiths and
made available by Dr Tindale. Catherine Wardrop and Julia Sideris assisted with the
final preparation of the illustrations and scale bars.

References

Kodela PG & Tindale MD (2001) Acacia subg. Acacia. Flora of Australia 11A: 196-207.

Maslin BR, Miller JT & Seigler DS (2003) Overview of the generic status of Acacia (Leguminosae:
Mimosoideae). Australian Systematic Botany 16: 1-18.

Maslin BR, Orchard AE & West JG (2003a) Nomenclatural and classification history of Acacia
(Leguminosae: Mimosoideae), and the implications on generic subdivision. Paper on
Worldwidewattle website at http://www.worldwidewattle.com/infogallery/taxonomy/
nomen-class.pdf

McNeill), Stuessy TF, Turland N) & Horandl E (2005) XVII International Botanical Congress:
preliminary mail vote and report of Congress action on nomenclature proposals. Taxon 54
(4): 1057-1064.

Orchard AE & Maslin BR (2003) Proposal to conserve the name Acacia (Leguminosae:
Mimosoideae) with a conserved type. Taxon 52: 362-363.

Pedley L (2002) A conspectus of Acacia subg. Acacia in Australia. Austrobailcya 6: 177-186.

Seigler DS&EbingerJE (2006) New combinations in the genus Vachdlia (Fabaceae: Mimosoideae)
from the New World. Phytologia 87 (3): 139-178. [dated Dec 2005, published 5 Jan 2006]

Tindale MD 8c Kodela PG (1996) Acacia valida (Fabaceae, Mimosoideae), a new species from
Western Australia and the Northern Territory, as well as the typifkation and revision of A.
pachyphloia. Australian Systematic Botany 9: 307-317.

Wardill TJ, Graham GC, Zalucki M, Palmer WA, Playford I & Scott KD (2006) The importance
of species identity in the biological control process: identifying the subspecies of Acacia
nilotica (Leguminosae: Mimosoideae) by genetic distance and the implications of biological
control. Journal of Biogeography 32:2145-2159.

Manuscript received 20 February 2006, accepted 1 March 2006

Telopea 11(2): 2006

245

Corrigenda — Telopea 11(1)

J.C. Millott & K.L. McDougall. A new species of Pomaderris
(Rhamnaceae) from the Central Tablelands of New South Wales.

Page 86: should read as follows.

Pomaderris graniticola (N.G.Walsh & F.Coates) K.L.McDougall & J.C.Millot, comb.
& stat. nov.

Pomaderris argyrophylla N.A. Wakef. subsp. graniticola N.G. Walsh & F. Coates,
Muelleria 10: 46 (1997).

Type: Queensland, Girraween National Park, c. 50 m W of Dr Roberts Waterhole,
towards its southern reach, N.G. Walsh 3883, 15.ix.1994 (holotype MEL; isotypes BRI,
CANB, NSW, UNE).

.

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Telopea

Journal of Plant Systematics

11 ( 2 ): 99-245 • 2006

ISSN 0312-9764

Viola silicestris, a new species in Viola section Erpetion from Australia

Kevin R. Thiele and Suzanne M. Prober 99
Philotheca papillata (Rutaceae), a new endangered species from
north-eastern New South Wales Ian R.H. Telford and Lachlan M. Copeland 105

Pimelea cremnophila (Thymelaeaceae), a new species from the New
England Tablelands escarpment of northern New South Wales

Lachlan M. Copeland and Ian R.H. Telford 111
Rediscovery of Prostanthera staurophylla F.Muell. and reinstatement
of P. teretifolia Maiden & Betche (Lamiaceae)

John T. Hunter, John B. Williams and Barry J. Conn 117
New combinations in Australasian Zostera (Zosteraceae)

Surrey W.L. Jacobs Donald LI. Lesand Michael L. Moody 127
Two new species of Aponogeton (Aponogetonaceae), and a key to species
from Australia Surrey W.L. Jacobs, Donald H. Les, Michael L. Moody

and C. Barre Hellciuist 129

Teucrium thieleanum (Labiatae), a new species from Victoria, Australia

Barry J. Conn 135

Haloragodendron gibsonii (Haloragaceae), a new species from the
Blue Mountains, New South Wales Peter G. Wilson and Michael L. Moody 141
Lysimachia (Myrsinaceae) in New South Wales P.G. Kodela 147

Three new species of Nymphaea (Nymphaeaceae) in Australia

Surrey W.L. Jacobs and C. Barre Hellquist 155
An anatomo-ecological experiment in Austrostipa aristiglumis , a lowland
Stipoid species. Mirta O. Arriaga and Surrey W.L. Jacobs 161

New taxa and combinations for Oedogonium and Bulbochaete
(Oedogoniales, Chlorophyceae) in Australia

Stephen Skinner and Timothy J. Entwisle 171
A review of the taxonomy and relationships of the Dendrobium speciosum
complex (Orchidaceae), and recognition of two new taxa

Peter B. Adams, Jacinta M. Burke and Sheryl D. Lawson 195
New combinations in the genus Vachellia (Fabaceae: Mimosoideae)
from Australia Phillip G. Kodela and Peter G. Wilson 233

Corrigenda — Telopea 11(1) 245

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