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TELOPEA

Volume 8(1) • December 1998

Royal Botanic Gardens 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 specialising in the
flora of New South Wales.

Telopea is published twice yearly.

Annual subscription for 1998-9

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Cover illustration

The Waratah, Telopea speciosissima (Sm.) R. Br., belongs to the family Proteaceae. The
species is endemic in eastern New South Wales and is the official State floral emblem.
Illustration by David Mackay

Telopea

A journal of plant systematics

CONTENTS

Dilhvynia glaucula (Fabaceae: Mirbelieae), a new species from the Southern
Tablelands, New South Wales Peter C. Jobson and Peter H. Weston 1

Aponogetonaceae of Australia, with descriptions of six new taxa

C. B. Hellquist and S.W. L. Jacobs 7

New combinations arising from a new classification of non-African

Restionaceae Barbara G. Briggs and L.A.S. Johnson 21

Two new rare species of Homoranthus (Myrtaceae: Chamelaucieae) from

the Northern Tablelands of New South Wales John T. Hunter 35

Nassella tenuissima (Gramineae) recorded from Australia, a potential new
weed related to Serrated Tussock

S.W.L. Jacobs, J. Everett and Maria Amelia Torres 41

Notes on Australian Meliaceae D.J. Mabberley 47

Revision of Livistona (Arecaceae) in Australia A.N. Rodd 49

SHORT COMMUNICATIONS

Amphipogon (Gramineae) does have microhairs

L. Watson, N.P. Barker and S.W.L. Jacobs 155

Eucalyptus canobolensis (Myrtaceae), a new combination for a former

subspecies of Eucalyptus rubida John T. Hunter 157

TELOPEA 8(1): 1-158 DECEMBER 1998

Scientific Editor
Peter Wilson

Typesetting and Production Assistance
Debby McGerty

Other members of Editorial Committee
Gwen Harden, Joy Everett, Karen Wilson

ISSN 0312-9764

Telopea 7(4) was distributed on 29 May 1998.

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

Dillwynia glaucula (Fabaceae: Mirbelieae),
a new species from the Southern Tablelands,
New South Wales

Peter C. Jobson and Peter H. Weston

Abstract

Jobson, Peter C. l ' 2 and Weston, Peter H. 1 PNational Herbarium ofNeiv South Wales, Royal Botanic Gardens,
Sydney, NSW 2000, Australia, 2 Dept. of Environmental Sciences, University of Technology, Sydney, Gore
Hill, NSW 2065, Australia) 1998. Dillwynia glaucula (Fabaceae: Mirbelieae), a new species from the
Southern Tablelands, New South Wales. Telopea 8(1): 1-5. Dillwynia glaucula Jobson & P.H. Weston,
a new species from the Southern Tablelands of New South Wales is described and its affinity with
Dillwynia sericea is discussed, along with its ecology, distribution and conservation status.

Introduction

In 1996, a specimen of an unknown Dillwynia was submitted to the Herbarium for
identification and was identified as a glabrous form of Dillwynia sericea A.Cunn. While
conducting fieldwork in October 1997, as part of the revision of Dillwynia for Flora of
Australia, we examined this plant in situ, discovering a localised but dense population
interspersed with D. sericea. This population proved to be distinct from D. sericea in a
number of morphological characters, in habit and in its preferred habitat. We describe
this new species below, along with its ecology, and assess its conservation status.

Taxonomy

Dillwynia glaucula Jobson & P.H. Weston, sp. nov.

D. sericeae A. Cunn. affinis sed usque ad 2.3 m alta crescens; rami ramulique omnino
glabri; folia glaucescentia, omnino glabra; inflorescentia axillaris; calyx omnino glaber;
fructus pubescentes, rubiginosi.

Holotype: New South Wales: Southern Tablelands: N side of Claypit Rd, 1.6 km SW
from Nerriga Road junction (Junction is 8 km S of Windellama), 35°05'43"S
149°52'29"E, P.C. Jobson 5103 & P.H. Weston, 20 Oct 1997 (NSW). Isotypes: CANB, K,
MEL, MO.

Erect shrub (0.2-)0.75-2.3 m high; taproot present and lacking lignotuber; bark
smooth, red-brown with conspicuous cream lenticels. Branches and branchlets
smooth, red-brown and entirely glabrous. Leaves spreading to reflexed, rarely loosely
appressed, linear, triquetrous, occasionally slightly twisted, glabrous, minutely
colliculate, young leaves glaucous, becoming bluish-green to green with age; petiole
c. 0.5 mm long; lamina with a longitudinal adaxial groove, 4-7 mm long, c.0.5 mm
wide, apex obtuse, often with an incurved tip; stipules absent. Inflorescences axillary,
often in upper axils of stem, 1-flowered. Peduncles 2.0-2.5 mm long; bracts ovate to
broad-ovate, 0.5-1.0 mm long, entire, cucullate, red-brown, ciliate with white hairs in
upper portion of margin; bracteoles ovate, c. 1 mm long, entire, yellow to brown.

Telopea 8(1): 1998

ciliate with white, straight to crisped hairs on upper portion of margin, attached to
pedicel 0.25-0.75 mm below calyx tube. Buds green with cucullate upper calyx lobes.
Calyx 10-ribbed, green, 3.5-6.0 mm long; calyx tube turbinate; lobes shorter than tube,
lower lobes broadly acute, upper lobes v-shaped notched, divergent, margins ciliate
with white hairs. Standard with lamina reniform with a deep v-notch separating lobes
and basal oblong claw; lamina 4.S-5.5 mm long, 9.0-11.0 mm wide; lobes obovate,
yellow with narrow red band (crescent) above claw; claw yellow-green, 4.0-5.0 mm
long, 2.0 mm wide. Wings narrow-obovate, partially obscuring keel, cucullate, obtuse,
auriculate on lower margin near base and clawed, yellow, 6.0-7.0 mm long, 1.5-2.0 mm
wide; claw 1.0-1.5 mm long, c. 0.75 mm wide. Keel longitudinally broad-ovate,
cymbicate, acute, red, 4.0-5.0 mm long, 2.0-3.0 mm wide; upper margin papillate; claw
c.l mm long. Stamens with filaments 2.0-4.0 mm long; anthers 0.5-0.75 mm long.
Gynoecium c. 4 mm long; ovary w'hite-pubescent, c.1.5 mm long; stipe glabrous c. 0.5 mm
long; style hooked, glabrous, c. 2 mm long; stigma capitate. Pod ovoid, turgid, red-
brown, pubescent with white hairs chiefly towards apex, 4.0-5.5 mm long, 3.0-4.0 mm
wide; petals caducous during late development of pod. Seeds ovoid, smooth, black to
dark olive with dark brown spots; aril cream, c. 2 mm long. (Fig. 1).

Phenology: flowers mainly in October but sporadic to January; fruits have been
collected from November to January.

Distribution: known from three areas, near Windellama, Michelago and Numeralla,
New South Wales. (Fig. 2).

Habitat: the Windellama populations of D. glaucula occur in dry sclerophyll
woodlands dominated by Eucalyptus rossii, E. macrorhyncha, E. pauciflora and
E. gregsoniana, with either a grassy or shrubby understorey on white kaolin-type clays.
Most of the southern (Michelago and Numeralla) populations occur in woodlands
dominated by £. rossii, E. nortonii and E. dives with a shrubby understorey. At one site,
the remnant roadside vegetation is open and shrubby, with Callitris endlicheri, Cassinia
aculeata, C. sp. aff. quinquefaria, Bossiaea riparia, Chrysocephalum sp., Wahlenbergia sp.,
Sanguisorba minor and various grasses. All southern populations occur on either rocky
sandstone hill tops or slaty outcrops. Tire altitudinal range of this species is between
580 m and 840 m.

Conservation status: prior to 1997, this species had only been collected three times. It
should be considered vulnerable (3V coding of Briggs & Leigh, 1996) as it does not
occur in any known reserve. Most populations are known from roadside reserves and
are potentially threatened by the widening of these roads. At one locality (Jobson 5384),
the remaining plant in the area had been heavily grazed, as had most of the
understorey vegetation. Although there has been clay mining in the Windellama area
in the past, the pit on Claypit Road appears to be disused.

Etymology: the epithet glaucula is from the Latin glaucus meaning 'bluish-green' and
-ula which is a diminutive suffix and refers to the blue-green colour of the leaves,
especially observed in the young leaves of the plant, and which tends to fade with age.

Notes: Dillwyniaglaucula most closely resembles D. sericea. In the key of Weston (1991),
D. glaucula keys out unsatisfactorily to either D. sp. C. or D. sericea. Dillwynia sp. C is
confined to the sandstone escarpments and cliff tops of the Budawang Ranges and
differs from D. glaucula in having larger, more robust leaves and larger flowers.
Dillwynia glaucula can be distinguished from D. sericea by its completely glabrous
calyx, leaves and new shoots (silky hairy in D. sericea) and by the blue-green tinge of
the leaves (dull green in D. sericea). In the Windellama area, D. sericea is sympatric but
is much smaller in stature (up to 1 m tall) than D. glaucula and has far less branching.
Dillwynia glaucula occurs there on a kaolin clay lens (often only a few metres in area)

Jobson and Weston, Dillwynia glaucula (Fabaceae) from the Southern Tablelands

Fig. 1. Dillwynia glaucula. a, habit; b, front view of flower; c, side view of flower; d, pod (all from
Jobson 5103 & Weston). Scale bars: a = 5 mm; b, c = 3 mm; d = 2 mm.

Telopea 8(1): 1998

that outcrops on the summits of small hills, with D. sericea occurring nearby on the
slopes of the same hill on soils derived from metasediments.

Specimens examined: New South Wales: Southern Tablelands: Cnr. of Claypit and Nerriga Rds,
8 km S of WindeUama, 35°05'21"S 149°53'05"E, PC. Jobson 5095& P.H. Weston ,20 Oct 1997 (fl.)(NSW,
NSW Ref. Coll., AD, BRI,), P.C. Jobson 5333 & B.M. Wiecek, 21 Nov 1997 (fr.)(NSW, CANB), T. Hayes
s.n., Jan 1996 (fr.) (NSW); E side of Claypit Rd, 2.2 km from Nerriga Rd junction, 35°06'05"S
149°52'39"E, P.C. Jobson 5705 & P.H. Weston, 20 Oct 1997 (fl.)(NSW); N side of Claypit Rd, 2.5 km
from Nerriga Rd junction, 35 o 06T2”S 149°52'36"E, PC. Jobson 5106 & P.H. Weston, 20 Oct 1997
(fl.)(NSW); 3.75 miles [6.0 km] from Michelago on Michelago - Queanbeyan Rd, 35°44'S 149°12'E,
C. W.E. Moore 1657 ,20 May 1952 (fr.) (CANB); 200-300 m E of Michelago - Queanbeyan Rd, 6.3 km
NNE of Michelago store, 35°40T2”S 149°12'09"E, PC. Jobson 5382 & P.H. Weston, 20 Jan 1998 (old
fl) (NSW); c. 2 km E of Michelago of the Canberra - Cooma Road, 35°43'S 149°12'E, R. Pullen 11,050,
3 Jan 1982 (fl. & fr.) (CANB, AD, PERTH); N side of Michelago - Queanbeyan Rd, 1.0 km E of
Michelago store, 35°42'44"S 149°10’06"E, PC. Jobson 5386 & P.H. Weston, 20 Jan 1998 (fr.) (NSW,
MEL), PC. Jobson 5389 & P.H. Weston, 20 Jan 1998 (fl.)(NSW); 4 km WSW by road from Numeralla
towards Cooma, 36°11'00"S 149°18'15"E, D.E. Albrecht 2244 ,15 Oct 1985 (fl.) (MEL).

Acknowledgments

Tim Hayes drew our attention to this species initially and has been most helpful in
subsequent discussions. Peter Wilson checked the Latin diagnosis, Nicola Oram drew
the illustration and Leonie Stanberg assisted in drawing the map. This project is
funded by an Australian Biological Resources Study Research Grant.

Jobson and Weston, Dillwynia glaucula (Fabaceae) from the Southern Tablelands

References

Briggs, J.D. & Leigh, J.H. (1996) Rare and Threatened Australian Plants, 1995 revised edition. (CSIRO:
Collingwood).

Weston, P.H. (1991) Dillwynia. Pp. 499-504 in Harden, G. J. (ed.) Flora of New South Wales, vol. 2.
(New South Wales University Press: Sydney).

Manuscript received 12 June 1998
Manuscript accepted 30 July 1998

Aponogetonaceae of Australia, with
descriptions of six new taxa

C. B. Hellquist and S.W. L. Jacobs

Abstract

Hellquist, C. B9 and Jacobs, S.W.L. 2 ('Department of Biology, Massachusetts College of Liberal Arts, North
Adams, MA 01247-4100, USA; 2 Royal Botanic Gardens, Sydney, NSW, Australia 2000) 1998.
Aponogetonaceae of Australia, with descriptions of six new taxa. Telopea 8(1): 7-19. Ten species of
Aponogeton (Aponogetonaceae) are recognised from Australia, five of which are described here for
the first time: Aponogeton kimberleyensis, A. lancesmithii, A. euryspermus, A. proliferus, and
A. vanbruggenii. Two subspecies of A. elongatus are also recognised: A. elongatus subsp. elongatus
and the newly described A. elongatus subsp. fluitans.

Introduction

While preparing the treatment of the family Aponogetonaceae for the Flora of Australia
it became obvious that neither the material being received from commercial suppliers
of aquarium plants nor the herbarium specimens could be adequately dealt with by
existing treatments. Bruggen (1969, 1973,1985) is the acknowledged authority on the
family and his pioneering studies have been responsible for an increased interest in
species of Aponogeton and a large increase in field observations and the number of
specimens available for study. Aston (1973) is the only Australian author to treat the
genus in any detail and she did not have the opportunity to study the genus in the
field. Bruggen (1969) was clearly aware of some of the problems as can be seen from
his treatment of A. elongatus where his forms are rather mixed; in one case he cites the
same specimen under two different forms and in another cites duplicates under
different forms. His notes clearly state that he was uncertain about this group and it is
indeed from this group that all of the new taxa are here separated. The problems with
the taxa seem to have been exacerbated by the use of floating versus submerged leaves
(Bruggen 1969) as a major key character. This is a good character for some of our
species but specimens from some tropical localities were being identified as one
species at one time of the year, and another species later in the year. The three species
from the wet/dry tropics described here (A. kimberleyensis, A. euryspermus and
A. vanbruggenii) produce submerged leaves in flowing or clear water and, as the water
becomes more turbid from mud or algae, or is shaded, produce more floating leaves.
This character is hard to quantify but, as we examined the specimens, it became
obvious that there were correlated seed characters that were easier to quantify.

Bruggen (1969,1985) was fortunate in being able to rely heavily on cultivated plants.
We had several opportunities to examine plants in the field and the benefit of help
from Lance Smith in cultivating many of the species. It appears that some of the
difficulties that we had with characters listed in Bruggen (1985) relate to the difference
between having fresh cultivated specimens available for study as opposed to relying
on dried and pickled herbarium specimens. In the same publication, Bouman (1985)
makes it clear that some of the embryo characters are not as clear-cut as presenting
them graphically may suggest. At least in the Australian species, germination seems
to be a continuous process. If ripe, shed seed can be collected from all species then

Telopea 8(1): 1998

comparisons of characters like the plumule can be useful. By contrast, the seed coat
characters can be useful over a wider range of maturity and have been important in
helping us delimit taxa.

The characters

Inflorescence: the inflorescence is usually either a solitary spike (most Australian
species) or forked (A. distachyos and A. hexatepalus and rarely in A. lancesmithii). In
A. hexatepalus there may be further branching. The flowers and inflorescences of all
native species are yellow or greenish yellow, the introduced A. distachyos being white or
pinkish.

The axis itself is often pithy and helps the inflorescence float. In some species growing
in flowing water, the tip of the axis is bent upwards, exposing the young flowers to the air.

It is only the introduced A. distachyos that has its flowers arranged fairly closely on one
side of each branch of the forked inflorescence. Tire native species have the flowers
arranged right around the axis. In A. bullosus, where the inflorescences are rarely
exposed, and in A. queenslandicus, which normally grows in still water, the flowers are
usually always tightly packed, hr A. hexatepalus, which also usually grows in still
water, the flowers are well-spaced. In the other native species the density of the
flowers depends on conditions. In still, shallow water with a stable water depth, the
flowers are usually densely packed. If there is a rise in water level, or an increase in
flow, after flowering has started, this is usually countered by an elongation of the axis
rather than by a further extension of the peduncle. In these specimens the flowers are
well-spaced. Consequently these characters are of only limited use in delimiting species.

Leaves: great variability exists within species and populations of the various taxa.
Many of the species, especially A. elongatus, A. euryspermus, and A. vanbruggenii, often
grow in fast-moving waters of creeks and rivers and, in these situations, often have
much longer leaves. The streams and rivers are also often heavily shaded. Plants
growing in deep shade usually have thin leaves that are much wider and thinner than
those growing in the sun. Leaves in the deep shade may tend to be more ovate than
broadly lanceolate. Most of the specimens have distinctive raised glands or punctae
along the leaves and occasionally the petioles. Leaf anatomy and morphology have
been summarised by Tomlinson (1982) and Baas (1985). Distinctive hydropoten, or
localised regions of flat cells that appear as blotches, occur on many plants (Tomlinson
1982). Leaf venation is not distinctive. Two species, A. bullosus and A, lancesmithii, have
bullate, or crinkled leaves, variously developed in the latter species.

Floating leaves are especially characteristic of A. distachyos, A. hexatepalus,
A. queenslandicus and A. elongatus subsp.fluitans. Many of the others will produce them
at certain times. However, A. bullosus, A. lancesmithii, and A. proliferus have not been
observed producing floating leaves, and A. elongatus subsp. elongatus produces them
only rarely.

Submersed leaf margins are either undulate or flat. Aponogeton elongatus, A. bullosus,
and A. lancesmithii have leaf margins that are almost consistently undulate.
Occasionally A. euryspermus and A vanbruggenii produce undulate leaves.

Submersed leaf tips and bases are not really useful characters. Most of the leaf tips are
obtuse, and the bases are cuneate, but many other variations and combinations of leaf
tips occur.

Floating leaf tips and bases are, however, useful in distinguishing A. qileenslandicus
with its characteristic cordate leaf bases.

Hellquist and Jacobs, Aponogetonaceae of Australia

Peduncle: in most species the length of the peduncle is directly dependant on both
water depth and speed of flow, length being increased with increases in both.
The exception is A. bullosus, in which the inflorescence is frequently never exposed
above the water, good fruit set being achieved without the flowers being exposed. In
other Australian species fruit is usually not set if the flowers are not exposed above
water.

The apex of the peduncle immediately below the inflorescence may be expanded or
not. This does not appear to be a reliable character as it seems to depend on the
environment, expanded apices being much more common in flowing water where the
inflorescences are not exposed. Species that commonly lack expanded peduncle apices
include A. bullosus, A. qucenslcmdicus and A. vanbruggenii.

Plumule: Bruggen (1985) states that the plumule may be covered by the cotyledonary
sheath and never be visible or it may be variously developed and visible in different
species. We found this character to be very variable on herbarium specimens and
apparently related to the stage of maturity, the more mature the seed the greater the
probability of finding a plumule or a groove. In all cases we found it preferable to
remove any seed coats before looking for a plumule. Because of the limited value of
this character in herbarium specimens we did not use it to delimit species but we
provide relevant information in the description.

Seeds: seeds provide the best diagnostic characters to distinguish many of the taxa.
There are differences in size, shape and number per carpel. Of the Australian species,
A. euryspermus has the largest seeds (to about 5 mm long, almost ovoid, and usually
only l-3(-5) per carpel), and A. kimberleyensis (1-1.5 mm) and A. elongatus (1-3 mm)
the smallest. A. queenslandicus has a large number (4-13) of almost cylindrical seeds
per carpel. A. vanbruggenii has seeds almost as long as those of A. euryspermus but they
are terete or flattened and about half as wide as the almost ovoid seeds of the latter.

Testa: the single or double seed coat or testa (Bouman 1985) is useful in many cases.
Seeds with the distinct double testa are those of A. queenslandicus, A. lancesmithii,
A. euryspermus, and A. vanbruggenii. A. queenslandicus has the outer layer tightly
adhering to the inner coat, while the other three are quite loose and easily detached.

Aponogeton

Perennial freshwater herbs; monoclinous, monoecious or dioecious, with tuber or
rhizome, stolon or corm and erect stem; entomophilous via beetles or bees, or
cleistogamous; with milky sap and raised glands or punctae. Leaves simple, sheathing
at base, alternate, petiolate or sessile, often containing hydropoten (localised regions
of flat cells appearing as blotches), floating, floating and submersed, or all submersed;
blades with parallel venation, main veins connected by cross veins, margins often
bounded by a thin parallel vein; tissue between veins may disintegrate with age,
leaving a network of veins separated by holes; axillary scales present. Inflorescence
pedunculate, spadix subtended by a caducous or persistent spathe, branched or single,
floating, projected above the water or remaining submerged. Flowers sessile,
zygomorphic, perfect or imperfect; perianth in one whorl of 1-6 parts or absent,
persistent, white to pink, yellow, green-blue or violet; stamens 6-18(-50), 2-loculed,
dehiscing by longitudinal slits; carpels separate, 2—6(—9), superior, sessile, placentation
basal. Fruit a follicle, with terminal, lateral, curved or straight beak. Seeds with
straight embryo, lacking endosperm, with single or double testa.

Telopea 8(1): 1998

Key to species

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

... 1. 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 . 2. 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 .

... 3. A. hexatepalus

4* Tepals 2; seeds 2.1-3.2 mm long; leaves all submersed .

4. 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 . 5. A. bullosus

6* Spathe caducous, to 1.5 cm long; inflorescence to 34 cm long; peduncle much broader
than inflorescence rachis . 4. A. lancesmithii

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

with obvious double testa . 6. A. queenslandicus

7* Plants mainly producing submersed leaves or submersed and floating leaves;
floating blades 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 .

... 7. 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. A. elongatus

9* Fruit elliptic and papery; seeds with a loose outer coat, easily separated from embryo;
floating leaves commonly formed . 10

10. Seeds broadly elliptic, nearly oval, (1.1—)1. 5—2.8 mm wide . 8. A. euryspermus

10* Seeds narrowly elliptic, 0.7-1.6 mm wide. 9. A. vanbruggenii

Hellquist and Jacobs, Aponogetonaceae of Australia

1. A. proliferus C.B. Hellquist & S.W.L. Jacobs, sp. nov.

A. elongato affinis, sed inflorescentiis raris, plerumque planta nova ad apicem pseudo-
pedunculi spathacei substitutis, differt.

Holotype: Cultivated by Lance Smith, Townsville, K.L. Wilson 8747, 11 July 1994
(NSW). Isotype: BRI.

Rooted, submersed, perennial aquatic. Tubers small, less than 1 cm; fibrous roots
abundant. Leaves submersed; blades linear, 18.5-31 cm long, (0.4-)1.3-1.6 cm wide,
dark green; margins flat to slightly undulate; base cuneate; apex obtuse to broadly
mucronulate; 7-veined; lacunae 0 or 1 on each side of the midrib; petiole to 13 cm long.
Peduncle to 32 cm long, broadening at base of inflorescence or proliferous offshoot.
Spathe to 1.5 cm long, partially persistent. Inflorescence yellow, emergent, single,
rarely formed, to c. 5.5 cm long; rachis c. 1.6 mm wide. Flowers widely spaced, mostly
bisexual, few unisexual, staminate; flowers turned in all directions. Perianth segments

2, spathulate, 1-veined, yellow. Stamens (l-)4(-6); anthers c. 0.4 mm wide, 0.4 mm
long, filaments c. 0.5 mm. long, broader at base. Carpels (0-)4—5(-9). Fruit not seen.
New plants commonly produced at tip of spathed peduncle-like structures.

Distribution: an extremely rare species now known only from the Innisfail region of
Northern Queensland. Presumably now restricted because of extensive clearing and
loss of habitat.

Specimens examined: Queensland: Innisfail district, creek through rainforest, S. Jacobs 7148
(NSW); cultivated at Kelso, Townsville by Lance Smith, S. Jacobs 8258,17 Aug 1997 (NSW).

This species is named for the proliferous shoots produced; from the Latin prolifer.

Despite the proliferous shoots this species is difficult to propagate and the proliferous
offshoots need to remain in contact with the parent plant until well established
(L. Smith, pers. comm.). This is the only yellow-flowered taxon of two proliferous
species known worldwide. The other is A. undulatus Roxb., a white-flowered species
from Asia (Bruggen 1985). Only one plant of A. proliferus has been seen in flower and
this was in cultivation. Its natural habitat seems to be narrow, shallow and heavily
shaded coastal streams.

2. A. distachyos C. Linnaeus f., Suppl .: 215 (1781)

Holotype: LINN 479.3 p.p. (Bruggen 1973)

Rooted, submersed and floating, perennial aquatic. Tuber to 6 cm long. Leaves
floating, blades ovate, narrowly elliptic, or narrowly lanceolate, 6-26(-32.5) cm long,
1.5-7.5(-8.6) cm wide; margins flat; base mostly obtuse or attenuate; apex obtuse,
acute, or acuminate, 7-9-veined; petiole to 100 cm long. Peduncle to 80 cm long, often
broadening at base of inflorescence. Spathe c. 3 cm long, caducous. Inflorescence
branched with two spikes to 5.5 cm long. Flowers secund, in 2 rows. Perianth
segments 1, white or pinkish, ±13-veined, 10-15(-30) mm long, 3.5-12 mm wide.
Stamens 8-16,3-4.5 mm long, anthers purplish black, filaments wider at base. Carpels
2-6. Infructescence to 7 cm long. Fruit to 22 mm long, 6 mm wide, with a curved or
straight terminal beak to 5 mm long. Seeds to 18 mm long, 7 mm wide, with a single
loose testa and small embryo.

Distribution: naturalised in southern South Australia, southern New South Wales,
and Victoria. This is the only introduced species of Aponogeton naturalised in
Australia. It is sometimes regarded as a minor weed though it is widely planted as an
ornamental in temperate areas because it is one of the few aquatic species that flowers
in the cooler months.

Telopea 8(1): 1998

Selected specimens examined: South Australia: Mount Lofty Range. Mylor, c. 20 km south-east
of Adelaide, N.N. Dormer 97, 2 July 1961 (MEL). New South Wales: Vineyard Creek, Rydalmere,
33°49'S, 151°02' R.G. Coveny 6142, D. Benson, L. Johnson & K. Wilson, (NSW); Royal Botanic Gardens,
Sydney, S. Jacobs 3345, 1 May 1978 (NSW). Victoria: c. 6 miles [9 km] south-east of Traralgon,
A. Eddy s.n., 29 May 1965 (MEL); Gisborne, in Jacksons Creek, where crossed by the Calder
Highway, H.I. Aston 1144 (MEL); Lome, Stony Creek, at crossing of the Great Ocean Road just
immediately north of the town, H.I. Aston 1585 (MEL); Lilydale district, approx. 35 km from
Melbourne city center, H.T. Clifford s.n., Oct 1948 (MEL); 1 mile [1.3 km] west of Ferntree gully,
Ferny Creek at the crossing of the road to Lysterfield, just as it leaves the main Melboume-Femtree
Gully road ,H.l. Aston 918, 21 Feb 1963 (MEL, NSW); Mornington Peninsula, north of Bewtons Rd,
1 km east of its junction with Moorooduc Rd, Bill Peel 51, 5 Sep 1981 (MEL); Otway Ranges,
c. 18 miles [25 km] south-west of Colac, c.lA miles [3 km] west of Carlisle, in the Gellibrand River,
H.I. Aston 1595 (MEL); south-eastern Otways, c. 9 miles [12 km] direct line north-east of Apollo Bay,
Great Ocean Road, c. A mile [1 km] east of Carisbrook Creek, H.I. Aston 1590,22 March 1967 (MEL);
Allsops Creek, 2 km from old Jeetho railway station; approx. 8 km NW of Kurumburra, /. Scott s.n.,
1 Oct 1980 (MEL); West Macedon, A.]. Tadgcll s.n., 1 May 1932 (MEL).

Grows in slow-flowing freshwater creeks and rivers, lakes, and garden ponds. Flowers
and fruits from September to May. This species grows best in the cooler waters and
during the cooler times of the year, hence is often most productive in spring and
autumn. This is widely planted as an ornamental water garden plant and is commonly
known as the Water Hawthorn, Cape Hawthorn or Cape Pondweed. Native to
southern South Africa.

3. A. hexatepalus H. Bruggen, Bluniea 17: 136 (1969)

Holotype: Western Australia: Trigwell, den Hartog 203, 26 Aug 1967 (L). Isotype: L.

Rooted, submerged and floating, perennial aquatic. Tubers elongate to 6.5 cm long, 2 cm
wide. Leaves floating, narrowly oblong, to 20 cm long, 0.3-0.7 cm wide, margins flat;
base narrowly cuneate; apex obtuse or blunt-cuneate; 5-9-veined, some indistinct;
petiole to 20 cm long. Peduncle to 40 cm long, not broadened or inflated at base of
inflorescence. Spathe 2-3 cm long, persistent. Inflorescence branched forming 2 spikes,
green (yellowish-green), to 17 cm long, laxly-flowered, sometimes further branching
present; rachis white, lower portion bare. Flowers turned in all directions. Perianth
segments 6, transverse broadly obovate, 1-veined, green. 0.5-1 mm long, 0.75-1.25 mm
wide. Stamens 6; anthers 0.3 mm long, 0.3 mm wide; filaments to 1.5 mm long, slightly
broader at base. Carpels 3(^4). Fruit (3.8-)5.0-7.0(-10.0) mm long, 1.9-3.2(-5.0) mm
wide, with a terminal beak. Seeds elliptic, (2.6-)5.1-6.2(-7.0) mm long, (1.2—)1.7-3.0 mm
wide, with a single testa, plumule present.

Distribution: endemic to southwest Western Australia in the vicinity of Perth. Now
rare due to loss of habitat.

Selected specimens examined: Western Australia: Kenwick, south of Perth, 32°0'55"S, 115°58'09"E,
S. Jacobs 6947, 16 Nov 1993 (NSW), G. Sainty 1890, 10 Oct 1993 (NSW); old school site, Waterloo’
NNE of Bunbury, 33°19'51"S, 115°45'27"E, S. Jacobs, 15 Nov 1993 (NSW).

Grows in freshwater ephemeral pools that contain water for 3 to 4 months a year.
Flowering and fruiting from August to November. This is the only species of
Aponogeton that consistently has 6 perianth segments.

4. A. Iancesmithii C.B. Hellquist & S.W.L. Jacobs, sp. nov.

A. elongato A. bullosoque affinis; ab A. elongato foliis plus minusve bullosis differt; ab
A. bulloso spatha inflorescentiae caduco, inflorescentia plerumque emergenti, differt; a
speciebus ambobus testis duabus seminium, testa exteriori longitudinaliter plicata
porcatave differt.

Hellquist and Jacobs, Aponogetonaceae of Australia

Holotype: Queensland: Woopen Creek at crossing 7 km west of Russell River bridge
along Woopen Creek Road, SW of Babinda, C.B. Hellquist 16197, C.E. Hellquist &
L. Smith, 6 Oct 1997 (NSW). Isotypes: B, BRI, GH, MEL, NASC, US, Z.

Rooted, submerged, perennial aquatic. Tuber ovate-elongate, to 5 cm long, 2-3 cm
wide. Leaves submersed; blades linear-lanceolate, 10-48 cm long 0.9-4 cm wide,
slightly to strongly bullate, green to dark green; margins slightly undulate; base
narrowly cuneate; apex obtuse or narrowly blunt-cuneate; 7-9-nerved; 0-2 rows of
lacunae on each side of midvein; petiole to 36 cm long. Peduncle to 74 cm long, 1.2-1.8
mm wide at base, broadening to 4.2-6.8 mm at base of inflorescence. Spathe 0.9-1.5 cm
long, 0.7-0.9 cm wide, caducous. Inflorescence yellow, often floating on surface with
the tip emergent, single, rarely branched, to c. 34 cm long, sparsely to tightly flowered;
rachis 1.5-4.5 mm wide. Flowers turned in all directions. Perianth segments 2,
spathulate, 1-veined, yellow, 1.5-1.7 mm long, 0.8-1.3 mm wide. Stamens (4-)6(-8);
filaments c. 1.5 mm long, broader at base. Carpels (2—)3—5(—7). Infructescence dense.
Fruit 3-3.9 mm long, 2.7-3.5 mm wide, with a somewhat straight or bent terminal
beak. Seeds 3-8 per fruit, 2.1—3.2 mm long, 0.6-1.0 mm wide, with double testa and
loose coat with longitudinal folds.

Distribution: endemic to the Russell River and its tributaries in the Innisfail District
of Far Northern Queensland. Grows in slow-to fast-flowing rivers.

Selected specimens examined: Queensland: Russell River, W.G. Trapnell 109,110 ,3 July 1960 (BRI);
Russell River-Bruce Highway, W. Irvine, s.n., 26 Aug 1986 (NSW); S. Jacobs 7240 14 Nov 1994 (NSW);
S. Jacobs, 8248, B. Hellquist, & J. Wiersema, 14 Aug 1997 (BRI, NASC, NSW); Stone Creek north of
Innisfail, 17°29.01'S, 146°00.97'E, S. Jacobs 7153 (NSW); Woopen Creek, a small tributary of Russell
River 8 miles [12 km] S of Babinda, 17°27'S, 145°55'E, K. Williams 216 ,16 Oct 1968 (BRI); Woopen
Creek, a tributary of the Russell River, approximately 16 km SSE of Babinda, G. Trapnell, s.n., July
1970 (BRI).

Grows in slow- to fast-flowing waters of streams. Flowers and Fruits from at least July
to November.

This species is named for Lance Smith, an aquatic plant propagator and grower from
Kelso (Townsville), Queensland who has been instrumental in propagating and
preserving a number of species of Aponogeton and providing us with critical specimens
and information.

Bruggen (1969) included the limited material of this species that he was able to
examine in A. bullosus.

Unlike A. bullosus, which produces fairly short peduncles and mostly submerged
inflorescences, A. lancesmithii produces very long inflorescences on inflated peduncles
that are longer than any other species in Australia. The inflorescences are usually
emergent. The leaves are often bullate. This character at first led us to believe that it
was a possible hybrid between A. elongatus and A. bullosus. However, the seeds of
A. lancesmithii are more similar to those of A. queenslandicus, A. vanbruggenii, and
A. euryspermus than to those of either A. elongatus and A. bullosus. A. lancesmithii seeds
have a double testa and a loose outer coat but they are generally smaller than the seeds
of A. vanbruggenii and A. macrospermus.

5. A. bullosus H. Bruggen, Blumea 17: (1939)

Holotype: Queensland: Ravenshoe, Millstream River, den Hartog 1049, 15 Nov 1967 (L).
Isotype: L.

Rooted, submerged, perennial aquatic. Tubers elongate to 2 cm long. Leaves
submersed; blades linear to linear-lanceolate, 7-40(-50) cm long, (0.5-)l-2(-2.5) cm
wide, strongly bullate, dark green; margins undulate; base narrowly cuneate; apex

Telopea 8(1): 1998

blunt-cuneate to obtuse, 3-5(-7)-veined; lacunae 0-2 rows on either side of midvein;
petiole 1-10(-18) cm long. Peduncle to 30 cm long, not noticeably thicker at base of
inflorescence. Spathe to 2.5 cm long, mostly persistent. Inflorescence yellow, single to

5.5 cm long, mostly remaining submerged. Flowers turned in all directions. Perianth
segments 2, spathulate or obovate, 1-veined, yellow, 0.75-2.7 mm long, 0.75-1.5 mm
wide. Stamens 6, filaments 1.2-1.6 mm long, slightly broader at base. Carpels (2-)3-5.
Infructescense dense, cone-shaped, usually submersed. Fruit 4.2-6 mm long, 3-4 mm
wide, with a terminal beak. Seeds 3-8 per fruit, 2.2-25 mm long, 1-1.6 mm wide, with
a simple testa. Embryo without plumule.

Distribution: confined to northern Queensland in fast-flowing rivers on and running
off the Atherton Tableland. The fruit consistently develop below the water surface.
Emergent flowers are not commonly seen.

Selected specimens examined: Queensland: Calleja Bridge, South Johnstone R., 17°35.62’S,
145°57.17'E, S. Jacobs 7147, 2 July 1996 (NSW); Cochable Creek, Forestry Reserve 756, B. Hyland
APO/2775 June 1962 (BRI); Five Mile Creek, 17°25'S, 145°40’E, A. W. Dockrill 1176, 20 June 1976 (BRI);
Malanda, North Johnstone River, B. Chirping 9674, 20 Sept 1945 (MEL); North Johnstone R.,
Jaggon-Butchers Creek road, 17°22.38'S, 145°37.98'E, S. Jacobs 8253, B. Hellquist & J. Wiersema,
15 Aug 1997 (BRI, NASC, NSW); tributary of N. Johnstone R., Jaggon-Butchers Creek road,
17°23.56'S, 145°37.31'E, S. Jacobs 8252, B. Hellquist & J. Wiersema, 15 Aug 1997 (BRI, NSW); National
Park 226, Palmerston Section, Henrietta Creek, near camping area, P.D. Boslock 7572, July 1994 (BRI);
Theresa Creek, Millaa Millaa, W.G. Trapnell 251, 17 June 1960 (BRI); The Millstream, Ravenshoe,
17°38’S, 145°29’E, S. Jacobs 4782 & G. Sainly (NSW); Josephine Creek, Bartle Frere to highway road,
17°25.82'S, 145°53.93'E, S. Jacobs 8249, B.Hellquist, J.Wiersema, 14 Aug 1997 (BRI, NASC, NSW).

Grows in cool rapidly flowing freshwater rivers and streams. Flowers and fruits from
June to October.

The flowers sometimes emerge for a very short time above the surface for pollination
and fall below the surface to develop fruit. When the flowers do not emerge they still
produce fruit. The infructescence is typically quite short, and conical with usually all
fruit developed. The distinguishing characteristic of this species is the extremely
bullate leaves. This species retains the spathe longer than any other Australian species
of Aponogeton.

6. A. queenslandicus H. Bruggen, Blumea 17:123-125 (1969)

Holotype: Rockhampton, Thozet s.n. (MEL). Isotype: MEL.

Rooted, submerged and floating, perennial aquatic. Tubers 1-5 cm long, 1.8-2.6 cm
wide, extremely strigose. Leaves submersed and floating; submersed blades
uncommon, elliptic to lanceolate, green, 10—13(—70) cm long, 2.9-4.5 cm wide; margins
flat; base obtuse, attenuate, or truncate; apex obtuse; 7-9-veined; 0-1 row of lacunae
on each side of midvein; petiole to 15 cm long. Floating leaves ovate or elliptic,
2.8-14(-27) cm long, 0.6-4.3 cm wide; light green to green, base mostly cordate, rarely
truncate or cuneate; apex obtuse, mucronulate, acuminate, rarely emarginate; 7-9-
veined; petiole to 66 cm long. Peduncle to 30(—73) cm long; 1.8-2.5 mm wide at base
often only slightly broadening or to 2-4.5 mm wide at base of inflorescence. Spathe to

1.5 cm long, caducous. Inflorescence yellow, emergent or floating, single, usually
tightly-flowered; rachis 15—27(—45) mm wide. Flowers turned in all directions.
Perianth segments 2, spathulate, 1-veined, 1.4—2 mm long, 0.5-1 mm wide. Stamens 6,
up to 3 mm long, slightly broader at base. Carpels (2—)3(—4). Infructescence dense.
Fruit 2.5-4.5 mm long, 2.3-3 mm wide, with a terminal, often curved, beak. Seeds
narrowly elliptic, 4—13 per fruit, 1.7-3 mm long, 0.6-1 mm wide, with a double testa;
outer testa loose, transparent and reticulate, inner testa brown and close fitting;
plumule absent.

Hellquist and Jacobs, Aponogetonaceae of Australia

Distribution: northeast Northern Territory and throughout Queensland south of Cape
York Peninsula, though more common west of the Great Divide and not common in
the driest areas.

Selected specimens examined: Northern Territory: 11 km from Borroloola, Daly Waters Road,
16°06'S, 136°13'E, S. Jacobs 1668, 18 May 1994 (NSW). Queensland: near Bowen, in waterhole in
Eucalyptus forest, S.T. Blake 18562, 29 Sept 1950 (BRI); Gilruth Plains, Cunnamulla, 190 m,
H.S. McKee 10350, 12 April 1963 (BRI); Dorunda Lake overflow, 1 km north of Dorunda homestead,
16°32'S, 141°49'E, V.J. Neldner 2926 & J.R. Clarkson, 12 June 1990 (BRI, MEL, NSW,); Dalrymple Shire,
Fletcher Creek near junction with Burdekin River, K.A. Williams 81081, 8 July 1981 (BRI); c. 10 km
N of the Gums, Condamine road, 27°14.63'S, 150°10.64'E, S. Jacobs 8356 & B. Hellquist, 28 Nov 1997
(BRI, NASC, NSW); 63 km SE of Home Hill, Bruce Hwy, 19°55' S, 147°50’ E, S. Jacobs 4086 16 May
1981 (NSW); 2.8 km from Kowanyama on the road to Shelfa Crossing, 15°28’S, 141°45'E,
J.R. Clarkson 9574 & V.J. Neldner, 26 May 1992 (BRI); c. 24 km N of Mareeba, Mt Molloy Road,
16°45'S, 145°20'E, S. Jacobs 4848 & G. Sainty, 1 May 1986 (MEL, NSW); 'Myall Park', Crater
Paddock, 27°13.39'S, 149°41.74'S. Jacobs 7131, 24 Apr 1994 (NSW); Tait River, C. Weld Birch & Ct.
Zelling s.n, 1892 (MEL); Fitzroy River Basin Brigalow Research Station, 20 miles [32 km] NW of
Theodore, R.W. Johnson 2619, 24 Apr, 1963 (BRI).

Grows in freshwater ephemeral habitats in drier regions, particularly gilgais in
Brigalow scrub, melon-holes, ponds, and roadside ditches. Flowers and fruits in
response to water levels and temperature, but more commonly during the warmer
months.

Aponogeton queenslandicus occasionally produces submersed leaves.

A population of A. vanbruggenii on the Atherton Tableland resembles A. queenslandicus
in producing almost all cordate-based floating leaves, but when the seeds are
examined it is clearly the former species.

7. A. kimberleyensis C.B. Hellquist & S.W.L. Jacobs, sp. nov.

A. euryspermo affinis, sed foliis submersis multo angustioribus, seminibus multo
minoribus, differt.

Holotype: Western Australia: water supply at Kuri Bay, 15°29'17"S, 124°29’54"E, A.A.
Mitchell 3501, 21 March 1994 (NSW). Isotype: Agriculture Broome.

Rooted, submerged, perennial aquatic. Leaves submersed; blades linear, rarely slightly
broadened, 24-32 cm long, 0.09-0.3 cm wide; dark green; margins flat; base long-
tapered, almost parallel-sided; apex obtuse to rarely retuse; 5-veined; lacunae band
prominent, to 2 mm wide, with up to 4 rows on each side of the midvein; petiole c. 5 cm
long. Peduncle to 32 cm long, 0.4-1 mm wide at base, broadening to 0.5-2.4 mm wide
at base of inflorescence. Spathe not seen. Inflorescence yellow, emergent, single, to 7.5 cm
long; rachis 0.5-2 mm wide. Flowers turned in all directions. Perianth segments 2,
spathulate, 1-veined, yellow, 1-1.4 mm long, 0.8 mm wide. Stamens 6; anthers c. 0.2 mm
long, 0.2 mm wide; filaments 1-1.5 mm long, broader at base. Carpels 3. Infructescence
slightly spaced along axis. Fruit 2.5-3.0 mm long, 2.0-2.8 mm wide, with an erect to
slightly curved terminal beak. Seeds (l-)2-3 per fruit, 0.7-1.4 mm long, 0.4-0.8 mm
wide; single testa with tight coat and fine reticulations.

Distribution: endemic to the western Kimberley of Western Australia.

Specimen examined: Western Australia: Langey, West Kimberley, 15°50’S, 124°23'E, G. Sainty s.n.,
25 May 1991 (NSW).

Known only from streams. Flowering and fruiting specimens from March and May.

This species is named for the Kimberley region of northwest Western Australia where
it grows.

Telopea 8(1): 1998

8. A. euryspermus C.B. Hellquist & S.W.L. Jacobs, sp. nov.

A. elongato affinis, sed seminibus multo majoribus, foliis interdum natantibus, differt.

Holotype: Western Australia: King Edward River Crossing, Mitchell Plateau Road,
14°54.07'S, 126°12.01’E, S. Jacobs 8030, 18 May 1996 (NSW). Isotypes: NASC, PERTH, Z.

Aponogeton elongatus forma strigosus H. Bruggen, Blumea 17: 133 (1969).

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

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 retuse; (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
(0.7-)2.2-4.8 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, 1-5 per fruit, usually 1 per small fruit,

2.3- 4.9 mm long, (l.l-)1.5-2.8 mm wide, thick outer testa easily removed, a second, if
present, tightly adhering and difficult to remove.

Distribution: northern portion of Western Australia from the Kimberley east to the
northern portion of the Northern Territory and Arnhem Land.

Selected specimens examined: Western Australia: Adcock Gorge, 16°56'S, 125°48’E, S. Jacobs 4388,
3 June 1982 (NSW); c. 3 km W of 'Ellenbrae' turnoff, Gibb River road, 15°59.47'S, 127°01.30'E,
S. Jacobs 8015, 17 May 1996 (NSW); feeder stream of the Drysdale River, 49 km north-west of 'Mt
Elizabeth' station, on track to Bachsten Creek, W.M. Molyncux & S.G. Forrester s.n., 30 June 1987
(MEL); 'Texas Downs' station, E of Mount John, K.A. Menkhorst 622, 28 July 1989 (DNA); c. 13 km
SE of 'Kimbolton' Hotel, 'Oobagooma' road 16°44'S, 123°54’E, S. Jacobs 5742, 26 May, 1988 (NSW);
Yampi Peninsula, in Trent River Gorge, 16°37'S, 123°42.5'E, PA. Fryxell, L.A. Craven & J. McD.
Steivart 4574, 31 May 1985 (MEL); 500 m west of Moochalabra Dam, 35 km south of Wyndham,
15°37'45"S, 128°05'05"E, A.A. Mitchell 3699, 27 July 1994 (NSW). Northern Territory: Arnhemland,
Mt Gilruth area, 13°03'S, 133°01'E, T.S. Henshall 1869, 6 June 1978 (DNA, MEL); lagoon at Jabiru,
12°40'S, 132°53'E, L.A. Craven 2277, 17 Feb 1973 (DNA, MEL); Kakadu Nat. Park, 13°32'S, 132°29'E,
/. Russell-Smith 8559, 7 Jan 1992 (DNA).

Grows in freshwater streams, rivers and lagoons. Flowering and fruiting December to
February and May to July.

This species is named for the large broad seed produced; from tire Greek eurys, broad
and spermus, seed.

Many specimens of Aponogeton from the tropics were originally assigned to A. elongatus
by Bruggen (1985) and others. These specimens represent taxa that initially produce
submerged leaves in flowing or clear water. Later in the year plants that we assign to
either A. euryspermus or A. vanbruggenii often develop floating leaves. These species
may be separated by seed shape and size; A. euryspermus has large broad seeds that
can become very thick when mature while A. vanbruggenii has longer and narrower

Hellquist and Jacobs, Aponogetonaceae of Australia

seeds with a distinct knob at one end. Both have seeds with a thin outer testa that is
easily removed. Both species are confined to the tropics with A. euryspermus more
westerly in its distribution and A. vanbruggenii more easterly.

9. A. vanbruggenii C.B. Hellquist & S.W.L. Jacobs sp. nov.

A. elongato affinis, sed seminibus plerumque complanatis, testis duabus, foliis
interdum natantibus, differt.

Holotype: Queensland: Mclvor River, 'Battle Camp' road, 15°07'13''S, 145°04'47"E,
S. Jacobs 8198, B. Hellquist & J. Wiersema (NSW). Isotypes: BRI, NASC, Z.

Rooted submersed perennial aquatic. Tuber ovate to elongate, to 0.5-4 cm long, 0.5-2.5 cm
wide. Leaves submerged and/or floating. Submersed blades lanceolate, elliptic, or
obovate, 7-36 cm long, 0.6-5 cm wide, green to maroon-green; margins slightly to
strongly undulate, rarely flat; base broadly cuneate, obtuse, rarely truncate; apex
obtuse to blunt-cuneate, 5-9(-ll)-veined; 0-2(-4) rows of lacunae on each side of the
midvein; petiole to 15(—23) cm long. Floating leaves elliptic or ovate to 12(-38) cm long,
0.7-2.5(-5) cm wide, green to maroon-green; margins flat; base cuneate, obtuse,
attenuate, or truncate, rarely cordate; apex blunt-cuneate, obtuse, mucronulate, rarely
cuspidate; 5-9 veined; petiole to 91 cm long. Peduncle to 50(-80) cm long, broadening
to 1.0-2.8(-4.2) mm wide at the base of inflorescence. Spathe to 1.5 cm long, caducous.
Inflorescence yellow, emergent or floating, single, tightly flowered to widely spaced
along rachis; rachis 1.1-3.2 mm wide. Flowers turned in all directions. Infructescence
moderately dense to spread out along rachis. Perianth segments 2, spathulate, rarely
obovate, 1-veined, yellow, 1.2-2.1 mm long, 0.7-1.5 mm wide. Stamens 6; anthers
c. 0.2-0.6 mm long, 0.2-0.6 mm wide; filaments 1-2.5 mm long, broader at base.
Carpels (2-)3(-5). Fruit elliptic, thin and papery; (2.7-)3.4-7.1 mm long, 2.2-4.5(-5.5)
mm wide, beak to 1 mm, mostly terminal, straight, bent, or reflexed, occasionally
lateral. Seeds obovate or narrowly elliptic (2-)3-5(-7) per fruit, 2.3-4.8 mm long, 0.7-1.6
mm wide, distinctly flattened or terete with a distinct knob at one end, with two testas;
outer testa loose and easily removed, inner testa usually easily removed.

Distribution: eastern tropical Northern Territory and Queensland.

Selected specimens examined: Northern Territory: Eastern Arnhem Land, Bralminar, 12°32’S,
135°55’E,y. Russell-Smith 8920, 28 Sept 1992 (DNA); Calvert River Gorge, 16°53'S, 137°22’E, P./C. Latz
10378, 30 Sept 1986 (DNA, MEL); 'Elsey' station, Roper River, 14°27 , S, 133°22'E, T.S. Henshall 2025,
19 June 1978 (DNA, MEL); King River at Victoria Highway Crossing, 14°42’S, 132°05'E, T.S.
Henshall 3547,29 March 1981 (DNA, MEL); 30 km E Mataranka, Jilkmingan area, 14°55'S, 133°26'E,
Wightman 4933 & D.R.L., 2 July 1990 (DNA).

Queensland: tributary of Gordon Creek, a tributary of Claudie River, 12°45'S, 143°15'E, A. W. Dockrill
459, 1 July 1972 (BRI); 6 km north of Mareeba, 16°56'S, 145°25'E, H.I. Aston 1961,25 May 1976 (MEL);
c. 16 km S of Mt Molloy, Mareeba road, 16°47.66'S, 145°21.49'E, S. Jacobs 8244, B. Hellquist &
j. Wiersema, 13 Aug 1997 (BRI, NASC, NSW); Pascoe River, 12°30'S, 143°16’E, AIM 635, 13 Aug 1979
(BRI); Port Douglas, Mowbray River c. 1 km S of Port Douglas turnoff, 2.2 km W of Port
Douglas-Cairns Highway, C.B. Hellquist 16194, C.E. Hellquist, L. Smith & E. Frye, 6 Oct 1997 (BRI,
NASC, NSW).

Grows in freshwater rivers, streams, still-water portions of lakes, and billabongs.
Flowers and fruits recorded from March to October and January.

This species is named to honour H.W.E. van Bruggen, the recognised world expert on
the genus Aponogeton.

A. vanbruggenii superficially resembles A. euryspermus (see discussion above) but
differs from that species in having smaller and narrower seed with a distinct knob at
one end. A. vanbruggenii, like A. euryspermus, also may regularly produce floating

Telopea 8(1): 1998

leaves in still or turbid waters, especially late in the season. A few populations of
A. vanbruggenii appear very similar to A. queenslandicus, with mainly floating leaves,
but the two species are easily distinguished by the seeds.

10. A. elongatus F. Muell. ex Bentham, FI. Austr. 7:188 (1878)

Holotype: Queensland: Brisbane River, F. Mueller s.n., Dec 1856 (K).

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 broadly
cuneate, obtuse, or rarely truncate; apex blunt-cuneate or obtuse; (5-)7-9(-ll)-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, 0.6-0.8 mm wide at base
broadening to 1.0-2.8(-4.2) 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, l-2(-2.7) mm long, 0.7-1.2 mm
wide. Stamens 6; anthers 0.3-0.4 mm long, 0.3A1.6 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.

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

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

10a. A. elongatus subsp. elongatus

As for species description except that floating leaves are rarely formed.

A. elongatus forma longifolius Bruggen, Blumea 17:10-11 (1969).

Distribution: coastal Queensland and coastal northeastern New South Wales.

Selected specimens examined: Queensland: Sheep Station Creek, Kilcoy, G. Trapnell s.n., 6 Feb 1957
(BR1); Portland Roads Road between Tozers Gap and the Middle Claudie Crossing, J.R. Clarkson
2923, 25 Feb 1980 (DNA, MEL); S arm of Maroochy River, W.G. Trapnell s.n., 20 Mar 1960 (BRI);
Yandina, J.H. Simmonds s.n., 1 Mar 1891 (BRI). New South Wales: Orara River, Coramba,
J.S. Boorman s.n., Nov 1912 (NSW); Nana Glen, Orara River, 30°08.01’S, 153°00.50'E, S. Jacobs 8346,

B. Hellquist, E. Bagley, 25 Nov 1997 (NASC, NSW); S of Lismore, S. Jacobs, 8354 & B. Hellquist,
27 Nov 1997 (NASC, NSW); Grafton, Miss Thornton s.n., (MEL 18784); Richmond River, Rudder s.n.,
Aug 1885 (NSW); Tintenbar, W. Baeuerlen, s.n., Oct 1894 (NSW).

Grows in freshwater portion of coastal rivers and streams. Flowers and fruits from
October to March.

10b. A. elongatus subsp. fluitans C.B. Hellquist & S.W.L. Jacobs, subsp. nov.

Subspecie elongato affinis, sed foliis plerumque natantibus differt.

Holotype: Queensland: Pine River near Petrie, S.T. Blake 2060, Dec 1930 (BRI).
Aponogeton elongatus forma latifolius H. Bruggen, Blumea 17:133,1969, (L).

Holotype: Queensland: Narangba, Sidling Creek, Gillieatt 151, 8 April 1964 (L). Isotype BRI.

Hellquist and Jacobs, Aponogetonaceae of Australia

Floating leaves common, 9-19 cm long, 2.2-3.5 cm wide, petiole to 90 cm long.

Distribution: localised in creeks and streams in the Petrie area of SE Queensland.

Selected specimens examined: Queensland: North Pine River, Petrie, 18 miles [30 km] N of
Brisbane, S.T. Blake 2060, Dec 1930 (BRI); Sideling Creek, Petrie, S.T. Blake 21022, 6 Oct 1959 (BRI,
MEL, NSW); Mt Coot-tha Botanic Gardens, Brisbane, S. Jacobs 2954, 7 Feb 1977 (NSW); Oyster Bay,
cultivated, originally from a Brisbane collector, A.N. Rodd & O. Campbell s.n., 18 Nov 1967 (NSW).

Grows in freshwater rivers and streams through rainforest. Flowers and fruits from
October through April.

This subspecies is named for the distinctive floating leaves; from the Latin fluito, float.

This subspecies might be mistaken for the newly described tropical species
A. euryspermus and A. vanbruggenii. Bruggen (1969) included two specimens of
A. vanbruggenii in his concept of A. elongatus forma latifolius. A. elongatus subsp.fluitans
is easily distinguished from the other two taxa by the distinctive seed shape and coat.
All three may produce floating leaves.

Aponogeton elongatus was previously considered a widely ranging species in Australia,
growing from the Kimberley region of Western Australia south and east to the
northeast corner of New South Wales. With this new treatment, specimens that we
here describe as A. euryspermus and A. vanbruggenii are removed from A. elongatus,
restricting it to the northeast portion of New South Wales and southeastern
Queensland, with a few populations in the Iron Range region of Cape York Peninsula.

Acknowledgments

We thank Lance Smith for his observations and specimens of various species under
cultivation, Karen Wilson for providing the Latin diagnoses, Nick Cuartero for
information on A. proliferus, Bill Lavarack from the Queensland National Parks for
help in obtaining collection permits, the International Water Lily Society for
contributing funds towards expenses for fieldwork, and the Royal Botanic Gardens,
Sydney for the support of C. B. Hellquist as a Visiting Research Fellow during his stay
in Sydney. We also thank John Clarkson for help in the field and for his comments on
the manuscript, and Frank Zofrea, Geoff Sainty, Greg Keighery and Nick Romanowski
for providing critical information and/or specimens.

References

Aston, H.I. (1973) Aquatic Plants of Australia. (Melbourne University Press: Melbourne).

Baas, P. (1985) Vegetative anatomy. Pp 3-4 in Bruggen, H.W.E. van. Monograph of the genus
Aponogeton (Aponogetonaceae). Bibliotheca Botanica Band XXXIII, Heft 137.

Bouman, F. (1985) Embryology. Pp 4-9 in Bruggen, H.W.E. van, Monograph of the genus
Aponogeton (Aponogetonaceae). Bibliotheca Botanica Band XXXIII, Heft 137.

Bruggen, H.W.E. van (1969) Revision of the genus Aponogeton (Aponogetonaceae): 111. The species
of Australia. Blumea 17:121-137.

Bruggen, H.W.E. van (1973) Revision of the genus Aponogeton (Aponogetonaceae): VI. The species
of Africa. Bull. / ard. Bot. Nal. Belg. 43: 193-233.

Bruggen, H.W.E. van (1985) Monograph of the genus Aponogeton (Aponogetonaceae). Bibliotheca
Botanica Band XXXIII, Heft 137.

Tomlinson, P.B. (1982) Anatomy of the Monocotyledons. Vll Helobiae (Alismatidae). (Claredon Press:
Oxford).

Manuscript received 18 February 1998
Manuscript accepted 30 July 1998

New combinations arising from a new
classification of non-African Restionaceae

Barbara G. Briggs and L.A.S. Johnson+

Abstract

Briggs, Barbara C. and Johnson, L.A.S. (Royal Botanic Gardens, Mrs Macquaries Road, Sydney, NSW 2000,
Australia) 1998. New combinations arising from a nezv classification of non-African Restionaceae. Telopea
8(1): 21-33. New combinations are made in accord with a new, broadly based classification of extra-
African Restionaceae. These involve 39 Australian species, of which two extend to New Guinea
and the Aru Islands, and one species in each of south-east Asia, New Zealand and Chile. The name
Baloskion Raf. is adopted for eight eastern Australian species excluded from Restio Rottb. and
Desmocladus Nees is adopted for a group of Western Australian species mostly transferred from
Loxocarx/a R. Br. The previously monotypic genera Meeboldina Suesseng. and Sporadanthus F. Muell.
are enlarged. Most other changes involve newly described genera. Four combinations replace
illegitimate epithets, two new combinations are made at subspecific rank, and lectotypes are
selected for 18 taxa.

Introduction

A new classification of the genera and species of Australian Restionaceae has been
developed through study of exomorphology, anatomy, pollen, seed ornamentation,
and flavonoids, with associated DNA sequence studies in progress. The classification
is outlined by Briggs and Johnson (1999) and Linder, Briggs and Johnson (1998). It has
led to the recognition of 16 new genera (Briggs & Johnson 1998) and has shown that
the species hitherto included in some of the genera are unnatural assemblages of taxa.
The largest group of inappropriately placed species have until now been referred to
Restio Rottb. These were thus given the name of a genus that, as now circumscribed
(Linder 1984,1985; Linder, Briggs & Johnson 1998; Briggs & Johnson 1999), is confined
to Africa and Madagascar. In addition, the epithets in use in some combinations were
illegitimate.

Forty two new combinations at specific rank are therefore provided here, together
with two new subspecific combinations. This will validate many of the names used in
the forthcoming book Australian Rushes — Biology, Identification and Conservation of
Restionaceae and allied families (Meney & Pate (eds) 1999). In addition, there still remain
45 undescribed species that we recognise in Australian Restionaceae, on which
manuscripts are being prepared.

Comparison of the features of the genera from which species have been removed and
those in which they are now placed will indicate many of the reasons for the new
placements. These features are summarised in Linder et al. (1998), Briggs and Johnson
(1999) and, for the newly described genera, in Briggs and Johnson (1998). The new
classification is, in general, supported by morphological and molecular cladistic
studies (Linder et al. in press; Briggs et al. in press). It will also be outlined in the
treatment in the Flora of Australia and future publications.

t Deceased 1 August 1997.

Telopea 8(1): 1998

New combinations and lectotypifications

Types of almost all taxa have been seen and will be cited in the treatment of
Restionaceae in the Flora of Australia, now in preparation. Types are therefore cited
here only for non-Australian species and in those cases where lectotypification is
desirable. Unusually large numbers of lectotypifications have been called for, since
both male and female plants have often been included in the type material of these
dioecious species. Choice of lectotypes has taken into account agreement with the
protologue and any comments therein, annotations by the author of the epithet and
certainty as to the identity of the specimen. Priority has been given to specimens that
exhibit a wide range of the features distinguishing the species and that are represented
in several herbaria. Female specimens are often selected since they show more of the
features characterising genera; but in some cases male specimens have been selected
where they show more distinctive specific features or agree more closely with the
protologue. Unless indicated otherwise, types cited here have been seen. Except for
the basionym, synonyms are not generally listed, apart from instances where another
name for the taxon has been in recent use. An index is provided of new combinations
and cited synonyms.

Acion

Acion B.G. Briggs & L.A.S. Johnson, Telopea 7: 353 (1998).

This Tasmanian genus consists of two species that were formerly included in Restio.
Type species: A. monocephalum (R. Br.) B.G. Briggs & L.A.S. Johnson.

Acion hookeri (D.I. Morris) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Restio hookeri D.I. Morris in M.R. Banks et al. (eds). Aspects Tasmanian Bot.,
Tribute to Winifred Curtis: 33 (1991).

Apodasmia

Apodasmia B.G. Briggs & L.A.S. Johnson, Telopea 7: 371 (1998).

A far-flung genus of three or four species: one (undescribed) in Western Australia and
one recognised in each of south-eastern Australia (including Tasmania), New Zealand
and Chile. A. chilensis and A. similis show such a close resemblance that their status
requires further study. They are, however, maintained here as distinct species. Type
species: A. brownii (Hook, f.) B.G. Briggs & L.A.S. Johnson, of south-eastern Australia.

Apodasmia chilensis (Gay) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Schoenodum chilense Gay, FI. chilena 6: 152 (1854).

Calopsis chilensis (Gay) Steud., Syn. pi. glum. 2: 258 (1855).

Leptocarpus chilensis (Gay) Mast., Monogr. Phan. 1: 341 (1878).

Type: Chile: ad fluv Rio nigro Arigue, Chili, Lechler 618 9 (P, ex Herb. Steudel).
Probable iso: Chile, (K ex P, annotated with the name 'Gay' and originally determined
Schoenodum chilense).

Apodasmia similis (Edgar) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Leptocarpus similis Edgar, New Zealand J. Bot. 6: 468 (1969).

Type: New Zealand: Brooklands lagoon, near mouth of Waimakariri R., north end
opposite Spencer Park, E. Edgar, 20 Dec 1967 (CHR, not seen).

Briggs and Johnson, New combinations in Restionaceae

Baloskion

Baloskion Raf., Flora telluriana 4: 32 (1838).

The name Baloskion is adopted for a group of eight eastern Australian species hitherto
included in Restio. This usage was foreshadowed by Quirico and Briggs (1993). The
extensive differences in anatomy between these and the African species that are
correctly placed in Restio were noted by Cutler (1969). A distinctive feature of Baloskion,
the adnation of the pedicel to the subtending glume, was clearly illustrated when its
first species was described by Labillardiere (1806, t. 227). All species require new
combinations, the type species of Baloskion having only an illegitimate combination
within that genus. Type species: B. dichotomum Raf., nom. illeg. = B. tetraphyllum
(Labill.) B.G. Briggs & L.A.S. Johnson.

Baloskion australe (R. Br.) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Restio australis R. Br., Prodr.: 245 (1810).

Type citation: (D.) v.v.

Type: Tasmania: In summitate Montis Tabularis [Mt Wellington] prope fl. Derwent,
R. Broivn (Bennett 5868), Apr. 1804 6 (lecto, here selected, BM). Residual syntypes: inter
Storm Bay passage, fl. Derwent, R. Brown, Apr. 1804 9 (BM); Derwent R., R. Brown (E);
van DJiemens] Land, R. Brown 6, 9 (K).

Baloskion fimbriatum (L.A.S. Johnson & O.D. Evans) B.G. Briggs & L.A.S. Johnson,

comb. nov.

Basionym: Restio fimbriatus L.A.S. Johnson & O.D. Evans, Contr. New South Wales Natl
Herb. 3: 210 (1963).

Baloskion gracile (R. Br.) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Restio gracilis R. Br., Prodr.: 245 (1810).

Baloskion longipes (L.A.S. Johnson & O.D. Evans) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Restio longipes L.A.S. Johnson & O.D. Evans, Contr. New South Wales Natl
Herb. 3: 208 (1963).

Baloskion pallens (R. Br.) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Restio pallens R. Br., Prodr.: 245 (1810).

Baloskion stenocoleum (L.A.S. Johnson & O.D. Evans) B.G. Briggs & L.A.S. Johnson,

comb. nov.

Basionym: Restio stenocoleus L.A.S. Johnson & O.D. Evans, Contr. New South Wales
Natl Herb. 3: 205 (1963).

Baloskion tenuiculme (S.T. Blake) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Restio tenuiculmis S.T. Blake, Contr. New South Wales Natl Herb. 4:198 (1963).
Baloskion tetraphyllum (Labill.) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Restio tetraphyllus Labill., Nov. Holl. pi. 2: 77 (1806).

Type citation: in capite Van-Diemen.

Type: Tasmania: Van Diemen, Labillardiere 9 (lecto, here selected, P, photo NSW).
Possible isolecto: in capite Van-Diemen 9 (BM). Residual syntypes: Van Diemen,
Labillardiere 6 (P, on same sheet as lecto); in capite Van-Diemen, Labillardiere 6 (BM);
Nova Hollandia, Labillardiere 6 (K).

Telopea 8 ( 1 ): 1998

Bnloskion dichotomum Raf., Flora telluriana 4: 32 (1838), nom. illeg.

Two subspecies are recognised:

Baloskion tetraphyllum (Labill.) B.G. Briggs & L.A.S. Johnson subsp. tetraphyllum

Baloskion tetraphyllum subsp. meiostachyum (L.A.S. Johnson & O.D. Evans) B.G.
Briggs & L.A.S. Johnson, comb. nov.

Basionym: Restio tetraphyllus subsp. meiostachyus L.A.S. Johnson & O.D. Evans, Contr.
New South Wales Natl Herb. 3: 220 (1963).

Chaetanthus

Chaetanthus R. Br., Prodr. 251 (1810).

The transfer of two species from Leptocarpus enlarges this genus, which occurs in the
south of Western Australia; it has hitherto included only a single species. Type species:
C. leptocarpoides R. Br.

Chaetanthus aristatus (R. Br.) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Leptocarpus aristatus R. Br., Prodr.: 250 (1810).

Chaetanthus tenellus (Nee s) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Restio tenellus Nees in Lehm., PI. Preiss. 2: 57 (1846).

Leptocarpus tenellus (Nees) F. Muell., Fragm. 8: 90 (1873).

Chordifex

Chordifex B.G. Briggs & L.A.S. Johnson, Telopea 7: 356 (1998).

A genus of 16 species in the south of Western Australia. Ten of the species were
previously referred to Restio, while five are undescribed. Type species: C. stenandrus
B.G. Briggs & L.A.S. Johnson.

Chordifex abortivus (Nees) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Restio abortivus Nees in Lehm., PI. Preiss. 2: 60 (1846).

Chordifex amblycoleus (F. Muell.) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Restio amblycoleus F. Muell., Fragm. 8: 65 (1873).

Type citation: In Australia occidentali, J. Drummond 66.

Type: Western Australia: Drummond 66 3 (MEL 14733 lecto, here selected; iso K).
Residual syntype: Drummond 66 9 (MEL 14730, iso K).

Chordifex chaunocoleus (F. Muell.) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Restio chaunocoleus F. Muell., Fragm. 8: 64 (1873).

Type citation: In Australia occidentali, J. Drummond 948, 949.

Type: Western Australia: Drummond 949 9 (lecto, here selected, MEL 14744, iso MEL
14742, B, BM, E, K). Residual syntype: Drummond 948 3 (MEL 14743, MEL 14745, iso
B, BM, E, K).

Chordifex crispatus (R. Br.) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Restio crispatus R. Br., Prodr.: 246 (1810).

Type citation: (M.) v.v.

Briggs and Johnson, New combinations in Restionaceae

Type: Western Australia: Bay 1 [Lucky Bay], R. Brown 6 (lecto, here selected BM; iso E,
K (two sheets), MEL 14749, 14750, 15016, P). Residual syntype 9 (BM, mounted on
same sheet as lecto).

Chordifex gracilior (F. Muell. ex Benth.) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Restio gracilior F. Muell. ex Benth., FI. austral. 7: 226 (1878). Bentham's
publication of Mueller's name (March 1878) predates that of Masters, Monogr. Phan.
1: 297 (June 1878).

Type citation: W. Australia, Drummond n. 68 and 71

Type: Western Australia: Drummond 71 <5 (lecto, here selected, K; iso MEL 14754;
probable iso MEL 14752-3,14755-7). Residual syntype: Swan River, Drummond 68 6 (K).

Chordifex isomorphus (K. W. Dixon & K. A. Meney) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Restio isomorphus K. W. Dixon & K. A. Meney, Nuytsia 9: 91 (1993).

Chordifex laxus (R. Br.) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Restio laxus R. Br., Prodr.: 245 (1810) ( Restio, species number 3), non R. laxus
R. Br. op. cit. p. 246 ( Restio, species 12). The latter was renamed Restio diffusus Sprengel,
Syst. Veg. 1:185 (1824), see below under Leptocarpus diffusus.

Type: The type sheet (BM, photo NSW) may include both & and $ pieces but this is
not certain since the spikelets of males and females are externally similar. The sheet is
marked 6 but the protologue refers to the styles, implying the presence of females. No
lectotypification is made here.

Chordifex leucoblepharus (Gilg) B.G. Briggs &t L.A.S. Johnson, comb. nov.

Basionym: Restio leucoblepharus Gilg, Bot. Jahrb. Syst. 35: 88 (1904).

Chordifex ornatus (Steud.) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Restio ornatus Steud., Syn. pi. glumac. 2: 256 (1855).

Chordifex sphacelatus (R. Br.) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Restio sphacelatus R. Br., Prodr.: 245 (1810).

Dapsilanthus

Dapsilanthus B.G. Briggs & L.A.S. Johnson, Telopea 7: 369 (1998).

A genus of four species; three in northern Australia and southern New Guinea (two of
these also in the Aru Islands), as well as one in south-east Asia (Malaysia, Cambodia,
Thailand, Vietnam and the south-eastern Chinese island of Hainan). Type species: D.
elatior (R. Br.) B.G. Briggs & L.A.S. Johnson.

Dapsilanthus disjunctus (Mast.) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Leptocarpus disjunctus Mast., J. Linn. Soc. Bot. 17: 344 (1879).

Type citation: in insula 'Phu', Cochin China, Godefroy-Lebceuf 928.

Type: Vietnam: Cochin-China, village de Bac, He de Phu Quoc, Godefroy-Lebceuf 928,
Sep [18]78 $ (holo K). (The protologue describes the species as monoecious; it is in
general dioecious. A possible isotype (P) is accompanied by the note Tower spikelets
9 upper S'.) We observed only $ flowers.

Dapsilanthus ramosus (R. Br.) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Leptocarpus ramosus R. Br., Prodr.: 250 (1810).

Telopea 8 ( 1 ): 1998

Type citation: (T.) B. v.s.

Type: Queensland: Endeavour R., Banks & Solander, 1770 $ (lecto, here selected, BM;
iso B, BRI, MEL 14439 p.p., NSW 78873). Residual syntype 8 (BM, on same sheet as
lectotype; iso MEL 14439 p.p., NSW 147064, P).

Dapsilanthus spathaceus (R. Br.) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Leptocarpus spathaceus R. Br., Prodr.: 250 (1810).

Type citation: (T.) v.v.

Type: northern Australia: Iter Austral., R. Brown (Bennett No. 5874p.p.), 1802-5 2 (lecto,
here selected, BM; iso BRI, K (2 sheets); probable iso MEL 15103 p.p. Residual syntype:
Iter Austral., R. Brown (Bennett No. 5874p.p.), 1802-5 8 (BM, iso BRI, K; probable iso
MEL 15103 p.p.).

Leptocarpus schultzii Benth., FI. austral. 7: 237 (1878).

Leptocarpus barbatus K. Bakker, FI. Males, ser. I, 5(4): 419 (1957).

Type: Aru Islands: P. Trangan, Cape Meroor, Buwalda 5531, 9 July 1938 8 (holo L; iso
BO, GH, K).

Desmocladus

Desmocladus Nees in Lehm., PI. Preiss. 2: 56 (1846).

The name adopted for this genus, although published in 1846, was not taken into use
until recently (Meney, Pate & Dixon 1996; Linder, Briggs & Johnson 1998; Briggs &
Johnson 1999). The 15 species occur in the south of Western Australia, with one species
also in Eyre Peninsula, South Australia. Except for D. glomeratus K.W. Dixon & K.A.
Meney and D. asper (see below), all the described species were previously included in
Loxocarya R. Br. which has markedly different features and is typified by L. cinerea R.
Bi. Nine of the species are undescribed. The type species is among those requiring a
new combination; its previous combination under Desmocladus being illegitimate.
Type species: D. brunonianus Nees, nom. illeg. = D. fasciculatus (R. Br.) B.G. Briggs &
L.A.S. Johnson.

Desmocladus asper (Nees) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Calorophus asper Nees in Lehm., PL Preiss. 2: 67 (1846).

Type citation:... haud longe ab ore maris (Perth) m. Septembri a. 1839 [Preiss 1716 p.p.]
et... praedia rustica v. ell. Barker et Lennard m. April a. 1840 [Preiss 1694].

Type: Western Australia: Preiss 1716 p.p. 8 (lecto, here selected, excluding material of
Hypolaena pubescens under this number, LD; iso MEL 14861 p.p., MO, P). Residual
syntype Preiss 1694 8 (LD, MEL 14861p.p., BM, K, MO).

Desmocladus fasciculatus (R. Br.) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Restio fasciculatus R. Br., Prodr.: 247 (1810).

Desmocladus brunonianus Nees in Lehm., PI. Preiss. 2: 56 (1846), nom. illeg., based on
Restio fasciculatus R. Br.

Loxocarya fasciculata (R. Br.) Benth., FI. austral. 7: 242 (1878).

Desmocladus flexuosus (R. Br.) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Restio flexuosus R. Br., Prodr.: 247 (1810).

Loxocari/a flexuosa (R. Br.) Benth., FI. austral. 7: 243 (1878).

Briggs and Johnson, New combinations in Restionaceae

Type citation: (M.) v.v.

Type: Western Australia: King George Sound, R. Brown 1802-5 (Bennett No. 5849 p.p.)
6 (lecto, here selected, BM, the lectotype consists of the fertile male specimens on the
sheet, excluding the vegetative piece on right of sheet which may be an allied species
of Desmocladus, and also the central specimen in the lower half of the sheet which is
Empodisma gradllimum (F. Muell.) L.A.S. Johnson & D.F. Cutler.; isolecto BM).

Desmocladus myriocladus (Gilg) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Loxocarya myriodada Gilg, Bot. Jahrb. Syst. 35: 90 (1904).

Type citation: in distr. Avon pr. Tammin ... (D. 5070); in distr. Eyre pr. Graspatch ... flor.
m. Nov. (D. 5296).

Type: Western Australia: Avon, Tammin, L. Diels 5070, 24 Nov. 1901 6 (lecto, here
selected, B; iso NSW). The residual syntype (examined in B) is an allied undescribed
species of Desmocladus. The lectotypification maintains the name for the more
widespread and abundant species.

Desmocladus virgatus ( Benth.) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Loxocarya virgata Benth., FI. austral. 7: 242 (1878).

Type citation: W. Australia, Drummond, n. 74 and 113. 'Another 113 however of the
same collector appears to be the male of some Hypolaena’ (Bentham).

Type: Western Australia: Swan River, Drummond 113p.p. 6 (lecto, here selected,
annotated by Bentham as Loxocarya virgata, K, photo NSW). Residual syntype: Swan
River, Drummond 74 6 (K).

Dielsia

Dielsia Gilg, Bot. Jahrb. Syst. 35: 88 (1904).

A distinctive monotypic genus of the south of Western Australia. The type species
requires a valid combination.

Dielsia stenostachya (W. Fitzg.) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Restio stenostachyus W. Fitzg., Proc. Linn. Soc. New South Wales 28: 108
(1903).'

Type citation: Burswood; in wet spots (2; in March 1900) near Causeway, Perth (6;
April 1901; W.V. Fitzgerald).

Type: Western Australia: Causeway near Perth, W.V. Fitzgerald, April 1901 6 (lecto,
here selected, NSW 91587; iso PERTH). Residual syntype: Burswood, E of Causeway,
Perth, W.V Fitzgerald, March 1901 2 (NSW 91959, iso PERTH) [this appears to be
Fitzgerald's 2 syntype, despite the discrepancy in date].

Dielsia cygnorum Gilg, Bot. Jahrb. Syst. 35: 88 (1904), nom. illeg.

Syntypes: Western Australia: in distr. Darling pr. Swan River (Bayswater), Pritzel 304,
May 1901 6 (B, iso AD, E, K, MO, NSW, PERTH [labelled 340], US); Diels 28166 13 May
1901 2 (B, iso K).

Harperia

Harperia W. Fitzg., J. W. Austral. Natural. Hist. Soc. 1: 34 (1904).

A genus of four species, one of them undescribed, in the south of Western Australia.
Type species: H. lateriflora W.V. Fitzg.

Telopea 8(1): 1998

Harperia confertospicatus (Steud.) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Restio confertospicatus Steud., Syn. pi. glumac. 2: 256 (1855).

Hypolaena

Hypolaena R. Br., Prodr.: 251 (1810).

A genus of eight species, three of them undescribed, in the south of Western Australia,
one of them (H. fastigiata R. Br.) also occurring in eastern Australia from southern
Queensland to South Australia and Tasmania. Lectotype species: H. fastigiata R. Br.

Hypolaena humilis (Gilg) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Leptocarpus humilis Gilg, Bot. Jahrb. Syst. (1904: 89).

Type citation: in distr. Stirling pr. Cranbrook ... fl. m. Sept. (D. 4433).

Type: Western Australia: Cranbrook, Diels 4433b, 24.9.1901 $ (lecto, here selected, B;
iso P). Residual syntype: Cranbrook, Diels 4433a, 24.9.1901 6 (B; iso P).

Lepidobolus

Lepidobolus Nees in Lehm., PI. Preiss. 2: 66 (1846).

A genus of eight species, two of them undescribed; one species including three
subspecies (one undescribed). Seven species occur in the south of Western Australia
and one (H. drapetocoleus F. Muell.) in the east of South Australia and in western
Victoria. Type species: L. preissianus Nees.

Lepidobolus preissianus Nees in Lehm., PI. Preiss 2: 66, (1846).

Note on typification: Preiss numbers 1755 and 1756 ex parte are given in the protologue
as syntypes of the species and Preiss 1757 as the type of var. volubilis. By contrast, Nees
(in the index of collections) in Lehm., PI. Preiss. 2: 408 (1846) and Masters, Monogr.
Phan. 1: 347 (1878) list Preiss 1757 as an example of the species and Preiss 1756 as var.
volubilis. The features of the specimens and Preiss's annotations of the sheets indicate
that the collection numbers cited in the protologue are correct.

Type citation: In arenosis clivuli Bellevue ad flumen Cygnorum m. Augusto a 1839 et
in... destrictus [districtus] York, m. Septembri a. 1839 i, ? Preiss No. 1755 et 1756 ex parte.

Type: Western Australia: Bellevue ... Preiss 1755 p.p. $ (lecto, here selected, LD; iso
MEL 14716 p.p.). Most sheets include 6 and 9 material (isolecto MEL 14711, $
material of MEL 14712-4 p.p.). Residual syntypes: Preiss 1756 6, $ (LD; iso MEL
14714); S material of Preiss 1755 (MEL 14712-4,14716 p.p.).

Two subspecies are recognised:

Lepidobolus preissianus subsp. preissianus

Lepidobolus preissianus Nees var. preissianus

Lepidobolus preissianus subsp. volubilis (Nees) B.G. Briggs & L.A.S. Johnson, comb,
et stat. nov.

Basionym: Lepidobolus preissianus Nees var. p volubilis Nees in Lehm., PI. Preiss. 2: 66
(1846).

Type citation: In districtu York cum No. 1756, Herb. Preiss No. 1757.

Type: Western Australia: Preiss 1757 ,10 Sep [18]39 6 (LD; iso MEL 14715).

Briggs and Johnson, New combinations in Restionaceae

Leptocarpus

Leptocarpus R. Br., Prodr.: 250 (1810: 250), (nom. cons.)

Currently the lectotype species of Leptocarpus is L. aristatus R. Br. but it is our intention
to seek conservation of a new type to avoid change to the name of the most
widespread of the species and to maintain the use of Chaetanthus R. Br. Our studies
have led us to restrict Leptocarpus, if typified by L. tenax (Labill.) R. Br., to three species.
All occur in the south of Western Australia, with L. tenax occurring also in eastern
Australia from southern Queensland to South Australia and Tasmania.

Leptocarpus diffusus (Spreng.) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Restio diffusus Spreng., Syst. Veg. 1:185 (1825) ( Restio laxus R. Br., Prodr.: 246
(1810) (Restio species number 12) non R. Br., Prodr.: 245 (1810) ( Restio species number 3)).

Type citation: (M.) v.v.

Type: Western Australia: 'King George III d Sd', R. Brown 6 (BM; iso P). The BM sheet
bears labels 'King George III d Sd' and 'Port Jackson', the latter stuck to a slip labelled
'12' and 'Bennett 5860', both locality labels with 'Restio laxus' in Brown's hand. The
material appears to represent a single collection of this Western Australian species,
and not to require lectotypification.

Loxocarya

Loxocarya R. Br., Prodr.: 249 (1810).

A genus of five species, one of them undescribed, in the south of Western Australia. As
noted above, five species previously referred to Loxocarya (although originally
described under various genera) are now transferred to Desmocladus. Type species:
L. cinerea R. Br.

Loxocarya striata (F. Muell.) B.G, Briggs & L.A.S. Johnson, comb. nov.

Basionym: Megalotheca striata F. Muell., Fragm. 8: 99 (1873).

[Restio megalotheca F. Muell. ex Benth., FI. austral. 7: 222 (1878), nom. illeg.J

Type citation: In Australiae plagis occidentalibus; /. Drummond.

Type: Western Australia: Drummond s.n. 9 (lecto, here selected, MEL 14774; iso K).
Residual syntypes: Drummond 186 $ (MEL 14776, 14778), 450 (MEL 14773, 14775).
Probable isotypes: Drummond 950, 1843 6 (BM, E), Drummond 951, 1843 9 (GH, P),
Swan R., Drummond 100 6 (K), Drummond 103 5 (E) 9 (K).

Meeboldina

Meeboldina Suess., Boissiera 7: 20 (1943).

This genus, of the south of Western Australia, was previously regarded as monotypic
but is now enlarged by the transfer of four species from Leptocarpus. A further six
species remain to be described. Type species: M. denmarkica Suess.

Meeboldina cana (Nees) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Leptocarpus canus Nees, Ann. Mag. Nat. Hist. 6: 50 (1841).

Type citation: ad Flumen Cygnorum lectae.

Telopea 8(1): 1998

Type: Drummond 1705 ex parte 2 (LD; iso CGE). Residual syntype: Drummond 1862 6
(LD; iso K).

L. catius was also published by Nees in Lehm. PL Preiss. 2: 64 (1846) with the citation
'L. et N.' [Lindl. & Nees] but with no reference to the earlier publication. The original
publication of the basionym had Nees as author but was communicated to the journal
by Lindley.

Meeboldina coangustata (Nees) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Leptocarpus coangustatus Nees in Lehm., PI. Preiss. 2: 65 (1846).

Type citation: 6 In depressis uliginosis prope Halfwayhouse, Darling's-range, m.
Septembri a. 1839 ..., Preiss 1708. 2 In uliginosis planitiei ad fluvium Cygnorum supra
oppidulum Perth, m. Octobri a. 1839 ... Preiss 1706. Drummond in Herb. Lindl.

Type: Western Australia: ad fluvium Cygnorum supra oppidulum Perth, Preiss 1706 2
(lecto, here selected, LD; iso MEL 14416, P). Residual syntypes: prope Halfwayhouse,
Darling's-range, Preiss 1708 (LD, B, P); Swan R., Drummond, 1839 2 (K).

Meeboldina crassipes (J.S. Pate & K.A. Meney) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Leptocarpus crassipes J.S. Pate & K.A. Meney, Telopea 6: 658 (1996).

Meeboldina scariosa (R. Br.) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Leptocarpus scariosus R. Br., Prodr.: 250 (1810).

Sporadanthus

Sporadanthus F. Muell., Trans. & Proc. New Zealand Inst. 6: 389 (1874).

This was considered a monotypic New Zealand endemic until the relationship was
recognised between S. traversii and the Australian species of Lepyrodia 'Group B' of
Johnson and Evans (1963). S. strictus and one undescribed species occur in the south
of Western Australia but the other five species now transferred occur in eastern
Australia from southern Queensland to Tasmania and western Victoria. Type species:
S. traversii (F. Muell.) F. Muell. ex Kirk.

Sporadanthus caudatus (L.A.S. Johnson & O.D. Evans) B.G. Briggs & L.A.S. Johnson,

comb. nov.

Basionym: Lepyrodia caudata L.A.S. Johnson & O.D. Evans, Contr. New South Wales
Natl Herb. 3: 226 (1963).

Sporadanthus gracilis (R. Br.) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Lepyrodia gracilis R. Br., Prodr.: 247 (1810).

Sporadanthus interruptus (F. Muell.) B.G. Briggs & L.A.S. Johnson, comb. nov.
Basionym: Lepyrodia interrupta F. Muell., Fragm. 8: 74 (1873).

Type citation: In insula Moreton's Island; F.M.

Type: Queensland: Moreton Island, F. Mueller, 1855 2 (lectotype, here selected, MEL
707408, iso MEL 707409-707411, BM, K). Residual syntype: <J (K, P).

Sporadanthus strictus (R. Br.) B.G. Briggs & L.A.S. Johnson, comb. nov.

Basionym: Lepyrodia stricta R. Br., Prodr.: 248 (1810).

Briggs and Johnson, New combinations in Restionaceae

Sporadanthus tasmanicus (Hook.f.) B.G. Briggs & L.A.S. Johnson, comb. nov.
Basionym: Lepyrodia tasmanica Hook, f., FI. Tasm. 2: 72 (1858).

Type citation: Gunn, 960, 1393.

Type: Tasmania: Detention R. nr Rocky Cape, Gunn 960, 16.12.[18]36 2 (lecto, here
selected, K). Residual syntypes: Detention R., Gunn 960, 16.12.[18]36 S (K); R.C. Gunn
960, 1837 <? (K); Gunn 960 (CGE ex Lindley); Tasmania, Lake St Clair, Gunn 1393,
7.1.[18]41 <S (K).

Acknowledgments

Many people assisted in the studies that led to the present paper. Thanks are due
especially to Carolyn Porter, Siegfried Krauss, Anna-Louise Quirico, Barbara Wiecek
and Louisa Murray. Karen Wilson assisted with photographs of specimens in several
herbaria. The opportunity to examine specimens on loan or in other herbaria is
gratefully acknowledged, as is the assistance of Anna Hallett and Miguel Garcia of the
Royal Botanic Gardens Sydney Library. Grants from the Australian Biological
Resources Study provided valuable technical help.

References

Briggs, B.G. & Johnson, L.A.S. (1999) A guide to a new classification of Restionaceae and allied
families. In Meney, K.A. & Pate, J.S. (eds) Australian Rushes — Biology, Identification and
Conservation of Restionaceae and alliedfamilies. (University of Western Australia Press: Nedlands).
Briggs, B.G. & Johnson, L.A.S. (1998) New genera and species of Australian Restionaceae (Poales).
Telopea 7(4): 345-373.

Briggs, B.G., Marchant, A.D., Gilmore, S. & Porter, C.L. (in press). A molecular phylogeny of
Restionaceae and allies. In Wilson, K.L. & Morrison, D. (eds) Systematics and Evolution of
Monocots (Proceedings 2nd Monocot Symposium).

Cutler, D.F. (1969) Juncales. In Metcalfe, C.R. (ed.) Anatomy of Monocotyledons, vol. 4. (Clarendon
Press: Oxford).

Johnson, L.A.S. & Evans, O.D. (1963) Intrageneric groups and new species in Lepyrodia. Contr. New
South Wales Natl Herb. 3: 223-227.

Labillardiere, J.J.H. de. (1806) Novae Hollandiae Plantarum Specimen, part 23.

Linder, PI.P. (1984) A phylogenetic classification of the genera of the African Restionaceae. Bothalia
15:11-76.

Linder, H.P. (1985) Conspectus of the African species of Restionaceae. Bothalia 15: 387-503.

Linder, H.P., Briggs, B.G. & Johnson, L.A.S. (1998) Restionaceae. Pp. 425-445 in Kubitzki, K. (ed.)

The Families and Genera of Vascular Plants, vol. 4. (Springer-Verlag: Berlin).

Linder, H.P., Briggs, B.G. & Johnson, L.A.S. (in press). Restionaceae —a morphological phylogeny.
In Wilson, K.L. & Morrison, D. (eds) Systematics and Evolution of Monocots (Proceedings 2nd
Monocot Symposium).

Meney, K.A. & Pate, J.S. (eds) (1999) Australian Rushes — Biology, Identification and Conservation of
Restionaceae and allied families. (University of Western Australia Press: Nedlands).

Meney, K.A., Pate, J.S. & Dixon, K.W. (1996) New species of Restionaceae from Western Australia.
Telopea 6: 649-666.

Quirico, A.L. & Briggs, B.G. (1993) Restionaceae. Pp. 396-407 in Harden, G.J. (ed.) Flora ofNeiu South
Wales, vol. 4. (New South Wales University Press: Kensington).

Manuscript received 18 June 1998
Manuscript accepted 30 July 1998

Telopea 8(1): 1998

Index to new combinations, basionyms and cited synonyms

New combinations are printed in boldface; synonyms are printed in italics.

Acion . 22

Acion hookeri. 22

Apodasmia . 22

Apodasmia chilensis. 22

Apodasmia similis . 22

Baloskion . 23

Baloskion australe. 23

Baloskion dichotomum . 24

Baloskion fimbriatum. 23

Baloskion gracile . 23

Baloskion longipes . 23

Baloskion pallens. 23

Baloskion stenocoleum . 23

Baloskion tenuiculme. 23

Baloskion tetraphyllum . 23

Baloskion tetraphyllum

subsp. meiostachyum . 24

Baloskion tetraphyllum

subsp. tetraphyllum . 24

Calopsis chilensis . 22

Calorophus asper . 26

Chaetanthus . 24

Chaetanthus aristatus . 24

Chaetanthus tenellus . 24

Chordifex . 24

Chordifex abortivus . 24

Chordifex amblycoleus . 24

Chordifex chaunocoleus . 24

Chordifex crispatus. 24

Chordifex gracilior . 25

Chordifex isomorphus . 25

Chordifex laxus . 25

Chordifex leucoblepharus . 25

Chordifex ornatus . 25

Chordifex sphacelatus . 25

Dapsilanthus . 25

Dapsilanthus disjunctus . 25

Dapsilanthus ramosus . 25

Dapsilanthus spathaceus . 26

Desmocladus. 26

Desmocladus asper . 26

Desmocladus brunonianus . 26

Desmocladus fasciculatus . 26

Desmocladus flexuosus . 26

Desmocladus myriocladus . 27

Desmocladus virgatus . 27

Dielsia. 27

Dielsia cygnorum . 27

Dielsia stenostachya . 27

Harperia. 27

Harperia confertospicatus . 28

Hypolaena . 28

Hypolaena humilis . 28

Lepidobolus. 28

Lepidobolus preissianus

subsp. preissianus . 28

Lepidobolus preissianus

subsp. volubilis . 28

Lepidobolus preissianus

var. preissianus . 28

Lepidobolus preissianus

var. |3 volubilis . 28

Briggs and Johnson, New combinations in Restionaceae 33

Leptocarpus. 29

Leptocarpus aristatus . 24

Leptocarpus barbatus . 26

Leptocarpus canus „. 29

Leptocarpus chilensis . 22

Leptocarpus coangustatus . 30

Leptocarpus crassipes . 30

Leptocarpus diffusus . 29

Leptocarpus disjunctus . 25

Leptocarpus Immilis . 28

Leptocarpus ramosus . 25

Leptocarpus scariosus . 30

Leptocarpus schultzii . 26

Leptocarpus similis . 22

Leptocarpus spathaceus . 26

Leptocarpus tenellus . 24

Lepyrodia caudata . 30

Lepyrodia gracilis . 30

Lepyrodia interrupta . 30

Lepyrodia stricta . 30

Lepyrodia tasmanica . 31

Loxocarya . 29

Loxocarya fasciculata . 26

Loxocarya flexuosa . 26

Loxocarya myrioclada . 27

Loxocarya striata . 29

Loxocarya virgata . 27

Meeboldina . 29

Meeboldina cana . 29

Meeboldina coangustata . 30

Meeboldina crassipes . 30

Meeboldina scariosa . 30

Megalotheca striata . 29

Restio abortivus . 24

Restio amblycoleus . 24

Restio australis . 23

Restio chaunocoleus . 24

Restio confertospicatus . 28

Restio crispatus . 24

Restio diffusus . 29

Restio fasciculatus . 26

Restio fimbriatus . 23

Restio flexuosus . 26

Restio gracilior . 25

Restio gracilis . 23

Restio hookeri . 22

Restio isomorphus . 25

Restio laxus . 25, 29

Restio leucoblepharus . 25

Restio longipes . 23

Restio megalotheca . 29

Restio ornatus . 25

Restio pallens . 23

Restio sphacelatus . 25

Restio stenocoleus . 23

Restio stenostachyus . 27

Restio tenellus . 24

Restio tenuiculmis . 23

Restio tetraphyllus . 23

Restio tetraphyllus

subsp. meiostachyus . 24

Schoenodum chilense . 22

Sporadanthus . 30

Sporadanthus caudatus . . 30

Sporadanthus gracilis. . 30

Sporadanthus interruptus . 30

Sporadanthus strictus. 30

Sporadanthus tasmanicus . 31

Two new rare species of Homoranthus
(Myrtaceae: Chamelaucieae) from the
Northern Tablelands of New South Wales

John T. Hunter

Abstract

Hunter, J.T. (149 Kentucky St, Armidale, NSW 2350, Australia) 1998. Tzvo new rare species of
Homoranthus (Myrtaceae: Chamelaucieae) from the Northern Tablelands of New South Wales. Telopea
8(1): 35-40. Two new species of Homoranthus (H. bornhardtiensis J.T.Hunter and H. croftianus
J.T.Hunter) from granitic outcrops on the Northern Tablelands of New South Wales are described
with assessments of their conservation status and abundance.

Introduction

During the last decade, several species of Homoranthus have been described in New
South Wales and Queensland (Byrnes 1981, Craven & Jones 1991). Many of these
species have come from the granitic outcrops that are common on the New England
Batholith. With the intensity of flora survey work being carried out in this region a
number of new distributional records and a potential new taxon of Homoranthus have
also been discovered (Hunter 1997, Hunter & Bruhl 1997, Clarke et al. 1998). During
recent surveys carried out as an assessment of properties offered to the New South
Wales National Parks and Wildlife Service two new Homoranthus taxa were discovered
growing on granitic outcrops. These are here described.

Key to species

The key to the species of Homoranthus presented by Craven & Jones (1991) should be
amended as follows.

1. At least some sepal margins laciniate (the divisions 1.5 mm or more long) or long

acuminately bifurcate

2. Flowering branchlets condensed into a corymbose head

3. Hypanthium 2-3 mm long, style 4-7 mm long . H. wilhelmii

3. Hypanthium 3.5-4.5 mm long, style 8-10 mm long . H. tropicus

2. Flowering branchlets not condensed into a corymbose head

4. Leaves 3-4.5 mm long (sepals with 6-10 laciniae) . H. darwinioides

4. Leaves 2.5-17 mm long (if less than 6 mm long then sepals with fewer than 5
laciniae

5. Leaf blade in transverse section sublinear . H. porteri

5. Leaf blade in transverse section subquadrate, suborbicular, subobtriangular
or subelliptic

Telopea 8(1): 1998

6. Sepals with 6-12 laciniae . H. cemuus

6. At least some sepals with 2-5 laciniae

7. Style 15-21 mm long, leaves 4.5-6 mm long, hypanthium 6-8 mm
long . H. decasetus

7. Style 1.5-10 mm long, leaves 2.5-11 mm long, hypanthium 2-6 mm
long

8. Style 6-10 mm long, leaves 7-11 mm long, hypanthium glabrous,
5-6 mm long . H. biflorus

8. Style 1.5-2.5 mm long, leaves 2.5-8 mm long, hypanthium with
trichomes, 2-4 mm long . H. croftianus

1. Sepal margin distally entire or erose to lacerate (if lacerate the divisions less than 0.5
mm long)

9. Leaves papillate. H. papillatus

9. Leaves glabrous

10. Leaves 3-5 mm thick (in side view axe-shaped)

11. Sepals 5-6.5 mm long, style 22-28 mm long. H. zeteticorum

11. Sepals 3^1 mm long, style 15-19 mm long. H. thomasii

10. Leaves 0.3-1.5 mm thick

12. Peduncles 9-16 mm long. H.floydii

12. Peduncles 0.7-3.5 mm long

13. Style 18-25 mm long. H. homoranthoides

13. Style 3.5-11 mm long

14. Sepals with the apex obtuse . H. decumbens

14. Sepals with the apex long acuminate

15. Leaf blade in side view incurved linear

16. Sepals less than 3 mm long

17. Leaves 3.5-7 mm long, the blade in transverse section
suboblong to subquadrate to subobtriangular; sepals
1-1.5 mm long. H. melanostictus

17. Leaves 7-22 mm long, the blade in transverse section
suborbicular to subelliptic to suboblong; sepals 1.5-3
mm long

16. Sepals 3.5-4 mm long . H. montanus

15. Leaf blade in side view lunate, scimitar-shaped, narrowly
obovate, to incurved linear-obovate

18. Hypanthium with multicellular trichomes between costae

19. Ovules 4-6 . H. virgatus

19. Ovules 8-10 . H. prolixus

18. Hypanthium glabrous

Hunter, New species of Homoranthus from NSW

20. Hypanthium smooth; style 6-9 mm long .... H. lurmtus

20. Hypanthium ruminate; style 3.5-5 mm long .

. H. bornlwrdtiensis

I. Homoranthus bornhardtiensis J.T.Hunter, sp. nov.

H. prolixo Craven & S.R.Jones affinis sed hypanthio ruminato brevique differt.

Holotype: New South Wales: Northern Tablelands: Bald Rock, 'Bornhardtia', adjacent
Ironbark Nature Reserve (30°21'58" 150°54'27"), J.T. Hunter 5550 & V.H. Hunter, 9 Nov
1997 (NSW). Isotypes: BRI, CANB, NE.

Shrub to 0.3 m tall, prostrate, spreading to 1 m diameter. Stems yellow to reddish when
young, turning grey-brown. Leaves opposite, decussate, 4.5-9 mm long, 0.6-0.9 mm
wide and thick, aromatic; blade incurved or rarely straight, linear to oblanceolate, in
transverse section subobtriangular, abaxially flat or concave, pale green to glaucous,
reddish to purplish in spring; apex curved, mucronate to apiculate; petiole 0.1-0.5 mm
long. Floivers solitary in leaf axils, 1-4 on undifferentiated branchlets, yellow. Peduncles
0.7-1.4 mm long, crowned between bracteoles; bracteoles caducous, 3.5-4.5 mm long,
red-brown. Hypanthium 5-costate, ruminate between the rounded costae on the ovary
region, 1.6-2.5 mm long, 1.1-1.7 mm wide. Sepals 1.2-2 mm long, 0.05-0.1 mm wide,
apex long and hair-like, the margins entire, gland-dotted at the base. Petals orbicular,
0.7-0.9 mm long, 0.8-1 mm wide, the margin entire. Stamens 10; filaments 0.2-0.4 mm
long; anthers 0.2-0.4 mm long; staminodes 20, free, 0.2-0.4 mm long. Style glabrous,
3.5-5 mm long; stigma capitate. Ovules 10. Fruit simple, dry indehiscent nut, 2.5-3.5 mm
long, 1.1-1.7 mm wide, caducous after seed set, orange brown. Flowering October to
November (Fig. lc).

Additional specimens examined: New South Wales: Northern Tablelands: Bald Rock,
'Bornhardtia', adjacent Ironbark Nature Reserve, 30°21’ 150°54’, Hunter 5551, Sheringham, & Davies,
1 July 1997 (NE); North Rock, 'Bornhardtia', adjacent Ironbark Nature Reserve, 30°20’ 150°54',

J. T. Hunter 5590 & V.H. Hunter, 2 Feb 1998 (NE).

Notes: this species was first recorded in Ironbark Nature Reserve by Greg Roberts
(Ranger, NPWS Armidale) soon after its dedication in 1985 but was considered to be
Homoranthus flavescens. As no collections were made at that time Craven & Jones (1991)
were unaware of it during their revision of the genus. This taxon was collected again
during an assessment of a property neighbouring Ironbark Nature Reserve that was
on offer to the New South Wales National Parks and Wildlife Service. This property
has since gone to public sale. This species can be distinguished from the similar
H. prolixus by the shorter ruminate rather than hairy hypanthium and the prostrate but
not spreading habit. H. bornhardtiensis does not flower as prolifically as H. prolixus.
Craven & Jones (1991) use hairs on styles as a generic character however, no hairs were
noted on collections of this taxon. The lack of stylar hairs may have implications for
reproductive biology and conservation.

Etymology: a bornhardt is a dome shaped monolith that is bald and steep-sided.
'Bornhardtia' is also tire name of the property from which the type specimen was
collected.

Distribution: at present this taxon appears to be restricted to a few isolated granitic
outcrops within the Ironbark Nature Reserve and neighbouring properties. Despite a
survey of the granitic outcrop areas in the region, no further collections of this species
have been made. If no further collections are forthcoming, the present distribution of
this species is restricted to a six by seven kilometre patch.

Telopea 8(1): 1998

Fig. 1 . Homoranthus croftianus. a, unopened flower, note trichomes on hypanthium and 1-3-
laciniate sepals; b, style with hairy neck. Homoranthus bornhardtiensis. c, flower with bracteoles
removed, note undivided sepals and ruminate hypanthium. Scale bar: a, b = 1 mm; c = 1.5 mm.
Material illustrated is from the types. Illustration courtesy of Vanessa H. Hunter.

Hunter, New species of Homoranthus from NSW

Habitat: found in open and exposed situations in shrubland and low woodland on
granitic outcrops between 650-970 m altitude. The annual precipitation at the nearby
property 'Granite Heights' is 773 mm. The taxon is found rooted within crevices of
bare rocky slopes and in shallow soil surrounding the bare slopes. The pH of the soils
is between 4.5 and 4.75. Associated species include Callitris endlicheri, Eucalyptus prava,
E. caleyi, Acacia cheelii, A. neriifolia, Calytrix tetragona, Leucopogon neoanglicus,
Ozothamnus obcordatus, Leptospermum novae-angliae, Cryptandra amara var. floribunda,
Prostanthera howelliae and Boronia anethifolia.

Conservation status: Homoranthus bornhardtiensis has a very restricted occurrence on
only a few disjunct and isolated granitic outcrops within the Ironbark Nature Reserve
and surrounding properties. The geographic distribution of this species appears to be
only within a 7 km radius and the population may comprise fewer than 500 individual
plants. A preliminary ROTAP code (Briggs & Leigh 1996) of 2VC is suggested for this
taxon.

2. Homoranthus croftianus J.T.Hunter, sp. nov.

H. bifloro Craven & S.R.Jones similis sed hypanthium trichomatibus multicellularibus,
sepalis distaliter 1-3-laciniatis differt.

Holotype: New South Wales: Northern Tablelands: Angoperran Hill (29°2T 151°54'),
J.T.Hunter 5167, P. Sheringham & P. Croft, 15 June 1997 (NSW). Isotypes: BRI, CANB,
HO, MEL, NE.

Erect shrub to 2 m tall by 1.5 m wide. Stems yellow to orange when young, turning
brown with age. Leaves opposite, decussate, 2.5-8 mm long, 0.4—0.7 mm wide, 0.6-0.9 mm
thick; blade incurved or rarely straight or recurved, linear, in transverse section
subobtriangular, abaxially flat or concave, dark green to pale green; apex curved or
straight, acuminate to cuspidate; petiole 0.1-0.5 mm long. Blowers solitary in axils on
undifferentiated branchlets, greenish to cream. Peduncles 0.7-1.4 mm long, crowned
between bracteoles; bracteoles caducous, gland dotted, 3.5M.5 mm long, red-brown.
Hypanthium 5-costate, with multicellular trichomes between the rounded costae on the
ovary region, 1.5-3 mm long, 1-1.5 mm wide. Sepals 2-3 mm long, 0.3-0.6 mm wide,
margins variously divided on individual flowers from undivided to 3-laciniate, gland-
dotted. Petals orbicular to widely ovate, 1-1.2 mm long, 1-1.4 mm wide, margins
entire. Stamens 10; filaments 0.1-0.3 mm long; anthers 0.3-0.5 mm long; staminodes 20,
free, 0.3-0.5 mm long. Style 1.5-2.5 mm long, swollen basally, with long trichomes on
the top third; stigma papillate. Fruit simple, dry indehiscent nut, 3.5M mm long,

I. 2-1.5 mm wide, caducous after seed set, orange brown. Flowering sporadically June
to August (Fig. 1 a, b).

Notes: this species was first discovered during an assessment of a property on offer to
the New South Wales National Parks and Wildlife Service. Despite much interest in
Bolivia Hill and surrounds by botanists, this species appears to have been overlooked
and not collected. The species can be distinguished by its erect habit, distinctive green
foliage, long, multicellular trichomes on the hypanthium and variable 1-3-laciniate sepals.

Etymology: named in honour of Peter Croft, a senior ranger at the Glen Innes District
National Parks and Wildlife Service. Peter has shown great enthusiasm for
understanding the plants and animals around any area he has worked and was
present when the type specimen was collected.

Distribution: at present, this taxon is restricted to the Bolivia Range, particularly
Angoperran Hill. This species is highly restricted and despite numerous searches of
surrounding areas, no further populations have been found.

Telopea 8(1): 1998

Habitat: found in open and exposed situations in shrubland and low woodland on
granite outcrops at around 1000 m altitude. Tire annual precipitation at Bolivia Hill is
around 830 nun. The taxon is found rooted within crevices of bare rocky slopes and in
the shallow soil surrounding the bare slopes. The pH of the soils is between 3.8 and
4.8. Associated species include Callitris endlicheri, Eucalyptus prava, £. dealbala,
Leucopogon neoanglicus, Micromyrtus sessilis, Kunzea bracteolata, Boronia anethifolia,
Leptospermum novae-angliae, Acacia viscidida, A. pycnostachya and Cryptandra lanosiflora.

Conservation status: Homoranthus croftianus has a very restricted occurrence on large
exposed granitic outcrops on and around Bolivia Hill. The geographic range at present
is only a few kilometres radius and the population is probably less than 500
individuals. At present the population is not represented in any conservation reserve
and a preliminary ROTAP code (Briggs & Leigh 1996) of 2V is suggested for this taxon.

Acknowledgments

I thank Paul Sheringham, Paul Davies, Peter Croft and Vanessa Hunter for various
assistance in the field. Barbara Bi'iggs and Jeremy Bruhl are thanked for allowing
access in the past to NSW and NE for examination of Homoranthus specimens. John
Williams is thanked for discussion of Homoranthus taxa.

References

Briggs, J.D. & Leigh, J.H. (1996) Rare or Threatened Australian Plants, revised edition. (CSIRO:
Collingwood).

Byrnes, N.B. (1981) Notes on the genus Homoranthus (Myrtaceae) in Australia. Austrobaileya 1:
372-275.

Craven, L.A. & Jones, S.R. (1991) A taxonomic review of Homoranthus and two new species of
Darwinia (both Myrtaceae, Chamelaucieae). Australian Systematic Botany 4: 513-533.

Clarke, P.J., Copeland, L.M., Hunter, J.T., Nano, C.E., Williams, J.B. & Wills, K.E. (1998) The
Vegetation and Plant Species of Torrington State Recreation Area. (Division of Botany, University
of New England: Armidale).

Hunter, J.T. (1997) A further record of Homoranthus lunatus Craven & S.R.Jones (Myrtaceae) for
northern New South Wales. Queensland Naturalist 35: 24-25.

Hunter, J.T. & Bruhl, J.J. (1997) Significant range extensions for 10 species of vascular plants in
northern New South Wales. Austrobaileya 4: 691-694.

Manuscript received 31 March 1998
Manuscript accepted 30 July 1998

Nassella tenuissima (Gramineae) recorded from
Australia, a potential new weed related to
Serrated Tussock

S.W.L. Jacobs, J. Everett and Marfa Amelia Torres

Abstract

Jacobs, S.W.L. 1 , Everett, J.' and Torres, Marta Amelia 2 0National Herbarium ofNeiv South Wales, Royal
Botanic Gardens, Sydney NSW 2000, Australia; 2 Departamento Cienttfico de Plantas Vasculares, Museo
de La Plata, Paseo del Bosque s/No., 1900 La Plata, Argentina) 1998. Nassella tenuissima (Gramineae)
recorded from Australia, a potential neiu weed related to Serrated Tussock. Telopea 8(1): 41-46. Nassella
tenuissima is recorded from the horticultural trade in New South Wales. It is very similar to
N. trichotoma (Serrated Tussock) and difficult to distinguish from that species. It is possible that
N. tenuissima has already become naturalised in Australia although not yet recorded. A description
is provided of N. tenuissima and the distinguishing features of the two species are Listed. The history
of establishment of N. tenuissima in New Zealand is reviewed; comments are made on i ts culti vation
in California; its potential weediness in Australia is assessed, and recommendations are made.

Introduction

Through the alertness of Ken Folkes, Weeds Officer for the Snowy River Shire Council,
a cultivated specimen he thought suspiciously similar to Serrated Tussock ( Nassella
trichotoma (Nees) Hack, ex Arechav.) was forwarded to Jim Dellow of NSW Agriculture
at Orange. The specimen was grown to flowering there in a glasshouse and forwarded
for our comments.

The specimen was then sent to Maria Amelia Torres in Argentina who identified it as
Nassella tenuissima (Trin.) Barkworth.

Nassella tenuissima is native to New Mexico, Texas, Mexico, Argentina and Chile, has
been cultivated in California and New Zealand, and also has been recorded from
South Africa (Connor & Ford 1996) where it is apparently only known from one
locality (N. Barker, pers. comm.).

Nassella tenuissima is closely related to N. trichotoma (Serrated Tussock), a serious
pasture and environmental weed in temperate Australia, and looks so similar,
especially when not flowering, that it could be mistaken in the field for N. trichotoma.
We thought it advisable to assess the potential seriousness of N. tenuissima as a weed
in Australia and to provide the information required for its detection and the
documentation of its behaviour.

Nassella tenuissima

The species can only be distinguished from Serrated Tussock when flowering.
N. tenuissima differs from Serrated Tussock in:

(i) its inflorescence (seedhead) not always being fully exserted and spread from the
sheath at maturity;

Telopea 8(1): 1998

(ii) the seedhead apparently not detaching from the uppermost culm node at maturity
(these characters require more observations over a range of conditions);

(iii) awns 5-9 cm long (2-3.5 cm in Serrated Tussock);

(iv) the lower glume slightly longer (9-10.5 mm vs 6-8.5 mm); (v) the body of the
lemma slightly longer (to 3 mm vs 2 mm);

(vi) and the floret slightly less gibbous (i.e., less asymmetrical).

The key to species of Nassella in Jacobs and Everett (1993) can be modified to produce
a key to all the species of Nassella recorded from Australia:

1 Summit of the lemma without a cylindrical corona; floret <3.5 mm long . 2

1* Summit of the lemma forming a conspicuous cylindrical corona 0.5-2.5 mm long
around the base of the awn; floret >3.5 mm long. 3

2 Awn 2-3.5 cm long; floret 1.5-2 mm long; inflorescence fully exserted and detaching

at maturity. N. trichotoma

2* Awn 4.5-9 cm long; floret 2-3 mm long; inflorescence often only partly exserted from
sheath, detaching or not detaching at maturity . N. tenuissima

3 Floret c. 4 mm long; awn 3.5-4 cm long, column scabrous with hairs <0.05 mm long

. N. hyalina

3* Floret 5.5-10 mm long; awn 4.5-8.5 cm long, column scabrous or pubescent with hairs

usually c. 0.2 mm long .'.. 4

4 Corona <1.5 mm long . 5

4* Corona >1.5 mm long . 6

5 Lower glume 8-10 mm long; floret 5.5-6 mm long; coronal spines c. 0.1 mm long

. N. megapotamia

5* Lower glume 14-18 mm long; floret 6-10 mm long; coronal spines c. 0.5 mm long

. N. neesiana

6 Corona 1.5-2.5 mm long, with a firm basal part 0.5-1 mm long, the upper

membranous part with hairs 1-1.5 mm long . N. leucotricha

6* Corona >3 mm long, with a firm basal part c. 3 mm long, the upper 0.5-1.9 mm
lacerate into narrow fingers, these in turn lacerate at the apex into fine hairs 0.5 mm
l° n 8. N. charruana

*Nassella tenuissima (Trin.) Barkworth (1990: 612); Torres (1997: 37).

Basionym: Stipa tenuissima Trinius (1836: 67); Caro (1966: 62-65).

Holotype: Argentina: Mendoza, Gillies (LE).

Densely caespitose perennial with very numerous, often sterile, intravaginal
innovations and wiry roots, the tussocks to 70 cm tall. Culms 2-4-noded, the nodes not
thickened; internodes terete, smooth, glabrous. Sheaths tight around the culms or
open towards the base of the plant, lightly striate, glabrous and smooth except for the
more or less scaberulous margins. Ligule opaque, chartaceous, obtuse or truncate,
0.5-2.5 mm long, usually glabrous and smooth, symmetrical or asymmetrical. Blades
straight or curving, convolute-filiform, setaceous, to c. 60 cm long, 0.25-0.5 mm diam..

Jacobs, Everett and Torres, Nassella tenuissima

the outer surface minutely and distantly scaberulous, not striate. Inflorescence
contracted and at first more or less included in the upper sheath', with few spikelets
towards tire tips of the ultimate branches, 15-25 cm long, green or purplish from the
colour of the glumes, the main axis somewhat compressed and plano-convex,
antrorsely scabrous on the edges; branches filiform, 2-3 together, scabrous-pubescent,
the lower up to 10 cm long, unequally branched, with spikelets borne on the ultimate
branchlets on long capillary pedicels. Spikelets narrowly lanceolate, 5-10 mm long
(excluding awn). Glumes much longer than the floret, unequal, narrowly lanceolate,
long-acuminate with a hair-like tip, 3-nerved with the lateral nerves shorter and
converging towards the median nerve, membranous and often purple, the apex thinly
hyaline; lower glume 8-10 mm long, upper glume 5-7 mm long. Floret ellipsoid to
obovate, gibbous, slightly compressed, without a corona, 2-3 mm long (excluding
awn), c. 1 mm wide. Lemma obscurely 5-nerved, antrorsely scabrous upwards, straw-
coloured, green or purplish, with an excentric filiform awn, the margins overlapping
and tightly enclosing the palea; awn more or less straight or flexuose, 5-9 cm long,
slightly twisted in the lower part, minutely scabrous, inconspicuously articulate with
the lemma, rather tardily deciduous; callus shortly oblique, obtuse or minutely
pointed, with white, erect, divergent hairs to 1.5 mm long. Palea thinly hyaline,
glabrous, nerveless, 30-50% as long as the lemma. Lodicules 2, thinly hyaline,
glabrous, nerveless, 0.25-0.75 mm long. Stamens 3, sometimes all c. 1 mm long and
mostly all exserted though sometimes not exserted; sometimes with a single anther
0.6-1.5 mm long and 2 c. 0.4 mm long or less, either all exserted, or sometimes the
2 smaller (?and sometimes the 3rd) not exserted. Caryopsis ellipsoid, slightly
compressed, 2-2.5 mm long; hilum linear, straight, inconspicuous, c. 60-70% of grain
length; embryo c. 50% grain length.

* Many species of the Stipeae are facultatively cleistogamous (Vickery et al. 1986) and
the behaviour of the florets can be influenced by prevailing conditions. There is
evidence of both cleistogamy and chasmogamy in N. tenuissima (Connor & Ford 1996,
1997). Both types of flower are in the same inflorescence. Cleistogamous flowers are
more precocious than the chasmogamous flowers and mature earlier. The only
specimen from Australia has partly exserted inflorescences, as seems to be the case in
New Zealand specimens, whereas there are reports of more exserted inflorescences in
Argentina (Connor & Ford 1996, 1997) though the lower part of the inflorescence is
always recorded as enclosed by the sheath. It is not yet clear whether any of these
inflorescences becomes detached before the spikelets disarticulate.

Natural distribution in Argentina

In Argentina it grows in the Central, Western, Southern and Northwestern regions,
extending from about 26°50'S to 47°34'S, and from near sea level to about 2900 m
altitude. Typical habitats include some zones of the Tampeana’ grass steppe,
Patagonian grass and shrub steppe, ground cover in various xeromorphic and
semiarid woodlands (e.g. the Calden woodlands of Prosopis caldenia Burk.), and
highland and, sometimes, alpine meadows. It is a species with a wide climatic and
habitat range.

Natural distribution and cultivation in USA

In Texas, N. tenuissima is reported (Hitchcock & Chase 1951; Correll & Johnston 1970;
Martin & Hutchins 1980; Connor & Ford 1996,1997) from open rocky flats and slopes,
on mountains, plateaux and arid valleys with an average rainfall of 300 mm, with soils
and sites of extreme variability. In the southern United States it grows over an altitude
range of 600-2350 m (M.E. Barkworth pers. com.). In Texas and New Mexico it is

Telopea 8(1): 1998

apparently eaten by cattle and goats but, in California, has proved difficult to eradicate
after spreading from cultivation (M.E. Barkworth pers. com.). It should be noted that
N. trichotoma is reportedly eaten by stock in South America (Vickery & Jacobs 1980).

History in New Zealand

It is fortunate that this is one of those rare occasions where the establishment of a
potential weed species has been well documented. Ford (1994) documented the first
recording of its spread from cultivation in New Zealand in a commercial nursery and
noted that it was not included in the treatment of stipoid grasses in New Zealand
(Jacobs et al. 1989). Gardner et al. (1996) supplied the first published identification,
reported a second locality of naturalisation and predicted that it would soon be
widespread. Connor and Ford (1996, 1997) reviewed the situation, including recent
changes in the classification and nomenclature in the stipoid grasses and pleaded for
vigilance in recording all introduced stipoid species. De Lange and Gardner (1997)
despairingly documented the continued spread of the species, both from known sites
and from previously unrecorded sites, and the apparent lack of action by both
responsible authorities and the nursery industry.

It appears that N. tenuissima was originally imported as Stipa tenuissima (a synonym of
N. tenuissima ) in mistake for the long-cultivated Stipa tenacissima (now Achnatherum
tenacissinm Barkworth; Barkworth 1993; Jacobs & Everett 1996), Esparto Grass.

History in Australia

The origin of the Australian material is still not entirely clear but the letter from Ken
Folkes accompanying the first specimen noted:

'Recently I received an ornamental grass which had been purchased several years ago
by a friend, from a Sydney nursery.'

The letter continues:

'This plant is marketed as a native grass, and was called "Elegant Spear Grass".'

Elegant Spear Grass, Austrostipa elegantissima (Labill.) Jacobs and Everett, is indeed
native and, as the name implies, is also a very attractive horticultural subject. It bears
no similarity to the recent introduction though it is in a related genus. It appears that
N. tenuissima was (?is) marketed under a wrong and misleading name, an action that
could prove to have dire consequences.

It seems most likely that the Australian material was imported from either California
or New Zealand, with perhaps the latter being more likely. It is possible that this
importation was made before the potential problems had been identified in New
Zealand and California. The fact that N. tenuissima was apparently sold in Australia
incorrectly labelled with the common name of an Australian native species may
indicate that this was not the case.

Although not recorded as naturalised within Australia, it is indistinguishable
vegetatively from Serrated Tussock and will only be recognised as distinct when
flowering and when observers have been made aware of the differences between the
two species.

While it is probably important to try and identify how N. tenuissima was introduced to
prevent a similar event being repeated, it is more important to document whether this
species has already become naturalised in Australia and to try to eradicate any such
plants, and to remove plants from propagation, sale and cultivation.

Jacobs, Everett and Torres, Nassella tenuissima

Assessment of potential weediness

In its natural habitat, N. tenuissima grows from about 20° to 35° North, and from 25° to
47° South, in areas with an average annual rainfall of 300 mm, and on extremely
variable soils. Reports from Wellington, New Zealand, suggest that the climate there
is too wet for it. Ford (1994) suggesting an upper rainfall limit of <1250 mm. Even a
very rough transfer of this information on to a map of Australia suggests that, all else
being equal (which it never is), N. tenuissima could become established over about 65%
of New South Wales (excluding the coastal strip and the northwest corner), 70% of
Victoria (excluding the higher rainfall areas), the southern 20% of South Australia, 20%
of Tasmania, up to 20% of Western Australia (the southwest corner), and possibly
some of the higher country or even elsewhere in southern Queensland. Whether it
becomes a serious weed in all or any of these areas will depend on various as yet
unknown biological characteristics and the particular land management regimes.
Although it is reportedly eaten by stock in its native range (M.E. Barkworth pers.
comm.), so is Serrated Tussock in its native range (Vickery & Jacobs 1980). The story of
Eragrostis curvula (African Love Grass) is also pertinent (Jacobs 1982) where
palatability varies within a species and can be dependent on the alternatives available.
The story in southern Africa for £. curvula was very different from that observed here.

Some consolation may also be taken by the apparent lack of long distance wind
dispersal in N. tenuissima but, as noted above, this is not definitely proved, and de
Lange and Gardner (1997) document establishment at sites in New Zealand
apparently well away from known plantings.

The information available suggests that N. tenuissima is potentially a serious weed. We
recommend that:

(i) N. tenuissima be banned from propagation and sale, and that nursery stocks be
destroyed;

(ii) N. tenuissima be removed where cultivated in public areas;

(iii) private land owners be encouraged to remove N. tenuissima from gardens;

(iv) experienced Weeds Officers be taught the differences between N. tenuissima and N.
trichotoma and asked to look for N. tenuissima as opportunities arise; and

(v) the situation be monitored and assessed as data become available.

Acknowledgments

We thank Ewen Cameron for providing us with relevant references from New
Zealand, Mary Barkworth for information about the species in North America, Nigel
Barker for information about the species in South Africa, and Jim Dellow for providing
the first good specimen.

References

Barkworth, M.E. (1990) Nassella (Gramineae, Stipeae): revised interpretation and nomenclatural
changes. Taxon 39: 597-614.

Barkworth, M.E. (1993) North American Stipeae (Gramineae) taxonomic changes and other
comments. Phytologia 74:1-25.

Caro, J.A. (1966) Las especies de Stipa (Gramineae) de la region Central Argentina. Kurtziana 3:
7-119.

Telopea 8(1): 1998

Connor, H.E. & Ford, K. (1996) Stipa tenuissima, and name changes in stipoid grasses. Weed
Identification Nezvs No. 20:1-5.

Connor, H.E. & Ford, K. (1997) Stipa tenuissima, and name changes in stipoid grasses. New Zealand
Botanical Society Newsletter 47:10-13.

Correll, D.S. & Johnston, M.C. (1970) Manual of the Vascular Plants of Texas. (Texas Research
Foundation: Renner).

de Lange, RJ. & Gardner, R.O. (1997) Stipa tenuissima: unwanted weedy grass on the move.
Auckland Botanical Society journal 52: 25-26.

Ford, K. (1994) Stipa sp., Nassella Tussock relation: "a potential weed!" Weed Identification Nezvs No.
13: 3-5.

Gardner, R.O., Champion, P.D. & Lange, P.J. de (1996) Andropogon virginicus and Stipa tenuissima.
Auckland Botanical Society journal 51: 31-33.

Hitchcock, A.S. & Chase, A. (1951) Manual of the grasses of the United States. United States
Department of Agriculture Miscellaneous Publication no. 200. (Govt. Printing Office:
Washington)

Jacobs, S.W.L. (1982) Classification of the Eragrostis curvula complex in Australia. Australian Plant
Introduction Reviexv 15: 5-14.

Jacobs, S.W.L. & Everett, J. (1993) Nassella. Pp. 638-639 in Harden, G.J. (ed.), Flora of New South Wales,
vol. 4. (New South Wales University Press: Sydney)

Jacobs, S.W.L. & Everett, J. (1996) Auslrostipa, a new genus, and new names for Australian species
formerly included in Stipa (Gramineae). Telopea 6: 579-595.

Jacobs, S.W.L., Everett, J., Connor, H.E., & Edgar, E. (1989) Stipoid grasses in New Zealand. New
Zealand journal of Botany 27: 569-582.

Martin, W.C. & Hutchins, C.R. (1980) A Flora ofNeiv Mexico, vol. 1. (J. Cramer: Vaduz).

Torres, M. A. (1997) Nassella (Gramineae) del noroeste de la Argentina. Monogr. Comis. Invest. Ci.
Prov. Buenos Aires 13: 5-45.

Vickery, J.W. & Jacobs, S.W.L. (1980) Nassella and Oryzopsis in New South Wales. Telopea 2: 17-23.
Vickery, J.W., Jacobs, S.W.L. & Everett, J. (1986) Taxonomic studies in Stipa (Poaceae). Telopea 3:
1-132.

Manuscript received 11 June 1998
Manuscript accepted 30 July 1998

Notes on Australian Meliaceae

D.J. Mabberley

Abstract

Mabberley, D.J., (Rijksuniversiteit Leiden, Netherlands and Royal Botanic Gardens Sydney, Mrs Macquaries
Road, Sydney NSW 2000, Australia) 1998. Notes on Australian Meliaceae. Telopea 8(1): 47-48.
Dysoxylum amooroides Miq. is lectotypified and Turraea decandra Blanco neotypified; Owenia capitis-
yorkii Domin is reduced to O. vernicosa; Synoum muelleri is recognised as a subspecies of
S. glandulosum — subsp. paniculosum (F. Muell.) Mabb., stat. nov.

Introduction

The following notes were intended as an Appendix to the Flora of Australia account of
Meliaceae, which was prepared in 1989, but has been unavoidably held up in
publication. As the findings are needed for other publications, they are presented here.

Dysoxylum Blume

Dysoxylum gaudichaudianum (A. Juss.) Miq.

Turraea decandra Blanco, FI. Filip.: 347 (1837).

Type (designated here): Philippines: Luzon, Manila, E D. Merrill Sp. Blancoanae 218;
neo: L; isoneo: A, BM, BO, G, K, NSW, P, PNH lost.

D. decandrum (Blanco) Merr., Philip. Gov. Lab. Publ. 27: 30 (1905).

D. amooroides Miq., Ann. Mus. Bot. Lugd.-Bat. 4:16 (1868).

Type (selected here):[Indonesia: south-west Irian Jaya,] A. Zippelius s.n .; lecto:
U; isolecto: L.

D. gaudichaudianum has been widely known in Australia as D. decandrum (Blanco)
Merr. or D. amooroides Miq., neither of which names has been precisely typified. As
Blanco's specimens are lost, Merrill collected a new series and his 'illustrative'
specimen collected near the type site of T. decandra is an appropriate choice for a
neotype, even though the PNH specimen itself is lost, as it is so well-represented as
duplicates in other herbaria. Of the syntype sheets quoted by Miquel in his description
of D. amooroides, the Zippelius sheet is represented in Miquel's own herbarium and
duplicated at Leiden, making it a suitable lectotype.

Owenia F. Muell.

Owenia vernicosa F. Muell., Fragm. 3: 15 (1862).

O. capitis-yorkii Domin, Biblioth. Bot. 22 (89 4 ): 854 (1927).

Type: Queensland: Cape York, W. Hann, Cape York Expedition 115 holo (?): PR/PRC,
n.v.[see below]; iso (holo?): K; syn. nov.

Domin described O. capitis-yorkii from a single leaf and drupe without its epicarp. He
compared it with O. acidula F. Muell. and O. vernicosa F. Muell., from which it appeared

Telopea 8 ( 1 ): 1998

to differ in leaflet characters. With more collections of O. vemicosa, the scrappy type
falls within the range exhibited by that species. According to the curators, there is no
extant material at PR or PRC and the fact that Domin mentions specifically the scraps
which match the Kew sheet suggests that the Harm specimen there (Haim specimens
not known to be preserved elsewhere) is the basis for the description.

Synoum A. Juss.

Synoum glandulosum (Sm.) A. Juss.

subsp. paniculosum (E Muell.) Mabb., stat. nov.

S. glandulosum var. paniculosum F. Muell., Fragm. 5: 145 (1866, as ‘Synoon glandulosum
var. paniculosa')

Type: Queensland: Rockingham Bay, 1865, J. Dallachy s.nr, holo: MEL; iso: BM, BRI, L.
S. muelleri C.DC., Monog. Phan. 1: 593, t. 7 fig. 10 (1878)

Type: Queensland: Rockingham Bay, 'Ferd. Muell. in h. Berol.'; holo: B, lost; iso?:
( J. Dallachy s.n.): BM, BRI, L, MEL.

The northernmost populations of S. glandulosum have very long thyrses and
pentamerous flowers; Mueller gave them varietal status and, probably based on a
duplicate of the same gathering, C. de Candolle recognised them at species level as
S. muelleri. Occasionally pentamerous flowers and rather intermediate inflorescence
sizes are found in trees further south so that it would seem more appropriate to
recognize the northern populations as a geographical subspecies rather than a
separate species.

Manuscript received 19 March 1998
Manuscript accepted 30 July 1998

Revision of Livistona (Arecaceae) in Australia

A.N. Rodd

Abstract

Rodd, A.N. (Royal Botanic Gardens, Mrs Macquaries Road, Sydney, NSW 2000, Australia; Present
address: 225 Trafalgar Street, Annandale, NSW 2038, Australia) 1998. Revision of Livistona (Arecaceae)
in Australia. Telopea 8(1): 49-153. A taxonomic revision of the Australian species of the palm genus
Livistona is presented, with a key to the species. Sixteen species are recognised, five of them
described here as new (L. fulva, L. kimberleyarta, L. lanuginosa, L. nitida and L. victoriae). One
new subspecies is recognised (L. inariae subsp. occidentalis) and a new combination made at
subspecific rank (L. mariae subsp. rigida) for Livistona rigida Becc.

Introduction

For the greater part of the period in which the Australian flora has been the subject of
study by Western science, the native palms were neglected by botanists. Firstly, for a
long period there were no resident taxonomists with an interest in palms. F. von
Mueller and F.M. Bailey both described a number of palm species, but they dealt with
them in a somewhat superficial manner, placing excessive reliance on very inadequate
collections. Secondly, the sparsely scattered palms of Australia were not central to the
interests of palm specialists such as Wendland and Drude, Beccari, and Burret, and
they lacked adequate material for study. Thirdly, there is the combination of the
limited economic value of Australian palms and the great difficulty of collecting and
preparing specimens, with most botanical collectors unwilling to invest tire time and
energy required to make a satisfactory specimen. The latter two considerations have
had the result that in Australian herbaria there is in total a very small number of
adequate specimens of native palms, by comparison with almost any other plant
group of equivalent size and diversity.

In the last two decades, though, Australian palms have been the subject of more
intensive taxonomic study. Notable contributions have been the revisions of
Ptychosperma (Essig 1978), Wodyetia (Irvine 1983), Oraniopsis (Dransfield, Irvine & Uhl
1985), Archontophoenix (Dowe & Hodel 1994) and Linospadix (Dowe & Irvine 1997). In
addition, the results of Irvine's unpublished studies of Calamus have been included in
Jones's (1984) book on palms. This brings to a total of 24 the number of Australian
palm species which could be said to be adequately covered by revisions, out of an
estimated total of 53 species in 19 genera (not including endemics of offshore
possessions such as Lord Howe Island); the present study will add another 16 species.

Past accounts of Australian Livistona are very incomplete and frequently in
disagreement with one another. Wendland and Drude's monograph Palmae
Australasicae (1875) has only five species: one of these is now known to be a Licuala,
and another, Mueller's L. leichhardtii, an illegitimate name purporting to replace two
valid names of Brown; this left only Brown's L. australis, L. humilis and L. inermis. The
circumscription of the last of these names was greatly enlarged by Wendland and
Drude to include east-coast plants now known as L. drudei and L. decipiens.

Telopea 8 ( 1 ): 1998

Bentham's Flora Australiensis (1878) recognises only Brown's three species. Bentham
shows by his notes and synonymy that he was aware of the faults of Wendland and
Drude's treatment, but chose to adopt an extremely cautious and conservative species
concept, perhaps excusable considering the materials he had at hand.

Drude (1893) in a rather discursive account attempted to clarify the taxa known up to
that time, including Mueller's poorly defined L. drudei and L. alfredii. He attempted a
comprehensive key based largely on fruit, in which some of the 'species' appear to be
given a ranking subordinate to that of others. Drude made some perceptive
observations but failed to formalise his conclusions.

It was Beccari who made by far the most valuable contributions to knowledge of
Livistona, both in Australia and beyond it. The posthumous brief account (Beccari 1921
in Latin and Italian) of Old World Coryphoid palms followed by the very lengthy
(1931) account in English, translated and edited by MarteUi, increased the number of
Australian species to 11 (of which 10 are still recognised in this treatment) and placed
their morphological study on a much firmer footing. It is a testimony to Beccari's
appreciation of the nature of variation in palms that he was able to derive such a sound
taxonomy from the few and scrappy collections available to him.

Historical

Although Livistona was not recognised as a genus until so described by Robert Brown
in 1810, the earliest botanical description of a Livistona appears to be that of Rumphius
(1741-55), published in 1741 though prepared the previous century. Rumphius
illustrated a palm from the Moluccas under the mononomial Saribus. Linnaeus (1753)
included this as one element in his circumscription of Conjplw Umbraculifera; the other
major element was Rheede's (1682) Codda-pana, characterised by a massive terminal
inflorescence. Linnaeus's successors soon realised that his Corypha umbraculifera was a
heterogeneous concept, and Lamarck (1786) separated the Rumphian element under
the name Corypha rotundifolia. Moore and Dransfield (1979) formalised this effective
lectotypification of C. umbraculifera. C. rotundifolia Lam. appears to be the earliest
binomial applying unambiguously to a Livistona. It is currently known as L. rotundifolia
(Lam.) Bl. ' J

Loureiro (1790) described Corypha saribus from Cochin-China, though his adoption of
this epithet bespeaks some confusion with Rumphius's Saribus, which in fact he cited
as a synonym; but recent palm specialists, e.g. Moore (1963a) do not seem to regard
this as a bar to its adoption as an epithet under Livistona (as L. saribus (Lour) Merr &
A. Chev.). Jacquin (1801) followed with the publication of Latania chinensis, now
known as Livistona chinensis (Jacq.) R. Br. ex Mart.

Brown's (1810) protologue of the genus Livistona recognised two species, L. humilis and
L. inermis, from northern Australia, collected by himself during the Voyage of the
Investigator and both beautifully illustrated by Ferdinand Bauer (but the illustrations
not published until 1838, by Martius). Moore (1963b) has designated L. humilis as
lectotype of Livistona. Brown's diagnosis of the genus hardly distinguishes it from
Corypha, under which genus he placed a third Australian species now placed in
Livistona, viz. L. australis (R. Br.) Mart.

The genus name was, in Brown's words '...in memoriam viri nobilis Patricii Murray
Baronis de Livistone, ...', continuing in a laudatory vein to recount how Murray had
over a thousand [species of?] plants laid out in his garden. According to Baines (1981)
this was prior to 1680 and the garden subsequently became the Edinburgh Botanic
Garden. It is clear that Brown deliberately latinized the Murrays' family seat of Livingston

Rodd, Revision of Livistona (Arecaceae)

to Livistonis (ablative Livistone), from the inferred stem of which he derived the
substantive Livistona.

Since Brown's publication of the genus, 58 additional binomials and 8 trinomials have
been published in Livistona (Table 1).

The genus Livistona

A description of the genus is not given here, partly because of my lack of knowledge
of its variation beyond Australia, and partly due to the existence of a very complete
description in Uhl and Dransfield (1987). They treat it as one of 12 genera of subtribe
Livistoninae, which in turn is one of the 4 subtribes of the tribe Corypheae. The
Livistoninae are distinguished by their 3 carpels being free but the styles united; they
have a wide distribution through the tropics and warmer temperate regions of eastern
Asia, Australasia and the Americas, though with large disjunctions in the latter; one
outlying genus ( Pritchardia ) occurs on mid-Pacific archipelagoes, and one outlying
Livistona species in the Horn of Africa and southern Arabia. Both the distribution and
morphological features of Livistoninae suggest they are one of the most ancient
groups of palms.

Livistona has arguably the largest number of unspecialised features of any genus in the
Livistoninae, or possibly in the subfamily Coryphoideae. Its closest ally appears to be
the tropical south-east Asian Pholidocarpus, distinguished by its compound leaf-
segments, broader-based carpels and very large, usually corky fruit, among other
features. However, some of the American genera such as Acoelorrhaphe, Brahea and
Copernicia appear to be almost as closely allied, with leaves almost identical in
structure to those of Livistona and flowers and inflorescence showing many
similarities.

Of the 30-odd Livistona species, 14 are Australian endemics, 2 extend from north¬
eastern Australia to lowland New Guinea, and the remainder are extra-Australian.
The northwestern limit of the genus in Asia appears to be Assam (L. jenkinsiana) and
the northeastern limit the southern islands of Japan (possibly as far north as Kyushu:
L. chinensis). It extends eastward in the Pacific only as far as the Solomon Islands
(L. woodfordii). It is doubtful that any major island of the Malay Archipelago has more
than 3 or possibly 4 native species, with the possible exception of New Guinea, which
has 4 named species (Johns & Hay 1984) and probably others as yet unnamed; but
there appears to be no species common to Papuasia and any other part of Malesia. Two
widespread and variable species, L. saribus and L. rotundifolia, account for most
occurrences of the genus in mainland Asia, the Sunda Shelf and the Philippines
(see Beccari 1921, 1935; Burret 1941; Whitmore 1973). Borneo, despite its size and
floristic riches, appears to have only one endemic species, the remarkable miniature
L. exigua. After Australia the next major region of diversity for the genus, if unnamed
taxa are taken into account, appears to be the lulls and mountains of mainland south¬
east Asia and the Malay Peninsula: according to Dransfield (personal communication
1998): 'Peninsular Malaysia has L. saribus, L. endauensis, L. tahanensis, L. speciosa and an
undescribed taxon that also occurs in Thailand and Lao. If we add in Thailand, then
there is at least one more taxon, and going further north we can add two or three more
in Vietnam and S. China.' The most remarkably disjunct occurrence of the genus has
only recently been demonstrated, with Dransfield and Uhl's (1983) transfer of the sole
species of Wissmannia Chiov. to its rightful place in Livistona, as L. carinensis. This
species occurs in coastal Djibouti and Somalia in the Horn of Africa, also in a few
sheltered valleys of the Hadramaut in South Yemen.

Telopea 8(1): 1998

Table 1. All known Livistona names, listed in chronological order. Those whose types are non-
Australian, or of unknown origin, are indicated by an asterisk.

Names that appear to be in current use are shown in bold type. Names now regarded as synonyms are
linked by the '=' sign to the name of the taxon to which they are believed to refer. This list does not
include orthographic variants (e.g. jenkinsii / jenkinsiana) where they are clearly references to the same species.

Livistona humilis R. Br. 1810
Livistona inermis R. Br. 1810
L. chinensis* (Jacq.) R. Br. ex Mart. 1838
[Latania chinensis Jacq. 1801]

L. mauritiana* Wall, ex Mart. 1838
L. rotundifolia * (Lam.) Mart. 1838
[Corypha rotundifolia Lam. 1786]

L. australis (R. Br.) Mart. 1838
[Corypha australis R. Br. 1810]

L. bissula* Mart. 1838
= Licuala sp.

L. gaudichaudii* Mart. 1838 = Pritchardia
gaudichaudii (Mart.) H. Wendl.

L. martii* Gaudich. 1842 = Pritchardia martii
(Gaudich.) H. Wendl.

L. jenkinsiana* W. Griff. 1844
L. spectabilis* W. Griff. 1844 = ?L hasseltii
L. subglobosa* (Hassk.) Mart. 1850
[Saribus subglobosus Hassk. 1842]

L cochinchinensis* (Bl.) Mart. 1850

[Saribus cochinchinensis Bl. 1836] = L. saribus
L oliviformis* (Hassk.) Mart. 1850

[Saribus oliviformis Hassk. 1842] = L. chinensis
L. altissima* Zoll. 1857 = L. rotundifolia
L. diepenhorstii* Hassk. 1858

= Pholidocarpus diepenhorstii (Hassk.) Burret
L. hasseltii* (Hassk.) Hassk. ex Miq. 1868
[Saribus hasseltii Hassk. 1842]

L. hoogendorpii* Teysm. & Binn. 1868
= ?L. hasseltii

L. leichhardtii F. Muell. 1874 = L. humilis
L. ramsayi F. Muell. 1874 = Licuala ramsayi
(F. Muell.) Domin
L. speciosa* Kurz 1874
L. papuana* Becc. 1877
L. mariae F Muell. 1878
L. filifera H. Wendl. 1878 (nom. nud.?)

L. moluccana* H. Wendl. 1878 = ?

L. occidentalis* Hook. f. 1887 (nom. nud.?)

= IBrahea dulcis [Becc. 1921]

L. kingiana* Becc. 1889 = Pholidocarpus kingianus
(Becc.) Ridley
L. enervis* Hort. 1891 = ?

L. alfredii F Muell. 1892
L. woodfordii* Ridley 1898
L. muelleri Bailey 1902
L. benthamii Bailey 1902
L. rupicola* Ridley 1903 = Maxburretia
rupicola (Ridley) Furtado

L. merrillii* Becc. 1904

L. vidalii* Becc. 1905 = Corypha utan Lam.

L microcarpa* Becc. 1907 = L rotundifolia
var. microcarpa

L. whitfordii* Becc. 1909 = ?L merrillii
L. mindorensis* Becc. 1909 = L. rotundifolia
var. mindorensis

L. inaequisecta* Becc. 1909 = ?L hasseltii

L. decipiens Becc. 1910
L. robinsoniana* Becc. 1911
L. chinensis var. boninensis* Becc. 1912
L. blancoi* Merr. 1918 = ?L rotundifolia
L. rotundifolia var. microcarpa* (Becc.) Becc.

1919 [L microcarpa Becc. 1907]

L. rotundifolia var. mindorensis* (Becc.)

1919 [L mindorensis Becc. 1909]

L. rotundifolia var. luzonensis* Becc. 1919
L saribus* (Lour.) Merr. ex A. Chev. 1919
[Corypha saribus Lour. 1790]

L. chinensis var. subglobosa* (Hassk.) Becc.

1921 [Saribus subglobosus Hassk. 1842]

L. tahanensis* Becc. 1921
L. decipiens var. polyantha Becc. 1921
= L. decipiens

L. holtzei Becc. 1921 = L. benthamii
L. lorophylla ('loriphylla') Becc. 1921
L. drudei F. Muell. ex Becc. 1921
L. rigida Becc. 1921 = L. mariae subsp. rigida
L. humilis var. sclerophylla Becc. 1921
= L. muelleri

L. humilis var. novoguineensis* Becc. 1921
= ?L muelleri

L. eastonii C.A. Gardner 1923
L. japonica* Nakai ex Masamune 1929
= ?L. chinensis var. subglobosa
L tonkinensis* Magalon 1930 = ?L saribus
L. brassii* Burret 1935 = L. muelleri
L. boninensis* (Becc.) Nakai 1935
= L chinensis var. boninensis
L. Crustacea* Burret 1939 = L muelleri
L melanocarpa* Burret 1939 = L. benthamii
L. beccariana* Burret 1941
L exigua* Dransfield 1977
L. fangkaiensis* X. W. Wei & M. Y. Xiao 1982
= L. saribus

L. carinensis* (Chiov.) Dransfield & Uhl 1983
[Hyphaene carinensis Chiov. 1929]

L. endauensis* Dransfield & K.M. Wong 1987

Rodd, Revision of Livistona (Arecaceae)

This pattern of distribution invites speculation as to the origins of the genus. Its
diversity in Australia and presence in northeastern Africa suggest that its origin might
be Gondwanan, and that it may have come into existence before the separation of
Africa. The difficulty with this explanation lies in the absence of Livistona, or indeed of
any closely allied genus, from the rest of Africa and for that matter from peninsular
India and Madagascar. Dransfield (1987) tentatively advances the alternative
hypothesis concerning the Livistoninae that: The distribution of the whole subtribe
together with a rich fossil record suggests its early differentiation in Laurasia followed
by later dispersal in the southern hemisphere.' Although he states the likelihood that
the Arecaceae originated before or around the time of separation of Laurasia from
Gondwana and then developed independently in both hemispheres, he suggests that
Australian Livistona 'may have originated in invasion from Sundaland; fragmentation
of the ranges of Livistona in Australia due to climatic deterioration may have led to
speciation. Far from being a Gondwanan relic, Livistona may, I now believe, be
an immigrant.'

If Dransfield is right, such an immigration would have to have taken place after the
collision of the Australian and Asian plates in the mid-Miocene (10-20 m.y.b.p): this
rather brief time-scale hardly seems sufficient to account for the great diversity of form
found in Australian Livistona, especially when taken in conjunction with the
concentration of most of the species in refugia often associated with far more ancient
land-surfaces, and the near-sympatric occurrences in some parts of Australia of such
clearly unrelated species as, for example, L. humilis and L. benthamii. A third
alternative, and one that is admittedly highly speculative, is that proto-Livistona had
already emerged by the time Laurasia and Gondwana began to separate, and its
progeny evolved independently but extremely slowly (by flowering-plant standards)
in both landmasses and their daughter lands. This would more adequately explain the
apparent relict status of many species, particularly the outliers, also the apparent
divide between the species on either side of Wallace's Line. Uhl and Dransfield (1987),
in their chapter on the fossil record, place the earliest appearance of palms in the early
Upper Cretaceous (Santonian) of North America, and in fact this is a fan palm equated
with the modern American genus Sabal in the Sabalinae, regarded as more specialised
than the Livistoninae at least in floral structure; this generic assignment may be
questionable, but it still suggests a great antiquity for the coryphoid palms.
Unfortunately there is little in the way of Australasian fossil evidence (Dowe 1995) to
support any general hypothesis as to the origins of Livistona. Except for the distinctive
and taxonomically isolated Nypa, there appear to have been no Cretaceous or Tertiary
macrofossils of fan palms reported as yet, and the fossil pollens, dating back as far as
the Palaeocene, are either of the spinulose type assigned tentatively to subfamily
Calamoideae, or are placed in the monosulcate form genus Arecipites. The latter covers
pollen types found in a diverse range of modern palms, including Livistona.

Morphology And Terminology

An excellent account of the morphology of palms, accounts of the major groups and
the genera, including features of inflorescence and flowers, is given by Uhl and
Dransfield (1987). Tomlinson (1990) also gives a detailed, though more functionally
oriented, account of palm morphology.

1. Growth-habit and trunk

With one exception, the Australian species of Livistona are invariably solitary¬
stemmed, with a straight erect trunk — i.e. solitary and pleonanthic, fitting 'Corner's
model' of Halle et al. (1978). Only L. inermis sometimes exhibits basal suckering

Telopea 8(1): iggg

('Tomlinson's model'); this appears to develop only in older plants, sometimes quite
profusely. Death of the original stem by age or fire is apparently followed in some
plants by elongation of several (up to 6) basal suckers, resulting in a plant with several
equal-aged trunks. Not all populations of L. inermis behave in this way, some
reportedly being consistently single-stemmed with no sign of suckering.

Trunk diameter varies within and between populations of a species, but usually within
limits of c. 25% of the median value for the species. Most of the Australian species can
be accommodated in one of three mutually exclusive (or almost so) diameter classes,
centred on trunk diameters of approximately 8, 15 and 30 cm at 1-2 m above the
ground. Most species conform to a predictable pattern of relationships between
diameter, height above ground, and age. In mature plants there is a basal swelling,
much more marked in some species than in others, varying from gently bulbous to
conical to flared like a trumpet; populations may show a certain degree of variation,
e.g. from conical to flared, and much variation in degree of swelling, so that this
character can only be described for each species in very approximate terms. In some
stands of L. australis in the Sydney district it can readily be observed that the palms
with greatest access to water, nutrients and light develop the most swollen trunks
during their phase of maximum growth-rate, and these may have bases up to twice the
thickness of less well-nourished palms in the same stand. Basal swelling of the trunk
normally extends upward in an obviously conical form to a point 1-5 m (varying with
species) above ground; above this the trunk appears superficially to be of constant
diameter, but measurements of individuals which have achieved close to maximum
height show a progressive though slight diminution of diameter continuing upward.

Old individuals of most species often display a phenomenon common in many palm
genera, namely new root growth arising from subsurface layers of the trunk base at a
point progressively further above the ground with age. This results in basal plates of
surface tissue being forced away from beneath, with a dense mound or pedestal of
exposed roots developing below (Fig. Id). In some cases such pedestals can be over
50 cm high (e.g. L. mariae), or over 80 cm diameter (e.g. L. drudei). This can only be
regarded as a significant character in these and one or two other species.

Surface pattern and texture of the trunk are moderately to very distinctive for each
species. Most of the distinctive features are consequences of the particular pattern of
aging, drooping and abscission the leaves undergo. On higher parts of the trunk,
where crown size has stabilised and leaf-fall is regular, the leaves tear off cleanly with
age leaving in most species a very regular pattern of thin sheath-scars which broaden
for part of their circumference into the more prominent petiole-scars, which of course
are arranged in the phyllotactic spiral. Sheath-scars and petiole-scars are often
inconspicuous on the lower trunk. The petiole-scars vary from very smooth and flush
with the trunk, to quite markedly stepped, i.e. with a projecting ledge at the lower
edge of the scar, or more rarely are rather ragged with a torn fibrous edge. The pattern
of scars on mature trunks is further complicated by the addition of inflorescence-scars.
In a number of species the lower trunk bears persistent petiole-stubs, the result of leaves
being shed from a weak point of the petiole some way out from the trunk, rather than
from the base of the sheath (Fig. 1). These stubs may be somewhat sporadically
scattered and confined to lowest 0.5-1 m, or regular and overlapping (e.g., in
L. eastonii; Fig. lc) covering the trunk to about half its ultimate height; in L. benthamii
they project almost at right-angles (Fig. la). This feature, although distinctive for a few
species, may not be expressed in some situations due to frequency of fires or of
strong winds.

A feature of Livistoim trunks that is developed to a more striking degree than in most
other palm genera is the presence of numerous vertically aligned fissures. As in most
fan palms the peripheral trunk tissues are dead from an early age, and the fissures

Rodd, Revision of Livistona (Arecaceae)

arise from the subsequent tangential shrinkage of this dead zone. They are often very
crowded, deep, sometimes widely gaping, of varied length, and always cut across the
horizontal markings (sheath and petiole scars) at random. As might be expected,
species with very slender trunks exhibit less Assuring. Old trunk surfaces, as on the
lower trunks of tall specimens, have generally undergone considerable erosion
resulting in rounding and bleaching of the ridges, so that the Assuring becomes
shallower and more obscure; in L. drudei the bases of older trunks often become almost
completely smooth.

2. Leaf

The leaf in Livistona, as in most other palms, is a large, complex organ in which
structural variadon is evident over more levels of fineness than are normally examined
in plant taxonomic studies. Its basic morphology, despite large differences in size
between species, is remarkably constant throughout the genus in Australia.

The costapalmate leaf of Livistona can conveniently be divided into three major parts
for purposes of discussion, namely the sheathing base, the petiole (including hastula)
and the lamina or blade. These are dealt with under these separate headings.

2.1. Sheathing base

Aspects of sheath morphology of palms are covered in Tomlinson (1964, 1990, 1995)
and Uhl and Dransfield (1987). The leaf base in Livistona does not conform exactly to
any one of Tomlinson's (1990) types, but combines the dead mat of vascular strands of
his Cocos-type with the elongated ligule of his Zombia-type. In the case of sheaths of
these types Tomlinson distinguishes between the thin part of the sheath that wraps
around the trunk apex, and the thickened part that forms a lower extension of the
petiole: the latter he terms the pseudopetiole. In Livistona this is sharply distinct from the
remainder of the sheath from an early stage of expansion, the distinction being in two
features: (1) except at the extreme base of the sheath, there is an abrupt thinning from
the pseudopetiole, which is concave-convex in transverse secAon, to the sheath which
is laminar; (2) at least in its upper half, the line of junction between sheath and
pseudopetiole is along the flat or concave inner face of the pseudopetiole, a few
millimetres in from its sharp, usually prickly margin; thus the pseudopetiole overlaps
the sheath (Fig. 2b). On older leaves the dry, fibrous sheath has usually torn away from
the pseudopetiole and, since the latter forms a smooth continuum with the peAole
itself, the only means of distinguishing it is the pair of intramarginal linear scars, often
rather inconspicuous, where the Abres have tom away. For this reason the concept of
the pseudopetiole has not been employed in the species descripAons, and the 'peAole'
is taken to extend right back to the junction with the trunk. The ligule in Livistona
usually persists as an opaque, smooth appendage, at least toward its apex, as it is not
subjected to the stretching and disintegration undergone by the lower sheath.
Nonetheless, it is often only on some of the younger sheaths that this apex remains
entire; it takes the form of a roughly triangular or trapeziform lobe, its margin
normally fringed with long, slender, entire scales, and its outer (abaxial) surface with
an indumentum of mostly appressed scales, the bases of which are sunken in shallow
pits, while the inner surface is always glabrous and more or less shiny. The size, shape,
colour and texture of the untorn ligule apices are characterisAc for each species,
though differences between species are not always very obvious. It is unfortunate that
the sheaths are seldom collected, though obtaining them is admittedly very difficult.
Whenever the opportunity arises of collecting from a felled palm, they should not
be omitted.

Telopea 8(1): 1998

2.2. Petiole

As used here, the term petiole will mean that part of the leaf extending from the point
of attachment to the trunk to where the lowermost segments of the blade diverge
(see discussion of pseudopetiole in previous section).The petiole exhibits a number of
taxonomically valuable characters in its margins, surfaces and cross-section. Its length
and width are also significant, though the variation of both within a species may be
surprisingly large; both these measures are to a degree proportional to the overall size
of the palm and its leaf. In the key and species descriptions length is taken to be the
total length of a fully expanded petiole, from point of attachment to lamina base. This
should be measured on the oldest leaves at the base of the crown or, more readily, on
fallen leaves. Even if it were possible to obtain a complete petiole on a leaf collected
from close to the centre of the crown, observations suggest that it will not have
attained its full length. Petiole width, for purposes of consistency of descriptions, is
measured as close as reasonably possible of the blade, though the taper is so slight in
the apical quarter of the petiole that the exact point of measurement is not critical.

The petiole transverse section (t.s.) is more or less characteristic of a species, though
showing a surprising degree of variation in some species. As with width, the t.s. is
taken to be that found in the apical quarter of the petiole, hr the majority of species its
profile is approximately plano-convex or slightly concavo-convex, though tending
toward a broad-based triangle with its (abaxial) apex rounded-off. Less common is an
unequally biconvex profile. There is frequently a slight adaxial keel (or less commonly
2 parallel keels), often slightly asymmetric and scarcely raised above the adaxial
surface; these are possibly compression artefacts from the leaf bud. The petiole margins
in t.s. vary in degree of attenuation and whether they are curved upward (i.e. giving
the petiole a somewhat crescentic t.s.) or outward; in some species this character seems
quite unreliable, both tendencies being observable even in the one collection. In
herbarium collections there is also the problem of to what extent the t.s. shape is the
result of collapse on drying; this appears to affect different species to different degrees,
presumably reflecting differences in quantity and distribution of structural tissues.
Degree of collapse depends also on age of leaf, the younger softer ones being more
unstable in drying than the older and harder.

The petiole margin shows a number of features of taxonomic value. Most conspicuous,
and traditionally regarded as most important, are the marginal prickles, found in all
species on at least some leaves. These are largest and most numerous close to the
petiole base but field observation has shown repeatedly that prickles are often almost
entirely absent from the petioles of mature, tall plants. It is interesting to note that
Cornett (1986), discussing tire occurrence of petiole spines in Washingtonia fi l if era in
California, reports that they are absent from the apical region of petioles on plants over
8 m tall, and almost entirely absent on plants over 14 m. He suggests an adaptive
explanation for their presence only on smaller plants, connected with the maximum
reach of now-extinct herbivores of the region (Brontosaurus?!). Petiole prickles do not
show very great variation between species. In some species at least some of the
prickles are curved either antrorsely or retrorsely, but such character-states are not
very constant. Marginal prickles grade into much smaller protuberances on petiole
margins, both toward the petiole apex and interpolated between prickles. The larger
of these are here termed pricklets and are usually distinctly flattened, though often
irregular in shape and usually blunt. The smallest are mere pustules, detectable only as
slight swellings of the margin, often of contrasting colour. Between protuberances
(including prickles) the margin is apparently knife-edged to the feel. Closer
examination shows it to be rounded, though with a very small radius; in some species
more than others a semi-cylindrical marginal callus of distinct colour is evident.

Surfaces of the petiole exhibit some of the most taxonomically useful characters.

Rodd, Revision of Livistona (Arecaceae)

For purposes of standardisation these are described for the apical region of the petiole
as this part is more likely to be present in herbarium specimens. Perhaps the most
obvious surface character is colour. This is usually the same on both surfaces and is
most commonly green, with the depth of colour and any glaucousness similar to that
of the leaf blade. Some species, though, develop a strong pinkish or red-brown colour,
though only as the leaves age and probably only in conditions of strong light. Towards
the petiole base there are often areas of much stronger red-brown or purplish
coloration, seen in most species but especially prominent in some (e.g. L. drudei).
Colour may be modified by indumentum, which is discussed below.

Pitting of the petiole surface provides interesting characters. In some species pits
appear to be shallowly impressed into otherwise smooth surfaces, possibly caused by
scales being pressed into the soft surface while the leaves are furled tightly in bud.
Patterns of pitting vary greatly on different parts of the petiole and between upper and
lower surfaces, with longitudinal zones of more prominent pitting often evident. A
particular type of pitting may appear characteristic of a species but is absent from one
or two collections.

Closely associated with pits on the petiole is the characteristic indumentum. In all
Livistonas this has as its fundamental unit a peltate scale, but in a wide variety of forms.
The scale is attached to the surface by a basal-cell mass, a dense mass of granular,
usually tanniniferous cells which persists long after the scale itself has been eroded
away; it appears then as a (usually) brown dot or streak on the petiole, varying much
in size and degree of protrusion, from minute and virtually flush with the surface, to
a prominent dark brown pustule. The scales themselves vary from membranous to
thick, and in overall outline from almost circular to very elongated; they are usually
asymmetrically attached, longer toward the petiole apex, and consist of sinuously
radiating, narrow, colourless cells which may be dotted sparsely to densely with
minute tannin-bodies giving them a brown coloration. Their margins are raggedly
denticulate to deeply laciniate, sometimes divided right to the base. The laciniae are
often branched and frequently become tightly twisted and tangled with laciniae of
adjacent scales, giving rise to a shaggy tomentum. On young leaves just emerging
from the bud the scales are often seen to be neatly seated in pits, in some species
characteristically in longitudinal rows in much elongated pits; tliis arrangement
persists in species with thick, persistent, non-laciniate scales, e.g. L muelleri and
L. decipiens, but in other species with caducous or laciniate scales their association with
pits is less obvious.

Distribution and size of scales varies considerably between upper and lower petiole
surface, from base to apex of petiole, and from marginal to medial region of either
surface. Scale type (as distinct from scale density or distribution) is one of the most
valuable and constant taxonomic characters, but its use demands attention to leaf age;
there is generally little resemblance between the petiole scales of a newly emerged leaf
and the equivalent scales (as to position) of a petiole which has been exposed for
several years.

The petiole scales characteristic of a species are generally also found, with certain
predictable modifications, on the leaf-sheath, the lamina ribs and costa, and the rachis
bracts. Hairs, as opposed to scales, appear to be confined to inflorescence axes if
present at all.

2.3. Hastula

At its apical extremity the petiole continues uninterrupted on its abaxial surface as the
costa of the lamina, though with an abrupt narrowing. On its adaxial side, though, this
continuum is interrupted by the hastula, a flap of tissue (Fig. 2d) in shape and position
somewhat like the ligule of a grass leaf, though doubt has been cast (Tomlinson 1990)

Telopea 8(1): 1998

on whether it is homologous with that organ — the term 'ligule' is instead used by
present-day palm workers for the extension of the leaf's basal sheath opposite its
junction with the petiole (used in this sense, it seems to me non-homologous with the
grass ligule and should rather be termed an ocren ; and it is difficult to see why the
hastula should not be homologous with the grass ligule, as some grasses have an
elongated and narrowed upper sheath that is not unlike the petiole of a fan palm, and
its junction with the plicate blade has a very similar conformation). The term 'hastula'
was coined by L. H. Bailey (1933); earlier palm writers mostly termed it a ligule. The
hastula in its shape, texture and orientation exhibits useful taxonomic characters,
though again there is often a surprising degree of variation within a species. From an
early stage of leaf expansion it undergoes partial necrosis and progressive attrition, so
that comparisons between leaf specimens of different age can be misleading. For
assessment of this, as of other leaf characters, all palm collections ideally should
include a range of leaf ages from the one crown, but in practice of course their bulk
militates against this ideal.

Hastula morphology can be subdivided into several elements for descriptive
purposes. The line of demarcation between the upper petiole surface and the hastula
is here termed the hastula base; it is asymmetric to varying degree and in shape varies
from an almost straight line to a narrow V, or commonly is in the form of an irregular
W, making the petiole apex 3-lobed. Invariably it has at least small lateral (basal) lobes,
representing the turning outward of the petiole margins which are decurrent onto the
margins of the lowermost lamina segments. Often there is no obvious break of surface
between petiole and hastula apart from a slight to somewhat abrupt upward tilt, or
there may be an abrupt convexity terminating the flat petiole surface.

The actual flap of tissue comprising the hastula is here termed the hastula rim. This is
rarely seen before it has undergone partial necrosis, but when new and undamaged it
is frequently several times longer than on mature leaves. At this initial stage its shape
is rather indeterminate and irregular, but generally much elongated with a ragged
edge. In L. decipiens it has been observed to be 15 cm or more long. The base of the rim
is thick and leathery, but laterally and apically it is rapidly attenuated into a thin, weak
membrane. Subsequent necrosis gives rise to a green inner zone, usually of even
width, and a necrotic margin which varies greatly in width and the pattern of its
attrition, both between and within a species, and indeed frequently between the leaves
of one tree. In some species, e.g. L. australis (Fig. 2a), the hastula is commonly semi-
cylindrical or almost cylindrical, though a wider sample may reveal this to be an
ontogenetic stage. It seems clear, too, that the cylindrical form is consequent upon a
leaf form in which the petiole is at near right-angles to the lamina and the segments
radiate in an almost complete circle. If the angle is somewhere intermediate between
90 and 180 degrees, then the hastula frequently takes the form of a squashed cylinder,
with lateral edges folded back over the petiole apex.

On the abaxial surface the petiole in all Livistona species flows smoothly into the costa,
at least with respect to its keel region; the margins are abruptly narrowed. There is no
evidence of the abaxial hastula (dorsal crest) found in some other coryphoid genera,
except for a coloured, obscurely stepped, V-shaped mark in L. fulva, cutting across the
costa at varying distance from the petiole. It may be significant that this is one of the
least strongly costapalmate species.

2.4. Lamina

The lamina (and indeed the whole leaf) in Livistona appears to have no characters by
which it can be distinguished from that of several other genera of the Livistoninae,
e.g. Brahea, though a majority of species share a facies of subtle characters which may
allow sterile plants to be recognised as Livistona, e.g. in collections of cultivated palms.

Rodd, Revision of Livistona (Arecaceae)

Aspects of its morphology under which characters and character-states are discussed
below are size, outline, undulation, division, segment shape, thickness, rib profile,
colour, texture, indumentum. Also significant is the costapnlnmte nature of the lamina
in Livistona — in effect it is not palmate in the narrowest sense, in that not all segments
radiate from the petiole apex but rather some of them arise from a central rib or costa
which forms a continuation of the petiole. Measurement of degree of costapalmateness
is discussed below.

Lamina size is one of the most useful and easily assessed characters. As might be
expected it shows a fair degree of correlation with overall size of the palm, which is to
a fair degree independent of environment: adverse conditions do not give rise to a
very much smaller lamina size, though often petioles are significantly shortened thus
giving a more compact crown. Only one measure of lamina size is used in species
descriptions, length from hastula base to leaf apex along central segments. This is most
readily gauged with a tape-measure, holding one end at the hastula base and grasping
a handful of the central segments which are then stretched to their maximum length
against the tape. In an undamaged leaf it will always be found that the longest
segments are either those closest to terminal on the costa or within 2-4 segments on
either side. The chief difficulty in measuring length arises from the necrosis and
subsequent attrition of the fine segment apices, more pronounced and earlier in onset
in some species than in others. If at least one newly expanded lamina is available for
a species, a rough extrapolation is possible for older leaves with damaged segment-
apices, based on width at the point of breakage. The reason for only this one measure
of size being employed is that all other possible measures appear to be correlated with
it, or can be dealt with under lamina outline. It was concluded early in tins study that
it is quite impractical to measure lamina length as the maximum distance from apices
of central (distal) segments to apices of the lowest (proximal), or most rearward¬
pointing segments.

Lamina length varies in Australian species from a minimum of around 0.3 m in
L. humilis to a maximum of around 2.2 m in L. marine. Within-species variation so far
observed is in the range 15-35% either side of the mean. Some specimens showing a
shorter lamina length have been excluded from species accounts if the collections were
sterile, as juvenile leaves are generally smaller (sometimes appearing large but with
segment tips less attenuated).

Lamina outline, here taken to mean the curve traced out by the segment apices together
with basal gap or sinus, is not easy to assess or quantify in most species. This is due
partly to the 3-dimensional shape of the lamina in most species, partly to the drying
artefact of contraction as in a folding fan, which greatly widens the basal sinus of
loosely pressed specimens.

There is rather little variation between species in length to width ratio of the lamina
but this may be difficult to assess because of the difficulty of measuring width. Clearly,
if the leaf was circular in outline the width would be twice the length as defined above.
Measurement reveals that this is never the case, since there is progressive shortening
of segment length from apical to basal segments. Lengths of all segments were
measured on several sample leaves for of two species, L. humilis and L. lorophylla: it
was found that segment length remains constant or increases slightly for the first few
segments (going outward from costa position at leaf apex), then falls off in an almost
linear progression to about half its maximum for the basal segments. For this purpose
length was measured from the hastula for all segments, even though, strictly speaking,
in costapalmate palms such as these, many of the apical segments do not actually
originate from the hastula but from the costa well above the hastula.

The pattern of dissection and folding of the lamina provides a number of important
characters. As illustrated by Dransfield (1977), the primary, or deepest divisions are

Telopea 8(1): 1998

along the adaxial ribs, i.e. each adaxial rib terminates at a sinus between two segments.
Each segment is additionally, shallowly to deeply bifurcated, the sinus between the
lobes terminating an abaxial rib.

One of the most obvious but difficult to quantify differences between species is in the
depth of dissection of the lamina. L. H. Bailey proposed the concept of the palman as one
means of expressing degree of dissection of a fan palm leaf. The palman is the central,
undivided part of the lamina, bounded by the sinuses of the primary segments. The
patterns of dissection of the mostly strongly costapalmate leaves of Livistona are such
that palman dimensions are difficult to measure, and other measures of dissection
have therefore been devised for this study. The degree of primary dissection has been
expressed (as a percentage) by comparing the length of the free portion of a segment
with its total length. Between about the second segment from the costa apex and at
least the fifth segment, the ratio of total segment length to length from hastula to the
intersegmental sinus remains moderately constant in any one leaf. Depth of dissection
has therefore been measured on one or more of the second to fifth segments depending
on which is available and relatively undamaged on a specimen. It should be regarded
as a relatively crude measure, but this is acceptable when considering the variation
encountered between trees and between populations. In some species the variation is
considerable, though useful contrasts still show up between most species-pairs which
might be confused on other characters. If anomalously low values are found for any
specimen it has been reassessed for possible juvenility (see discussion of this aspect
below), and if suspected to be juvenile it has been excluded.

A feature that is rare or at least poorly expressed in the Australian species is the
presence of 'compound segments', consisting of groups of two or more segments in
which the primary sinuses are displaced to much closer to the leaf's perimeter. In a
number of the species, close examination of the smallest segments close to the base of
the lamina will reveal some compound segments; in L. fidva they are often present
among the medium-sized segments midway to the lamina apex, though not at all
conspicuous; the same applies to some specimens of L. australis.

Depth of bifurcation of segments is expressed by comparing the length of the bifurcated
portion to the free length of the segment. It is even more affected by juvenility than is
depth of dissection. The majority of species have deeply bifurcated segments in the
adult state, the lobes accouxating for around 50% or more of the segment's free length.
Only 2 species, L. muelleri and L. fulva, consistently have very shortly bifurcated
segments (under 10% of free length). A remarkable feature of some specimens of
L. drudei is an apparent secondary bifurcation of the lobes of up to a few centimetres
(occasionally more than 10 cm) in depth.

Width of segments has been taken as the maximum found on any one specimen,
measured at the widest point above the intersegmental sinuses but below the point of
bifurcation. This also shows considerable variation within a species, sometimes even
between different leaves from the one tree, but the ranges obtained still show useful
distinctions between species. Width is at a maximum between about the second and
fifth segments either side of costa, falling off somewhat erratically among the very
apical segments, and in the opposite direction falling off toward the lamina base in a
fairly regular progression.

Outline of the individual segments includes some useful characters, apart from the
crude quantitative measures just discussed. In some species the free part of the
segment below point of bifurcation is parallel-sided, in others it may taper slightly
toward the apex, in others slightly toward the base. This tapering appears consequent
upon depth of dissection since the segments of deeply dissected leaves will fit better
against one another if tapered toward the base, whereas very shallowly dissected

Rodd, Revision of Livistona (Arecaceae)

leaves demand a taper in the reverse direction. Tapering is not always consistent
within a species. Variation is far greater in the lobes both in outline, and also in
direction to a minor degree. In some species they diverge noticeably from the point of
bifurcation, though this should be confirmed by pressing the segment against a flat
surface. In most species the outer margins of the segments continue in a straight line
and the inner margins of the lobes form a narrow V, but in some the inner margins
appear more convergent. Outline of the individual lobes varies in a number of ways:
an interesting phenomenon seen on some (even most) specimens of nearly all species
is a curious 'step' in either the inner or outer margins of each pair of lobes. Even in the
one collection, this step can vary greatly both in presence and position between
different leaves and between segments of the one leaf. When the two lobes of a
segment are folded together, as in bud, the steps coincide perfectly. This, combined
with its erratic occurrence, suggests that it is in a sense an 'artefact' of compression of
the leaves against one another in bud, and is of doubtful taxonomic value.

Apices of the lobes vary greatly in their degree of attenuation, some being finely
aristate or even hairlike, others tapering evenly to a fine or more abrupt point. Usually
the lower, narrower segments (closest to petiole) have more finely attenuate lobes, but
for purposes of standardising descriptions only those of the upper, broader segments
are considered. Considerable variation is also found in degree of necrosis, brittleness,
and consequent attrition of apices and the earliness of its onset in relation to leaf
expansion. This pattern appears characteristic of a species.

A distinctive feature of many fan-palm leaves is the presence of thread-like
appendages in the intersegmental sinuses, terminating the adaxial ribs. Attention has
been drawn to these by morphologists such as Eames (1953) and their relationship to
the development of plications in the leaf primordium explained. They are evident in
nearly all the Livistona species examined, but vary in thickness, texture, indumentum
and persistence. Their length is initially indefinite (up to a maximum of the total
lamina length) but their early necrosis and consequent brittleness results in short
remnants only being found on fully expanded leaves. Nonetheless each species seems
to retain a characteristic length of appendage, within rather broad limits. In the light
of the explanations of their origin it is hard to see why they do not occur also in the
sinuses between the segment lobes, terminating the abaxial ribs, but this does not
appear to be the case.

Another major aspect of the overall plan of the leaf, so far hardly considered, is the
degree to which it is costapalmate. The costa is most evident on the abaxial surface
because it is roughly triangular in transverse section, displaying a broad face on the
abaxial leaf surface. Many of the lower segments join the margins of the petiole apex
rather than the costa but there is no point at which it is possible to say that one
segment arises from the petiole apex but the next, more distal segment arises from the
costa. There is progressive stretching apically of the costal 'internodes', or intervals
between the points of attachment of segment midribs. Toward the costa apex the final
2-4 'nodes' are widely but often rather unevenly spaced and the costa finally
terminates in a sinus between the two co-apical segments, which are sometimes of
greatly reduced width and irregularly formed compared with the third to fifth
segments out from the co-apical ones. The position of this uppermost sinus relative to
total lamina length is found to be quite erratic on leaves from the one tree, leading to
the conclusion that overall costa length as a proportion of lamina length is not a very
useful taxonomic character. Since the great majority of segments in all species join the
costa/petiole-apex in the basal 10% of total lamina length, a more useful measure of
costapalmateness is given by the proportion of all segments which join it beyond that
region. This proportion (expressed as a percentage) was calculated for three
representative specimens of each species (Table 2).

Telopea 8(1): 1998

Table 2. Leaf laminae of Livistona species

Collection

Lamina
length
in cm

Total no. of
segments

No. arising
above basal

10% of
lamina length

Percentage
arising
above
basal 10%

L mariae subsp. mariae

Rodd 3215

L. mariae subsp. occidentalis

218

15.1

Rodd 2868

176

22.8

L. mariae subsp. rigida

Rodd 2937

L. lanuginosa

161

17.9

Rodd 3759

129

16.7

Irvine 1912
leaf 1

188

21.8

Irvine 1912
leaf 2

163

17.2

L. benthamii

Specht 1187

15.6

Jacobs 1790*

10.0

Rodd 2904

20.3

L. drudei

Rodd 3135
leaf 1

140+20

22.7

Rodd 3135
leaf 2

150+10

17.4

Irvine 1832*

125

12.7

Irvine 1833*

100

13.2

L. australis

Rodd s.n.,

28.viii 75

11.5

Rodd 3615

121

10.6

Irvine 1928

9.6

L. nitida

Rodd 3055

177

14.5

Rodd 3051

173

12.7

Telford 5648*

105

15.9

L. decipiens

Rodd 3045

185

21.7

Rodd3117

c. 180

22.5

Rodd 3069

165

16.1

L. fulva

Rodd 1166

4.7

Rodd 3062

6.3

Rodd 3063

30*

6.7

Rodd, Revision of Livistona (Arecaceae) 63

Collection

Lamina
length
in cm

Total no. of
segments

No. arising
above basal

10% of
lamina length

Percentage
arising
above
basal 10%

L. muelleri

Hind 748

1.8

Hind 216

3.3

Rodd 3170

5.2

Rodd 3159
leaf 1

1.8

Rodd 3159
leaf 2

5.2

L. humilis

Benson 950

9.4

Gittins 2646

12.9

Rodd 2926

12.5

L. eastonii

Maconochie 1228*

19.5

Symon 10254

18.5

Rodd 2887

20.0

L. alfredii

Rodd 2847

103

23.3

Rodd 2848

130

28*

17.8

Wark A

122

11.1

L. victoriae

Rodd 2934
leaf 1

112

17.0

Rodd 2934
leaf 2

107

19.2

Rodd 2800*

10. 2

Parker 1065*

12.2

L. kimberleyana

Symon 10154*

18.2

Rodd 2866
leaf 1

17.5

Rodd 2866
leaf 2

20.9

L. lorophylla

Kenneally 2162

27.3

Telford 6404

18.7

Rodd 2871

22.9

L. inermis

Telford 8114

13.6

Telford 8064

3.0

Benson 952

9.5

Symon 10347

7.9

* Specimens possibly from juvenile plants.

Telopea 8(1): 1998

Table 2 shows that this measure of costapalmateness varies within a fairly narrow
range for all the Australian species. Although the minimum value found was 1.8%
(some plants of L. muelleri ), most of the collections of all but two species showed
values in the range 8-25%. Thus, the degree of costapalmateness among the Australian
species is all of a similar order, and this is borne out by a careful subjective assessment.

A few species, e.g. L. decipiens, appear superficially to be very strongly costapalmate,
but this is an illusion caused by the very deep dissection of the lamina, which serves
to emphasise the costal region. It is also evident that the measure of costapalmateness
shows large variation within a species. Some of this variation is due to operation of
chance factors, exaggerated by the very small sample surveyed. If, for example, the
value were based on only three or four segments out of 30, then one less (which may
just miss being included) will cause a major increase in the value; sometimes two
segments, one each side of the costa, both barely fall outside the basal zone.

Degree of costapalmateness is nonetheless shown to be quantifiable by this measure.
The lamina is here described as weakly, moderately, or strongly costapalmate, depending
on whether respectively 0-7%, 7-16%, or over 16% of its segments join the costa
beyond its basal 10% zone. These boundaries are chosen to give the most useful
discrimination between species. By this criterion L. muelleri and L. fulva are weakly
costapalmate; L. humilis, L. australis and L. nitida are moderately costapalmate; and
L. eastonii, L. kimberleyana and L. lorophylla are strongly costapalmate. Other species can
be described as weakly to moderately, or moderately to strongly costapalmate.

Another lamina character which emerges in Table 2 is number of segments in the
lamina. It will be seen that this is reasonably constant for each species, except where
juvenility is suspected, and thus appears taxonomically useful. It is not always easy to
count accurately, in that the basal segments are sometimes of irregular width,
suggesting that some may be fused together, or are sometimes vestigial. There are
sometimes also one or two vestigial apical segments. However, an error in counting of
± 5% is regarded as acceptable for purposes of description.

One of the most striking characteristics of the fresh whole leaf in most of the
Australian species is the pattern of undulation of the lamina. This takes the form of
undulations, which are best seen in a tangential cross-section of the lamina at right-
angles to all the segments (e.g., by cutting a small circle out of the centre of a fresh
leaf). Both number of undulations and their amplitude vary between species, and
amplitude also varies between collections of a species depending on both maturity
and exposure. The most extreme undulations are on very mature palms in the most
exposed, sunny places. Invariably there is a median trough along the costa (viewed
from above), which in most species is gently to strongly downward-curving (Fig. 9a),
adding a further dimension to undulation of the lamina. On either side of the trough
there are one to three crests of undulations, referred to in the descriptions as 'adaxial
undulations'. Approaching the basal margins of the lamina the undulations become
tighter and usually the margins are sharply inflexed or involute, so that some basal
segments are resupinate. In several species there is an additional, very tight fold of the
last two to three segments back in the other direction, i.e., the resupination is reversed
by a double fold. The spatial relationship of these basal folds with the petiole apex and
hastula is complex, and their morphogenesis is almost impossible to visualise. To
speculate on the adaptive value of undulation, its exaggeration under exposed, high
light conditions makes it likely that it is a means of reducing absorption of solar
radiation, and possibly also of strengthening the lamina against wind damage. There
is some degree of correlation between lamina size and degree of undulation.

Another feature which affects the three-dimensional shape of the leaf in the palm
crown is drooping of the segments. In some species, most notably L. decipiens and

Rodd, Revision of Livistona (Arecaceae)

L. australis, the segment lobes all hang vertically, undergoing a moderately sharp bend
just above the point of bifurcation. This would appear to be the result of three
characteristics of such leaves: (1) very long lobes; (2) thin lamina texture, lacking
mechanical strength; and (3) the V-shaped section of the segment below the point of
bifurcation, imparting mechanical strength in this region. Most other species have the
segment lobes drooping to some lesser degree, but a few, most notably L. muelleri, have
segments which are stiff and straight for their whole length; this correlates with very
short segment lobes.

The ribs of the lamina present a number of characters, chiefly in their cross-sectional
shape and dimensions. In most species they are roughly rectangular in t.s., between 1.5
and 5 mm deep and 0.5 to 1.5 mm wide. Generally the abaxial ribs are slightly larger
in t.s. than the adaxial, though this is not always the case. The summit of a rib is
usually squared off more or less sharply, but often one edge is more rounded than the
other. In some species the ribs are more elliptical in t.s., or subtrapeziform, with the
narrower dimension adjacent to the leaf surface. The measurements in descriptions are
expressed as depth, which is to be taken as the amount by which the rib stands above
the leaf surface, and width, which is the greatest width. These have been measured
with a dial caliper on dried specimens, at approximately their mid-points (which are
much closer to the hastula on the adaxial ribs), choosing the more robust ribs on any
one lamina.

Lamina thickness was also measured with a dial caliper, and shows less variation
between species and more within a species than might be expected, given the
differences in rigidity that are readily perceived. It may be that rigidity of the lamina
derives largely from the ribs. The greatest thickness found was 0.45 mm, the least
around 0.15 mm, though on a given leaf measurement of thickness was found to be
repeatable only to around 0.05 mm. Measurement was standardised at a cut end about
halfway between base and apex, with the narrow blades of the caliper paralleling and
between major veins which are slightly thickened. Care was taken to avoid excessive
pressure on the blades. Surprisingly little difference in thickness was found between
fresh and dried leaves when both were available. Variation between different parts of
the one leaf was slight, except that there was detected a tendency for thickness to
increase somewhat towards the lobe apices.

Leaf indumentum is of great importance. The Australian species exhibit a great
diversity of indumentum type, with sometimes more than one type present on a leaf,
as well as great variation in density and distribution over the various parts of a leaf.
Two main classes of indumentum can be recognised: (a) narrow chaffy scales
('ramenta'), and (b) sessile, more or less disc-like, closely appressed peltate scales,
often sunken in elongated pits. Scales of the first type are mostly shed or wear off as
the leaf ages, especially on the lamina, and it is on the petioles and basal parts of the
costa and ribs, mainly on the abaxial surface, that they are densest and most persistent.
The sessile scales of the second type are attached more securely and are more or less
identical to the scales of the upper petioles, under which heading (2.2 above) their
morphology is discussed.

The nature of the cuticle and associated surface layers of wax on the leaf lamina are
significant. The species with very thick-textured, xeromorphic, bluish or greyish
leaves, e.g. L. alfredii and L. eastonii, seem to have the cuticle impregnated with wax
which is not readily rubbed off. Other species, e.g. L. mariae and L. lanuginosa, have a
very thick but soft layer of wax which seems quite distinct from the cuticle and easily
rubbed off; it is usually most noticeable on the abaxial surface, but a thinner almost
transparent layer may also be present on the adaxial surface. In some species the wax
layer is very thin and only detectable on the undersurface of newer leaves, e.g.
L. muelleri and L. fulva. Most species of more mesic environments, e.g. L. benthamii,

Telopea 8(1): 1998

L. drudei and L. australis, have no detectable glaucousness of the leaves, the surfaces
being deep green or brownish green and more or less glossy.

Venation of the lamina is of limited use as a key character but does show a
characteristic range for each species. Each intercostal area of lamina has longitudinal
veins of several orders of prominence in an alternating sequence (see anatomical
diagrams). The prominence of these veins and their spacing varies within fairly
narrow limits, with the least prominent in those species with xeromorphic, very
opaque leaves. The number of prominent veins which can be counted across a half¬
segment (interval between an adaxial and abaxial rib) varies between certain limits for
each species, but the overlap between species is so large that it is rarely of use as a
distinguishing character.

The longitudinal veins are invariably connected at regular intervals by transverse
veinlets ('commisures') which show varying degrees of irregularity in length,
orientation and curvature. In relation to the longitudinal veins some are steeply
oblique (though direction of slope may frequently reverse), others almost at right
angles. A veinlet may bridge the interval between adjacent prominent longitudinal
veins, or may pass beneath two or more such veins, sometimes crossing nearly all the
longitudinal veins in a half-segment. They may be virtually straight or are often
curved into a slight S-bend (sigmoid); in longer veins the S-bends may combine to give
a wiggling form. Like the longitudinal veins, the transverse veinlets are most
prominent in the species with thin-textured, deep green leaves, where there often
show up strongly by their whitish colour and translucency. Unfortunately their
appearance in the fresh leaf has not been consistently recorded.

In transverse section (Fig. 3, 4) there are consistent differences in the shape and
distance between major vascular bundles; continuity and number of layers of the
hypodermis of upper and lower surfaces and presence of buttresses of the hypodermis
extending to the major vascular bundles; and the shape, size and distribution of the
fibre strands.

3. Inflorescence

Inflorescences nearly always appear several in the palm crown at the one time, among
the bases of living leaves. In all Australian Livistona species, as in most coryphoid
palms, the flowering and fruiting cycle is completed, while the subtending leaves of
inflorescences are still close to the centre of the crown, and the leaves remain alive long
after the inflorescences have fallen (unless fire-killed); thus the inflorescences at all
stages are intrafoliar rather than infrafoliar. Inflorescence length as an absolute
measure varies considerably in most species, and for purposes of characterising the
species is best measured by relating it to leaf length, e.g. shorter than petiole;
exceeding petiole but shorter than lamina; and so on. Inflorescence attitude is of some
significance, though this changes from earliest to later anthesis and even more so
between anthesis and fruiting, when weight of fruit results in most inflorescences
being pendulous.

The basic pattern of inflorescence architecture in Livistona is one with an elongated
first-order axis (counting trunk as zero-order, as in Tomlinson & Soderholm 1975)
which is dominant over a number of equally spaced, more or less evenly sized,
second-order branch systems (Fig. 2c). These branch systems, together with the similar
branch-system terminating the first-order axis, are here termed partial inflorescences,
and there may be as many as 15 partial inflorescences in one inflorescence. When a
whole inflorescence is dissected they are found to be arranged in a secund-distichous
orientation. The first-order axis, or rachis is roughly triangular in section with a rounded
adaxial side and two abaxial faces, from which the second-order axes branch alternately.

Rodd, Revision of Livistona (Arecaceae)

This variation from a strictly spiral phyllotactic arrangement is similar to that
described by Tomlinson and Moore (1968) in first-order branches of Nannorrhops
inflorescences, a variation that they ascribe to the 'requirements of available space'
such that 'any inherent phyllotactic arrangement is suppressed'. They also note that
there is a gradual change from distichous to spiral arrangement of second-order
branches toward the distal ends of the first-order branch, though the spiral is often
obscure. A similar change has also been observed in all Livistona inflorescences examined.

In all but three, or possibly four, of the Australian species the inflorescence conforms
to the pattern described above, and it seems reasonable to assume this represents the
primitive state for the genus. One departure from the pattern is found in L. inermis ; in
this species there is a reduction in the number of partial inflorescences accompanied
by enlargement of the basal one to a size subequal to the remainder of the
inflorescence, resulting in a very short and broad inflorescence (Fig. 20d); this state is
also referred to by Uhl and Dransfield (1987, p. 17) for the non Australian L. woodfordii
and L. rotundifolia (neither of which appears to have a close relationship to L. inermis)
Another departure is suppression of lower branches on an axis, seen in an extreme
form in L. humilis (Fig. 13c), in which the rachis is long and cane-like with up to 8 lower
nodes bearing sterile bracts only, i.e. with no lateral partial inflorescences, so that in
effect only the terminal partial inflorescence remains. This form of inflorescence occurs
only on fruit-bearing plants of L. humilis, in contrast to the functionally male plants
which have the conventional inflorescence structure for the genus.

Of significance in distinguishing species is the number of further orders of branching
in the partial inflorescences: this varies from 2 in L. benthamii to a maximum of 5
(usually 4) in L. australis, though the highest number for each species is usually found
only on one or two of the major branches of the lowest partial inflorescence. In the
species descriptions this is expressed in the style 'partial inflorescence ... branched to
n further orders...', but the total number of orders of branching is then n + 2 if the
palm trunk is treated as the zero-order axis.

What appear superficially to be the ultimate, flower-bearing branches, i.e. the
rachillae, are slender and spiciform and mostly over 2 cm long; but, more evidently in
some species than others, close examination reveals that the flowers themselves are
borne in clusters, the axis of each cluster representing a further order of extremely
condensed branches, in some cases approaching zero in length and reduced to the
merest knob on the apparent rachilla (which is, in fact, hereafter termed the rachilla,
following the normal convention in palm descriptions). In some species these ultimate
axes, hereafter referred to as the cluster axes, mostly bear only a single flower, but there
is always some proportion of them bearing 2 or more flowers. The size, shape and
other features of the cluster axes are of considerable taxonomic significance, as are also
the number of flowers per cluster (expressed as a range), and the width of spacing of
clusters of along the rachis in some cases.

Together with their flowers, the cluster axes are referred to as cincinni by Uhl and
Dransfield (1987), who refer (p. 19) to Uhl (1969) in which developmental studies were
used to demonstrate the sympodial nature of apparently equivalent structures in
another Coryphoid genus, Nannorrhops. In some Australian Livistona species, however
(for example L. dccipiens — Fig. 11), the cluster axes can be quite elongated and even
appear at times to grade into the rachillae, which may call into question their
sympodial nature. Other species, though, exhibit a constant sessile flower cluster
resembling that illustrated by Uhl & Dransfield (p. 192) for L. chinensis, while L. mariae
has flowers arranged secundly on the outer face of a sharply deflexed and flattened
cluster axis, appressed against the rachilla; this arrangement (Fig. 6) superficially
resembles the acervulus of some Hyophorbeae as described by Uhl and Dransfield
(p. 21) except that the cluster axis is not adnate to the rachilla.

Telopea 8(1): 1998

Several characters of the rachillae are useful in discriminating species, notably their
length (expressed as a range), thickness (though accrescence in lruit must be kept in
mind), colour, features such as wrinkles or striation, and indumentum. With regard to
the last, it seems a common feature of inflorescence axes, in contrast to bracts, to be
quite glabrous and indeed the scales characteristic of foliar structures are absent from
all such axes. But in a minority of species the axes are noticeably papillose, and in a
very few they are very shortly pubescent (e.g. L. humilis). The degree of papillosity or
pubescence is usually greater on the rachillae than on thicker axes.

Of importance in interpretation of Livistonn inflorescences are the bracts. As leaf-
homologues, they share the general palm character of being closed, i.e. tubular, though
this is not obvious when the bracts are much reduced. The inflorescence rachis at least
bears well developed tubular bracts which overlap one another (Fig. 2c). The
lowermost bract on each axis is the prophyll, identifiable by its adaxially displaced
two keels, as well as by its position and the fact that it is empty, i.e. does not subtend
a branch. The prophyll of the primary axis resembles the other tubular bracts except
that it is usually shorter and thicker in texture. In most Australian Livistoun species
there are no empty peduncular bracts apart from the prophyll: the exceptions are the
'male' inflorescences of L. humilis (noted above) with up to 8 tightly overlapping
empty bracts. Apart from the prophyll, the other bracts clothing the primary axis,
termed rachis bracts following Uhl and Dransfield, are also 2-keeIed but with keels
fully opposite one another and less prominent, sometimes barely detectable. The
rachis bracts of most species are 2-lipped at the mouth with identical lips positioned
dorsally and ventrally, at least on the distichous part of the axis, and two laterally
positioned keels each terminating in acute sinuses between the lips. The lip apices are
generally acute or acuminate, though often torn. The rachis bract apices of L.fulva are
exceptional, irregularly laciniate with several to many points of equal length; it is
difficult to detect whether these result from tearing of tire bract apex, or are a normal
morphogenetic feature.

The character of the rachis bracts is strongly influenced by their texture and by the
tightness with which they sheathe the axis. Aspects of texture are thickness, rigidity
(proportion of mechanical tissues to parenchyma), fissility, and earliness or lateness of
tissue necrosis. Three extremes are exemplified in the following species: (1) L. mariae
— bracts tightly sheathing, thick, moderately rigid, hardly splitting, remaining green
at least to early fruit (except at apices); (2) L. alfredii, L. victoriae — bracts loosely
sheathing, thick and very rigid, usually much buckled with irregular splits and tears,
tissues dead by anthesis; (3) L. benthamii — bracts tightly sheathing, thin, smooth and
straight, remaining green at least in lower part up to about anthesis.

Prophylls of the primary branches (partial inflorescences) are themselves tightly
enclosed within the rachis bracts, with only their upper parts protruding. In texture,
colour, indumentum and shape of apices they resemble closely the rachis bracts.
Progressing further outward through the orders of branches there is a rapid reduction
in bract size and complexity, so that the bracts subtending the rachillae are small,
triangular, membranous organs with no apparent tubular base. The flower clusters
have even smaller subtending bracts, termed here the cluster-bracts — usually
displaced to one side. Their proper shape is usually only discernible pre-anthesis, as
they rapidly shrivel and disappear leaving only a basal rim that is often barely
detectable by the time of early fruit development. The bracteoles associated with
individual flowers of a cluster are often more difficult to detect and even more difficult
to assign to a particular flower in the cluster, such is the degree of displacement they
appear to have undergone. In the relatively few collections in which bud-stage
inflorescences are available, both cluster-bracts and bracteoles are seen to be very
delicate and membranous, with the apex drawn out into a fine hairlike point, and in

Rodd, Revision of Livistona (Arecaceae)

some cases apparently laciniate. Because of their absence or vestigial presence in most
available collections, they are not useful in distinguishing species.

4. Flower

Flower structure is fairly uniform in the Australian species and conforms to the
general pattern found in most of the 12 genera of Livistoninae (see Uhl & Dransfield
1987). Overall shape of the flower is noted in the descriptions, e.g. funnel-shaped, cup¬
shaped, rotate, but these should be interpreted with some caution as it is possible they
change between the late pre-anthesis and early post-anthesis stages and, given the
paucity of flowering collections of most species, it has not been possible to compare
these stages. Overall length of flower may also be of value, though only within broad
limits; it has been found to vary by as much as 150% (minimum to maximum) in
species for which a number of flowering collections were available. In the case of
rotate flowers, length was measured from the very base of the flower along the back
of a petal to its apex.

The calyx is always much shorter than the corolla, and the sepals much thinner than
the petals; at the base they are fused into a shallow cup. The base of the calyx runs into
a stalk-like structure, up to about 1 mm long in some species, which is sharply
constricted at the point where it attaches to a projection of the cluster axis. This is
always the point of abscission when unfertilised flowers are shed. The stalk-like
structure is here termed the anthopodium following Briggs and Johnson (1979), who
define it as the intemode between the flower and the most distal node below the
flower. Since 'pedicel', as used by Uhl and Dransfield (1987), is described as sometimes
bearing prophylls, it does not always equate with pedicel, and anthopodium is the
more precise term. The base of the anthopodium is generally trigonous and concave in
Australian Livistona species, the angles corresponding to the bases of the sepals
(Fig. 12). Its junction with the cluster axis (or a minute projection thereof) is usually
very constricted, concealed beneath its concave base.

The valvate petals are nearly always thickened at their edges and apices and are often
somewhat cymbiform. A distinctive feature is the 'sculpting' of their inner faces,
usually with 2 longitudinal ridges which separate 3 concave depressions, the median
one of which may extend as a hollow under the thickened apex. There would seem to
be a correspondence between the petal concavities and the stamens, suggesting that
they result from tight packing of the flower parts in early bud stage, though
differential expansion usually results in the petal concavities being much larger than
the anther locules. They are rather too constant a feature to be of much taxonomic
value, and such variability as they do show seems to be strongly correlated with petal
thickness. A feature of the petals that does not appear to have been reported
previously is their basal auricles, present to at least some degree on nearly all species
examined. They are not truly basal, since they arise just above the point of fusion of
the petal bases, in the inter-petal sinuses. These auricles are not obvious, being mostly
membranous and tightly overlapping with those of adjacent petals, and usually need
prising apart with a fine needle before they become visible. The species in which they
are most readily seen is L. inermis (Fig. 21). Their direction varies from retrorse, i.e.
giving the free part of the petal a sagittate outline, to laterally diverging, and their
outline from acute-triangular to broadly rounded; they are commonly larger on one
side of the petal than on the other.

The stamens show few distinctive characters apart from the dimensions of their
filaments and anthers. The short, flattened filaments are both connate basally for up to
half their length and adnate to the petals for about the same amount; they are rather
abruptly acuminate at the apex, and show some variation between species in the

Telopea 8(1): 1998

degree of angularity or 'shouldering' at the point where they start to narrow — the
term shouldered has been used where this angularity is well developed, e.g. in
L. humitis (Fig. 14); in contrast, a species with unshouldered filaments is L. australis
(Fig. 10). Stamen length within a species appears to vary with petal length, so it
seemed most appropriate to express it as a proportion (expressed here as a fraction,
e.g. %) of petal length, both measured in an opened-out flower from the base of the
petal tube. The range among the Australian species is not large, between % and %>, and
this character may not be very significant. And finally there is anther size, expressed
here only as length since their shape is fairly uniform: in most species the anthers are
around 0.4 mm long, but those of L. lorophylla examined were only 0.2 mm, and at the
other extreme L. australis anthers were 0.5-0.7 mm long — this is also the largest-
flowered of the Australian species.

The carpels likewise show few distinctive characters at specific level. As with the
stamens, it seemed most appropriate to describe their size in relative terms, in this case
as a proportion of stamen length. Some differences were observed in shape of the
ovary portion, in style length and in the abruptness or otherwise of the carpel's
constriction into the style.

5. Fruit

The fruits of Australian Livistona species are of rather uniform type, but they provide
a rich suite of taxonomic characters. Fruit size varies considerably, with diameter as
small as 8 mm in ripe fruit of L. humilis, to as large as 35 mm in L. lanuginosa and
L. alfredii. Variation is considerable between individuals of a species, for example 13-22 mm
in L. australis, but fruit size is still a useful diagnostic character for some species. Fruit
shape in the majority of species is globose, the main exceptions being the shortly
ellipsoid or obovoid fruits of some of the more slender northern species, viz. L. humilis,
L. inermis, L. eastonii, L. lorophylla and L. kimberleyana. Fruits of some other species, viz.
L. benthamii, L. drudei, L. muelleri, are somewhat pyriform. A frequent feature is a slight
flattening of the ventral (adaxial) side of the fruit, corresponding to the base of the seed
coat intrusion into the endosperm, with the result that the diameter measured
dorsiventrally is slightly less than the diameter in the lateral plane. In some species the
ventral suture is evident in the form of a slight crease even in the ripe fruit; less
common is an apical mucro (ventrally displaced to varying degree) representing the
stigma. In common with most other tricarpellate, apocarpous genera Livistona
normally aborts 2 of the carpels at an early stage, and their vestiges may sometimes be
observed at the base of the fruit. A search among large numbers of fallen fruit will
sometimes turn up the odd one in which 2, or in very rare cases all 3, of the carpels
have developed, joined together at the base only, though it is more common to find the
additional carpels have developed only partially, their enlargement evidently
suppressed at an intermediate stage of growth. It is very doubtful that such aberrant
fruits have any taxonomic significance at species level.

The stalk of the mature fruit is more complex in structure than may appear at first
glance. In most species it is a compound structure consisting of three main elements,
all undergoing considerable enlargement post-anthesis. The calyx is persistent,
sometimes remaining attached to fallen fruits; between the calyx and fruit base there
is usually a fleshy pad of tissue representing the expanded remnants of the fused
petals and filaments. Below the calyx, the anthopodium also enlarges greatly,
especially in diameter, with its lower rim swelling downward to overlap and press
hard against the similarly enlarged but slightly more woody cluster axis. It is very rare
for more than one fruit to develop from the flowers of each cluster, hence the cluster
axis appears part of a single fruit stalk; and in fact only a small minority of the clusters

Rodd, Revision of Livistona (Arecaceae)

on a rachilla ever bear a fruit. The compound fruit stalk is commonly up to double the
length of its combined elements at anthesis.

Colour of the ripe fruits appears to be predominantly black or brownish-black in
Australian Livistona. The china-blue colour seen in some Asiatic species such as
L. chinensis or L. saribus does not occur, nor the bright red of (immature?) L. rotundifolia.
The ripe colour of some of the northern species is not yet known with certainty, in
particular that of L. victoriae, of which only immature or very old fallen fruits have
been collected, all of which are brown, though the label notes of one collection suggest
they may ripen to black. Other surface characters of the epicarp may be significant, for
example presence or absence of a glaucous bloom, a high gloss as a opposed to a matt
surface (usually minutely wrinkled), and the presence in some species of small corky
dots or pore-like structures (apparently confined to the larger-fruited species). There is
some variation in epicarp texture, some species having very thin, delicate skin while
in others it is thicker and tougher.

The mesocarp exhibits a range of useful features. Generally the fresh colour and
texture have not been recorded, though there is a strong suggestion that species fall
into two broad classes of 'thick and juicy' and 'thin and fatty' (but with some
intermediate). Characters that are well preserved in the dry state include the
abundance, size and arrangement of fibres or stone-cells; presence/abundance of
reddish-brown tannin-bodies (possibly an artefact of drying); and texture, colour
and/or patterning of inner face of mesocarp. Thickness of mesocarp in the dry state
has been measured for each species and appears constant within certain limits.

The endocarp of most Livistona species is straw-coloured, thin, brittle and somewhat
granular in texture, but in a few species it is somewhat tougher with a springy texture.
There is some variation in the degree to which it remains adherent to the mesocarp
after ripening and drying. When examined under magnification its inner or surface
may show some patterning, which in a few cases seems to be characteristic of a
species, though there is a tendency for the any patterning of the inner surface to reflect
that of the seed surface. Endocarp thickness has been measured for all species of which
ripe fruit was available.

6. Seed

Seed morphology has been much emphasised by past workers, though the basic
structure of the seed is remarkably uniform in Australian Livistona. Its shape varies
from subspherical, usually somewhat flattened at the base of the seed coat intrusion,
to moderately ellipsoid in those species with elongated fruit, though the
length:diameter ratio is never more than 1.5:1. Tire hilum and raphe are inconspicuous
in all but the largest-fruited species, but in nearly all species there is a more or less
circular patch on the ventral side that is shinier and darker brown than the rest of the
seed surface — this corresponds to the base of the intrusion of dark-coloured seed coat
into the whitish endosperm, which characteristically takes the form of a cylinder
expanded to varying degree at its apex into a somewhat fungoid shape, and always
lying directly opposite the embryo position (Fig. 2a). The shape, size and orientation
of the intrusion can only be discovered by splitting or cutting the seed in both
transverse and longitudinal planes, the latter through the embryo position. Because of
the difficulty of detecting the raphe this is not always easy in practice and in most
cases is best done with sharp secateurs on a ripe but fairly fresh (or half-dried) fruit in
which the mesocarp, endocarp and seed are still adherent to one another, using the
stalk position and any hint of a ventral suture as guides for orienting the cuts.

Features of the seed coat intrusion have been described in detail in the species accounts.
There is a strong correlation between the seed size and the relative volume and

Telopea 8(1): 1998

complexity of shape of the intrusion, which is doubtless explainable in terms of seed
physiology (exchange of gases or solutes from endosperm?). Its dimensions have been
expressed in terms of how far toward the opposite seed wall it extends (as a
proportion of the dorsiventral diameter), and what proportion of the endosperm its
width occupies (i.e. of the lateral diameter), and likewise what proportion of the seed
length it occupies, all as approximate fractions, e.g. %. But another feature of
significance is the position from which the intrusion penetrates: in some species this is
from a median position relative to the fruit stalk (i.e. at 9 o'clock if the fruit stalk is
taken as 6 o'clock), while in others it may be from closer to the base or the apex
(e.g. 10 o'clock or 7 o'clock). The embryo, which lies opposite the intrusion in a small
pocket in the endosperm, varies in position in a similar way, generally between the
2 o'clock and 5 o'clock positions relative to the fruit stalk. These features of intrusion
and embryo position appear to be constant for each species within quite narrow limits.

7. Evidence of dioecy (or androdioecy)

Uhl and Dransfield (1987) describe Livistona as 'hermaphroditic (rarely dioecious)'.
Some of the Australian species display clear signs of sexual differentiation among the
palms of a population, although the nature of this sexuality is still unclear. It finds its
strongest expression in L. humilis, which conveniently is also the species in which it is
most easily observed, flowering as it often does at only 1-2 m above the ground.
Observation of populations of this species reveals a striking dimorphism of
inflorescence structure among plants, with approximately half of them producing very
elongated inflorescences with only a single, terminal partial inflorescence, lateral
branching being suppressed and the rachis clothed in a long series of sterile bracts.
The remaining plants have an inflorescence structure conforming to the normal
Livistona pattern, with each rachis bract subtending a partial inflorescence. It was my
observation that only inflorescences of the first type bear fruit, and that each plant
consistently produces only one type of inflorescence, evidenced by the previous years'
inflorescences either fallen on the ground or persisting on the palm. Examination of
flowers found on the two inflorescence types reveals that the 'female' (fruit-bearing)
plants have flowers with apparently empty anther locules, though in other respects
more or less identical to flowers on 'male' plants, the stamens being of normal form.
Flowers of the 'male' plants appear to have have their full complement of stamens and
carpels, the latter containing apparent ovules.

The other species for which my observations clearly support sexual differentiation is
L. australis, readily observable in large populations in the fringes of the Sydney
metropolitan area. Although this species produces inflorescences of similar overall
structure on all individuals, many palms consistently fail to bear fruit, evidenced by
the fallen old inflorescences on which none of the rachillae have thickened fruit-stalk
remains, hr contrast, fruiting individuals consistently accumulate beneath them old
inflorescences with thickened fruit-stalk remains, as well as having thicker, more
decay-resistant rachillae, doubtless a consequence of the conducting and mechanical
tissues that develop post-anthesis to support the masses of fruit. At flowering time,
normally early to mid-August in the Sydney area, some trees can also be seen bearing
ripe fruit from the previous year's flowering, but showing no sign of flowering in the
current season. This observation lends partial support to that of Orscheg and Parsons
(1996b), who postulate an 18-month interval between flowering and fruit fall in the
southernmost stand of L. australis, and at least 2 years between successive flowerings
of individual trees. They make no mention of sexual differentiation among trees. In the
Sydney populations observed by me, there are slight but consistent differences
between fruiting and non-fruiting trees in arrangement of flowers on rachillae, and in
texture and indumentum of rachis bracts: the 'male' rachillae have closely crowded

Rodd, Revision of Livistona (Arecaceae)

flower clusters that are predominantly 3—4-flowered, while 'female' rachillae have
more widely spaced clusters that are predominantly 1-2-flowered; 'male' rachis bracts
are papery with dense patches of tangled scales near their apices, while 'female' bracts
are more leathery and with much sparser scales. Flowers of both kinds appear to have
fully developed stamens and carpels. In the description of L. australis below, no
attempt has been made to separate character-states of the two 'sexes', since for most
flowering collections there is no record of whether or not the tree bears fruit.

Brief observations during some of my collecting trips suggest that several other
species exhibit some degree of functional sexual differentiation: these include
L. alfredii, L. eastonii and L.fulva. It is possible, indeed, that this is a feature of all the
Australian species, or even of the genus as a whole. It remains an interesting area for
further study.

Relationships of the Australian Species

Among the 16 Australian species recognised here, no clear-cut pattern of affinities is
readily discernible, except that several pairs of allied species can be identified, notably
L. australis-L. nitida; L. mariae-L. lanuginosa; and L. lorophylla-L. kimberleyana. As
previously noted, all the Australian species except L. imielleri and L. benthamii are
endemic and even these do not extend beyond the Papuan lowlands, part of the
Australian plate. Present knowledge does not allow any confident statement as to
affinities between the Australian and other extra-Australian species, but neither is
there any strong evidence to contradict a tentative hypothesis that the Australian and
the extra-Australian species belong to distinct clades, or at least that there may be
several clades, of which none is shared between Australia (including the Papuan
lowland) and other regions.

Only the most tentative groupings have been attempted among the Australian species,
and it is quite likely that some of the features holding their component species
together are the result of convergence in similar environments. Four informally
designated groups are outlined below

A. 'Mariae Group' -— stout-trunked palms of alluvial flood-channels and soaks in
semi-arid environments, fronds large with thick ribs and thick wax layer on
underside, inflorescences long, stout and rigid with numerous partial inflorescences.
An apparently natural group consisting of species 1. L. mariae and 2. L. lanuginosa.

B. 'East Coast Group' — green-leaved, mesomorphic palms of rainforest margins, wet
sclerophyll forest, swamp-forest and river channels. A probably heterogeneous group,
having in common deep green leaves with drooping segments, and small to medium¬
sized, globose to subpyriform black fruit. Consists of species 3. L. benthamii, 4. L. drudei,
5. L. australis, 6. L. nitida, 7. L. decipiens, and 8. L. fulva (this last species is the most
anomalous in this group, the others all having glossier leaves, glabrous beneath and
with deeply bilobed segments; it may have affinities with species 9 of the
following group).

C. 'Arafura Group' — small to medium-sized palms forming very extensive stands on
deep sandy or lateritic interfluves in strongly monsoonal environments of far northern
Australia (extending to Papuan lowlands in the case of L. muelleri). They appear to
form a natural group, characterised by smallish fronds with stiff, rather shallowly
bilobed segments, the petioles with an indumentum of persistent, crustose basal-
masses of scales, arranged in lines; fruits are smallish, blue-black, slightly to strongly
ellipsoid or pyriform. Consists of species 9. L. muelleri, 10. L. humilis, and 11. L. eastonii.

D. 'North-western Group'— small to moderately large palms of cliff-ledges, ravines
and banks of non-perennial streams in semi-arid environments, the leaves deeply

Telopea 8(1): 1998

dissected with deeply bilobed segments, bluish or greyish-green with a hard, waxy
cuticle, inflorescences with rather few (3—8) partial inflorescences, rachis bracts early-
necrotic and parchment-like. All are from the ranges of the Kimberley and Pilbara
Regions of Western Australia except L. inermis from rocky escarpments of the Northern
Territory 'Top End'. Possibly a heterogeneous group, consisting of species 12. L. inermis,
13. L. lorophylla, 14. L. kimberleyana, 15. L. victoriae, and 16. L. alfredii.,

Notes on the Key

• All leaf characters used apply only to adult leaves, defined here as leaves produced
after the first flowering.

• Division of the lamina is measured on the widest segments, usually those between
4 and 8 segments away from the costa apex on either side

• Lamina length is measured from base of hastula to the longest segment apices

• Petiole width is measured just below the hastula; other petiole features such as t.s.,
color and indumentum are also taken to be at this point

Key to the Australian species of Livistona

1 Leaf blade very deeply divided leaving a central undivided portion not exceeding
one tenth of total length (measured along broadest segments, at c. 2 o'clock position
relative to apex of costa)

2 Leaves deep green or somewhat brownish green, more or less glossy at least on
upper side, strikingly costapalmate with long pinnatisect central portion, costa
conspicuous from above as well as from beneath

3 Small slender palm with trunk rarely over 6 m high or 10 cm diameter; leaf

blade under 80 cm long with 50 or fewer segments, these under 15 mm wide
below bifurcation and bifurcated to c. % their free length; inflorescence with
5-6 partial inflorescences; flowers in sessile clusters; fruits obovate, under
10 mm long. 13. L. lorophylla

3* Robust palm, trunk to 12 m or more high, 20-25 cm diameter; leaf blade
120 cm or more long with over 60 segments, these over 18 mm wide below
bifurcation and bifurcated for about half their free length; inflorescence with
7-12 partial inflorescences; flower clusters with cluster axis to 5 mm long;
fruits globose, mostly 12 mm or more in diameter . 7. L. decipiens

2* Leaves not shiny, pale greyish green on both surfaces with hard wax-
impregnated cuticle, only moderately costapalmate with costa short and
requiring close examination to detect it from above

4 Very slender palm, sometimes multi-stemmed, with trunk mostly under 10 m
tall and under 10 cm diameter; leaf blade mostly under 70 cm long with
segments under 15 mm wide below bifurcation; inflorescence under 80 cm
long with only 2-3 partial inflorescences, the lowest subequal to the main axis

... 12. L. inermis

4* Palm to 15 m or more tall, trunk c. 15 cm diameter, always solitary; leaf blade
mostly 80-100 cm long with segments 20-35 mm wide; inflorescence over
100 cm long with c. 8 partial inflorescences. 14. L. kimberleyana

1* Leaf blade divided less deeply, leaving a central undivided portion always greater
than one tenth of total length

Rodd, Revision of Livistona (Arecaceae)

5 Leaf blade small to medium-sized, seldom exceeding 100 cm in length; segment
lobe apices not long-attenuate and not or only slightly drooping

6 Leaf segments bifurcated to at least half of their free length, the lobes long-
attenuate and concurrent; blade with broad basal notch (120-200 degrees);
fruits usually markedly longer than wide; trunk under 15 cm diameter,
generally remaining clothed in old petiole stubs

7 Trunk exceeding 6 m with age, over 10 cm diameter, abruptly flared out

into a bulblike base, covered in closely appressed petiole stubs in a
strongly geometric pattern; leaf blade usually over 60 cm long, glaucous
when young, with more than 40 segments; inflorescences not strongly
dimorphic (on different plants), approximately equalling leaves;
rachillae glabrous... U- L. eastonii

7* Trunk rarely attaining 6 m, under 10 cm diameter, only slightly flared at
base, petiole stubs persisting unevenly, not neatly geometric; leaf blade
under 55 cm long, never glaucous, with fewer than 40 segments;
inflorescences markedly dimorphic, those on fruiting plants c. twice leaf
length with lateral branching suppressed and only a small terminal
partial inflorescence, those on non-fruiting plants c. 114 times leaf length
with 4 or more partial inflorescences; rachillae bristly-pubescent with
hairs c. 0.1 mm long . 10. L. humilis

6* Leaf segments shallowly bifurcated, lobes mostly under 5 cm long; blade
with very narrow basal notch (mostly under 60 degrees), appearing circular
in outline; fruits globose or only slightly ellipsoid; trunk mostly over 15 cm
diameter, generally free of old petiole stubs

8 Leaf blade c. 100 cm long, underside thinly glaucous and clothed in short

golden-brown appressed hairs at least when young; segments bifurcated
to a depth of c. 2-2.5 cm into rigid, converging points; inflorescences
shorter than petioles, the tubular bracts of main axis rather loose, grey-
brown, their apices irregularly laciniate (not 2-lobed); fruits globose,
over 13 mm diameter . 8. L.fulva

8* Leaf blade c. 70 cm long, underside thinly glaucous, glabrous except for
whitish scurfy scales on ribs; segments bifurcated to a depth of up to
5 cm into rigid parallel points; inflorescences equalling or slightly
exceeding petioles, the tubular bracts of main axis rich chestnut-brown,
their apices acutely to acuminately 2-lobed; fruits ovoid-ellipsoid, under
13 mm long ....... 9. L. muelleri

5* Leaf blade medium-sized to large, if under 100 cm in length then segment lobe

apices long-attenuate and at least moderately drooping

9 Leaf blade pale greyish-green to quite glaucous above

10 Leaf blade grey-glaucous on both sides with hard wax-impregnated
cuticle; inflorescence rachis bracts glabrous to sparsely scaly

11 Trunk c. 30 cm diameter at 1 m above ground; partial inflorescences
branched to 2 or 3 further orders, rachillae mostly over 4 cm long
. 16. L. alfredii

11* Trunk c. 20 cm diameter at 1 m above ground; partial inflorescences

branched to 4 further orders, rachillae mostly under 3 cm long .

. 15. L. victoriae

Telopea 8(1): 1998

10* Leaf blade grey-green to somewhat yellowish-green above, thickly
pruinose below with layer of soft, delicate wax; inflorescence rachis
bracts mostly clothed with dense white, woolly scales .

. 2. L. lanuginosa

9* Leaf blade glossy mid- to deep green above

12 Leaf blade strongly pruinose beneath with layer of soft, delicate wax;
inflorescence axis straight and ascending, bearing 10 or more partial
inflorescences . 1. L. marine

12* Leaf blade not pruinose or glaucous beneath, more or less concolorous
though usually less glossy beneath

13 Widest primary leaf segments mostly less than 30 mm wide below
point of bifurcation; fruit slightly pyriform, under 12 mm diameter

14 Trunk (of mature palm) smooth toward base without persistent
petiole stubs; leaf blade mostly over 100 cm long, drying dull
greyish-brown above, paler brown beneath; inflorescence rachis

bracts coarsely striate with regularly cuspidate apices.

. 4. L. drudei

14* Trunk with persistent, widely diverging petiole stubs toward
base (unless exposed to fires); leaf blade seldom more than 100
cm long, drying to somewhat glossy pinkish-brown above
contrasting with olive brown beneath; inflorescence rachis bracts
finely and obscurely striate with narrowly triangular apices
... 3. L. benthamii

13* Widest primary leaf segments mostly more than 30 mm wide below
point of bifurcation; fruit quite spherical, over 12 mm diameter

15 Leaf blades mostly under 140 cm long, the segments bifurcated

for little more than half their free length; fruits dull blackish
often with brownish or reddish tones; mesocarp sparsely fibrous,
inner face of endocarp not reticulate. 5. L. australis

15* Leaf blades mostly over 140 cm long, the segments bifurcated for
about two-thirds their free length; fruits glossy black; inner layer
of mesocarp densely fibrous, inner face of endocarp reticulately
patterned . 6. L. nitida

Treatment of species

1. L. mariae F. Muell.

Mueller (1874: [221], 283; 1875: 222; 1878: 55; 1892: 112); Drude (1893: 39); Dammer
(1905: 297); Beccari (1921: 19; 1931: 92); Lothian (1958: 92); Moore (1963: 150); Latz
(1975: 189).

Type citation: 'in valle "Glen of Palms" montium Gillii; Emestus Giles'.

Type: Corresponding to this citation are two sheets in MEL — the first is a whole leaf,
roughly folded, labelled 'Gills Range, Giles'; the second consists of a few segments
only, labelled 'Gills (Macdonnells) Range, E. Giles'. The former of these I here
designate as lectotype of L. mariae. [Mueller's mixed concept of 1878 is discussed
under Note 1, below, and also under L. alfredii ]

Saribtis mariae (F. Muell.) O. Kuntze, Rev. Gen. PI. 2: 736 (1891).

Rodd, Revision of Livistona (Arecaceae)

Trunk solitary, straight or frequently curved or kinked, to c. 28 m high, c. 40 cm
diameter at 1.5 m above ground, tapering gradually but perceptibly upward to a final
constant diameter of c. 30 cm, broadening below into a gently bulbous base c. 50 cm
diameter sitting on a pedestal of densely massed roots sometimes 50 cm or more high.
Surface moderately smooth but with stepped sheath scars, more strongly stepped
petiole scars; vertical fissures numerous, some short and closed, others long, deep and
gaping; petiole-stubs present mainly at extreme base, short, semi-appressed.

Crown moderately open, globose to somewhat umbrella-shaped, consisting of c. 35-50
spreading to drooping massive leaves, their blades extending in same plane as petioles
but with pendulous or at least nodding segment-lobes. Ligules prominent, pale
straw-coloured aging to dirty white, bearing dense shaggy whitish scales.

Petiole 150-260 cm long, 20—45 mm wide, triangular in t.s. with rounded keel, slightly
concave above sometimes with 1 or 2 slight raised ridges. Margins minutely and
irregularly denticulate in apical 'A to %, with frequent small dark brown calli, some
projecting, others slightly depressed; toward base armed with blackish, blunt, patent
prickles to c. 3 mm long. Surfaces yellowish-green, the upper with very crowded,
deep, subparallel, linear pits, up to 10 mm long, most abundant on median band of
petiole, on younger leaves obscured by dense shaggy white scales, these
semi-appressed, mostly with elongated bases, laciniately dissected, the laciniae
narrowly straplike, c. 5 mm long, mostly straight but helically twisted and apically
tufted; older petioles with smaller scale-remnants persisting; lower surface similar but
pits shallower, sparser; more sparsely scaly, scales densest close to margins, laciniae
mostly under 3 mm. Hastula flattish; base shallowly 3-lobed, the central lobe longest;
rim at c. 30° to costa, to c. 15 mm wide, ± triangular, acute, with disintegrating necrotic
margin.

Lamina 130-220 cm long, c. 0.3 mm thick, moderately to strongly costapalmate with
broad basal sinus, fairly rigid but somewhat brittle, easily split; slightly to moderately
contorted with 1 gentle adaxial undulation either side of the slightly deflexed costa
and base on either side slightly involute. Segments 37—43 either side of costa; largest
segments 38-45 mm wide, free for 45-55% of their length, bifurcated for 45-65% of free
length, the lobes parallel to slightly diverging, tapering into fine but not aristate or
threadlike apices, slightly bristly, early-necrotic, mostly broken-off and split,
lntersegmental appendages present, to c. 30 cm long, moderately thick and tough,
bearing short woolly scales Ribs: abaxial ribs large and prominent, in t.s. oblong to
trapeziform (narrowing to lamina), sharply square-edged or slightly rounded on one
side, 0.8-1.2 mm thick, 1.8-2.5 mm deep; adaxial ribs more narrowly oblong, c. 0.7 mm
thick, 1.5-2.5 mm deep. Venation: major longitudinal veins 7-9 either side of abaxial
rib, moderately prominent on both surfaces; transverse veins evident but not
prominent above, moderately prominent beneath, rather widely spaced, mostly
near-orthogonal with respect to longitudinal veins, mostly discontinuous, generally
sigmoid or more finely sinuous. Surfaces: upper pale to deep glossy green, variably
glaucous with delicate waxy bloom, glabrous; lower dull pale green with coating of
thick, soft wax, glabrous except for whitish scales on bases of ribs.

Inflorescences mostly shorter than petioles, though sometimes approaching 2.5 m in
length, very stiff and straight. Partial inflorescences c. 10-14, subequal, short relative
to rachis, stiff and forward-pointing to moderately recurved, each branched to 4
further orders; rachillae 3-8 cm long, c. 0.8-1.0 mm thick, glabrous, irregularly striate,
minutely white-papillose. Rachis bracts cylindrical or slightly flattened, larger ones
4-5 cm diameter, tightly sheathing, smooth or with few coarse transverse wrinkles,
rigidly coriaceous, remaining entire and generally untorn through to fruit, pale green
aging yellowish, sparsely to densely covered in shaggy white loosely appressed scales,
these deeply laciniate with straight, helically twisted laciniae to c. 6 mm long, or
subentire and strongly undulate, tufted at apices; bract-apices ovate-acuminate,
sometimes drawn out into a long slender point. Flower-clusters 0.5-2 mm apart,

Telopea 8(1): 1998

3-6-flowered; cluster axis to 1.3 mm long, fleshy, arcuately deflexed with flowers
tightly crowded secundly on adaxial (outer) side. Cluster-bract short and broad, c. 0.3 mm
long, lacerately toothed or entire and broadly acute, membranous, hidden beneath
cluster axis. Bracteoles 1(?) per flower, very short, crowded.

Flowers 1.5-1.8 mm long, globose-campanulate. Anthopodium c. 0.15 mm long,
minutely recessed at base with sharp narrow rim, minutely trigonous. Sepals closely
appressed to petals, strongly concave, membranous, 0.8-1.0 mm long, connate for
c. A their length with rather narrow, angular sinuses, free portion ovate, broadly acute
to somewhat rounded, sometimes awned with one or more delicate hairlike laciniae
c. 0.3 mm long. Petals fleshy, strongly concave, 1.2-1.5 mm long, connate for c. Ko their
length, suborbicular to broadly oblong, broadly acute with strongly thickened apex,
auriculate at base, the auricles minute, retrorse, narrow; inner face with 3 deep
cavities. Stamens c. % as long as petals, connate for c. % their length; filaments very
small, rather membranous, triangular-acuminate, distinctly shouldered a little above
base, thence tapering very smoothly into a slender apex; anthers c. 0.4 mm long, about
the same wide. Carpels from % as long as to subequal to stamens; ovaries
globose-obov'oid, strongly gibbous abaxially, gradually to abruptly contracting into
styles 0.2-0.3 mm long.

Fruit ± spherical to slightly flattened, 13-17 mm diameter; suture sometimes evident;
base quite rounded or with very slight conical protrusion, Epicarp glossy black or with
patches of mahogany-brown, with fairly conspicuous scattered pale brown spots
0.2-0.5 mm diameter, some with slits or pores. Mesocarp 2-2.5 mm thick when fresh
(c. 1.5 mm dry), purplish green, thick and fleshy, fairly soft; granular in texture when
dry ; containing thick, fairly crowded, moderately straight fibres largely confined to
inner half of mesocarp and very prominent on its inner face in old fallen fruits.
Endocarp 0.2-0.25 mm thick, straw-coloured, brittle, adhering fairly closely to
mesocarp; inner surface smooth with conspicuous regular pattern of very close, pale
orange-brown dots, outer surface sometimes deeper orange-brown, obscurely
verruculose with a similar patterned appearance. Seed markedly flattened ventrally,
c. 7.5-12 mm long and wide; surface pale grey-brown, dull and minutely rough with
very fine pattern of spots and radiating whitish cells except for shiny deeper brown
hilum. Intrusion short and broad, penetrating transversely from slightly above median
position for % seed diameter, in l.s. with broadly flared base, usually constricted at
middle, shallowly to deeply 2-lobed or unlobed, usually lumpy in outline, occupying
A or slightly more of seed length; in t.s. less flared at base, more markedly 2-lobed,
occupying less than 'A of seed width. Embryo sub-basal (c. 5 o'clock relative to stalk).

Distribution: see under subspecies below.

Ecology: essentially a palm of stream-channels, associated with permanently flowing
water or permanent pools or soaks; also in rocky beds of sandstone gorges or
occasionally on lower ledges of cliffs, but in these situations always on permanent
soaks. It is associated with a variety of vegetation types, e.g. fringing forest/woodland
of Eucalyptus camalduletisis in Central Australia, gallery-forest of Melaleuca leucadendra,
Nauclea orientalis, Ficus racemosa, Pandamts aquaticus in northwest Queensland and the
Northern Territory. In the Durack Range it was observed in close association with
Livistona kimberleyana while at Kapalga in Arnhem Land it was reported to occur with
L. benthamii.

Notes

1. Mueller first referred to this species without name in 1874 in a brief paragraph
appended to his description of Livistona leichhardtii. He postulated that Giles's
collection from the 'Glen of Palms' (consisting of a single leaf without petiole) might
possibly be conspecific with his L. leichhardtii (now regarded as an illegitimate name);

Rodd, Revision of Livistona (Arecaceae)

he refers to its characters in the most sketchy way, viz., 'sub hastis clavisque barbarorum
segmenta usque 4' longa exhibet'.

2. Mueller's next mention of this palm was in the 'Additamenta' of the same volume
of the Fragmenta, also issued in 1874, in the form of a brief statement: 'Palma Gilesii
huic non conspecifica, sed fortasse congenerica, et tunc L. Mariae dicanda'. Chapman
(1991) treats this as the place of publication of the name Livistona marine, commenting
'Combination made by reference to 8: 221...'. Chapman's acceptance of this as place of
publication is endorsed here, even though Mueller's diagnosis rather doubtfully
meets the requirements of the present Code.

3. By the time Mueller came to publish a full botanical account in the Fragmenta,
another occurrence of a Livistona had been discovered by Forrest, also distantly
isolated from any other palms, and Mueller concluded (wrongly, as he later admitted)
that it represented the same species.

4. The more ample 1878 description of L. mariae in Mueller's Fragmenta was based on
Giles's leaf specimen and flowers and fruits collected by Forrest in northwestern
Australia. Mueller acknowledged these were unconnected but nonetheless concluded
'sed plantam ex utraque regione conspecificam arbitror'. As early as 1879 he was sent
fruits from the Palm Valley Livistona by the missionary Kempe, and his growing
doubts culminated in his publication of the 1892 note in the Victorian Naturalist,
generally regarded as the protologue of L. alfredii (see also notes under that species).
If Mueller's 1874 mention of L. mariae is not regarded as valid publication of the name,
then this 1878 description would have to be regarded as the protologue. But that
would leave unresolved the problem of typification of the name L. mariae, due to the
mixture of elements on which this later description is based, which is a pragmatic
argument for allowing the 1874 publication of the name.

5. Chapman's (1991) note under L. mariae, 'Also published by Mueller in Gard. Chron.
(June 1879) 790/ is misleading, in that this article has no descriptive material except
for: 'Von Muller hopes shortly to succeed in introducing Livistona Mariae into Europe,
where its peculiar pale green colour will contrast well with its more highly coloured
eastern congeners.' Moreover he appears to have overlooked Mueller's 1878
description (see previous paragraph).

6. Subsequent authors such as Drude (1893) and Beccari (1921, 1931), following
Mueller's 1892 note, interpreted L. mariae in the present sense, firmly establishing its
application to the Central Australian palm. C.A. Gardner (1923), however, seemed
totally confused, applying L. mariae to the Fortescue River palm and L. alfredii to a
Kimberley palm (see notes under L. alfredii and L. kimberleyana).

7. Beccari (1921, 1931) failed to perceive any resemblance between his L. rigida and
L. mariae, perhaps understandably in view of the fragmentary material he had
available of both. It seems strange nonetheless for him to have speculated that his new
species might prove identical with Brown's L. inermis, a species far smaller in leaf
dimensions, which in fact he described quite accurately; perhaps he was too
influenced by their geographic proximity.

8. Tire decision here (see below) to treat the Central Australian palms as conspecific
with the various northern Australian populations is perhaps somewhat controversial.
However, although there are slight but apparently consistent differences in leaves,
rachis bracts and fruits, the resemblances in detail in most organs are so striking as to
suggest overwhelmingly that this is all the one species. Unfortunately this conclusion
is at present based on very inadequate collections, and a clear picture of variation must
await comprehensive sampling of adult leaves, flowers and mature fruits from all
populations. In particular the Western Australian plants are very poorly known.

Telopea 8(1): 1998

But, as with any disjunct occurrences of slightly differing plants whose relationship is
clearly one of vicariance, the decision to separate or combine them at species level
must in the end be arbitrary.

9. The most useful solution at this stage would seem to be to treat the entities that can
be detected within L. marine s. lat. as subspecies. The following subspecific classification
is therefore proposed.

Key to Subspecies

1 Adult leaf blade mostly exceeding 2 m, segments drooping, bifurcated for about %

of their length; upper surface glossy green; fruit c. 15 mm diameter.

.. subsp. marine

1* Adult leaf blade mostly under 1.8 m, segments stiff or slightly drooping, bifurcated
for about A of their length; upper surface thinly glaucous, fruit c. 13 mm or less
diameter

2 Rachis bracts with moderately dense scales especially toward apices .

. subsp. rigida

2* Rachis bracts virtually glabrous except for a few sparsely scattered white scales
near apex . subsp. occidentals

subsp. mariae

The type subspecies is characterised in the field by its greener, more glossy upper leaf
surfaces and longer, more drooping leaf segments. Table 3 details the comparative leaf
dimensions and ratios of this and the other two subspecies. The rachis bracts are rather
densely clothed with persistent white scales, though usually with some relatively bare
areas especially on the lower part of the bract. Ripe fruits are 13-16 mm in diameter,
virtually spherical and glossy black; seeds are 10.5-12.0 mm long (the smallest
diameter may be under 10 mm). (Fig. 3a, 5a)

Distribution: Central Australia: Restricted to Palm Valley, on the Finke River where it
cuts through the Macdonnell Range, west of Alice Springs. A relict occurrence of only
about 1500 adult palms, about 12 km separating its furthest limits (Latz 1975).

Conservation status: 2VCa (Briggs & Leigh 1996)

Specimens examined: Northern Territory: Central South: Gills (Macdonnells) Range, Giles s.n.
(MEL); Palm Valley, Mitchell 84, 25 Sep 1974 (DNA); Palm Valley, beside Palm Creek near ruins of
Chalet, Rodd 3215 & Jacobs, 29 May 1976 (NSW, DNA); Palm Valley, Lothian 315 ,1954 (AD). Palm
Valley, Cleland s.n., 17 Aug 1929 (AD).

subsp. rigida (Becc.) A.N. Rodd, comb. nov.

Livistona rigida Becc., Webbia 5(1): 19 (1921).

Type citation: 'Albert River nel Golfo di Carpentaria (Erb. di Kew). Collettore ignoto.'

Type: Albert River, Ferd. Mueller; holo K. [Mueller may have distributed the specimen
from MEL and is not necessarily the collector.]

Differs from subspecies mariae in its slightly shorter leaves with shorter bifurcation of
segments, more glaucous upper leaf surface, less dense indumentum of shaggy white
scales on rachis bracts, and slightly smaller fruit and seed. Schneider s.n. has fallen
mature fruit 14 mm long, 12-13 mm wide. Seed 10 mm diam; inner mesocarp with
conspicuous dense straight fibres. For differences from subsp. occidentalis see under
that subspecies. (Fig. 3b, 5b-c, 6)

Rodd, Revision of Livistona (Arecaceae)

Table 3. Comparison of selected characters of subspecies of L. mariae.

Note: Leaf dimensions are based largely on my own collections Rodd 2868, 2937 and 3275, as
none of the other available collections have undoubtedly adult leaves.

subspecies

mariae

lamina

c. 220 cm

length

depth of

c. 120 cm

primary

(55%)

division

depth of

c. 75 cm

bifurcation

(63%)

upper

glossy

surface

main

densely

inflorescence

scaly

bracts

fruit

13-16 mm

length

seed

10.5-12.0 mm

length

rigida

occidentalis

c. 160 cm

c. 175 cm

c. 80 cm

c. 40cm

(50%)

(48%)

c. 40 cm

c. 40cm

(50%)

(48%)

glaucous

less densely

almost

scaly

glabrous

14 mm

unknown

10 mm

unknown

Distribution: this subspecies is known from several disjunct areas: 1. far northwest
Queensland, along sections of the Albert, Nicholson, Gregory and Leichhardt Rivers
and Lawn Hill Creek, also reportedly on coastal sands opposite Mornington Island
this is the most extensive occurrence, extending southeast almost to Mount Isa; 2.
upper Roper River, Northern Territory, in a small area close to Mataranka; 3. Kapalga,
near the East Alligator River, Arnhem Land; and 4. Upper Goyder River in
northeastern Arnhem Land — the last two, each known from a single collection, are
about 2 degrees of latitude north of Mataranka; the intervening region may not have
been adequately investigated for palms.

Conservation status: none assigned by Briggs and Leigh (1996); 3RCa is suggested.

Specimens examined: Queensland: Burke: Babbling Brook, Lawn Hill Stn, Lutz 1619, 24 July 1971
(DNA, CANB); Lawn Hill Creek, Rodd 2937, 2938, Dec 1974 (NSW, BRI, K); Lawn Hill Creek, SW
of Burketown, Gittins, June 1963 (BRI); Lake Julius Dam below dam wall, [collector?], 12 Feb 1977
(BRI).

Northern Territory: Darwin and Gulf: Kapalga, Dunlop 5052 & McKean, 14 Feb 1979, (DNA,
CANB, NSW); Goyder River, Dunlop 7259,2 Nov 1987 (DNA, NSW); Roper Highway, 20 k E Stuart
Highway, Latz 7828, 19 June 1978 (DNA, NSW); Warlock Ponds Bridge, S of Mataranka, Sijnum
10358, 8 May 1975 (NSW); Mataranka, Maloney s.n., 13 July 1975 (NSW); Mataranka Station, Salt
Creek, Schneider s.n., 5 July 1964 (AD).

subsp. occidentalis Rodd, subsp. nov.

Differt a subsp. mariae et subsp. rigida bracteis majoribus inflorescentiae fere glabris.

Type: Western Australia: SE base of Mt King, Durack Range, Rodd 2868, Dec 1974; holo
NSW; iso PERTH, K, BH.

Telopea 8(1): 1998

The only character so far known in which this differs from both the other subspecies
is its almost glabrous rachis bracts. In leaf characters it matches subsp. rigida quite
closely, notably in its overall blade length, depth of dissection, length of secondary
lobes, and moderately glaucous upper surface of lamina, in all of which these two
subspecies differ from subsp. marine. Comparisons of flowers and mature fruits are not
yet possible, due to lack of material. Recognition of this population as a separate
subspecies on the basis of a single wild collection (and one from cultivation) is
arguably rather premature, but represents a middle course between simply
submerging it in subsp. rigida and treating it as a distinct species. (Fig. 3c, 5d)

Distribution: as so far known, this subspecies occurs in the gorges of the Durack and
King Leopold Ranges, central Kimberley region. Western Australia. In the Durack
Range it has been collected at Mt King and near El Questro Homestead, and seen from
air in gorges between these points. In the King Leopold Ranges it has been collected at
Mt Gladys, and apparently similar palms are reported by Dr Peter Wilson (NSW) in
Brownrigg Gorge near the very head of the Lennard River in the King Leopold Range.

Conservation status: none assigned by Briggs and Leigh (1996); a coding of 3VCi is
suggested.

Specimens examined: Western Australia: Bens Springs, 3 km E of El Questro Homestead, Kenneally
10965, 29 June 1989 (PERTH).

Cultivated: Queensland: Townsville Palmetum, Dowe 117, 9 Dec 1993 [ex seed from Broome
Botanical Society: Mt Gladys, Leopold Range, T. Willing s.n., May 1987] (NSW).

2. L. lanuginosa AN. Rodd, sp. nov.

Palma magna affinis L. marine habitu similique: folia valde undulata costa elongata
valde deflexa; lamina profunde dissecta, griseo-viridia, infra velamine crasso cerae,
petiolo supra squamas densas albo-lanuginosas ferenti. Inflorescentiae foliis breviores,
axe principali recto crassoque, bracteis vaginatis squamas albo-lanuginosas dense
ferentibus. Fructus magni, usque ad 3.5 cm diametro, atrocastanei ubi maturi, punctis
minutis suberosis sparsis ferentes, mesocarpio crasso.

Type: Queensland: Glenroy Creek, 75 km SE of Ravenswood, 20°35'S, 147°10’E, A.K.
Irvine 1912, 25 Aug 1978; holo NSW; iso QRS, BRI, BH, MEL.

[Livistona sp. 'Cape River' (Jones 1984, Dowe 1990)]

Trunk solitary, straight, to c. 15 (20?) m tall, c. 35 cm diameter at 1.5 m above ground,
broadening gradually downward and flaring widely at very base to 50-60 cm
diameter above a slight pedestal of exposed roots. Surface rather corky on lower trunk,
usually with persistent appressed petiole stubs to 20 cm long on basal 1-2 m, as well
as some step-like petiole scars; upper trunk surface very rough, with distinct annular
sheath-scars; vertical fissures numerous, shallow on lower trunk, deep and
zig-zagging on upper (some very long).

Crown ± globose or somewhat umbrella-shaped, rather open, consisting of c. 35-45
strongly ascending to horizontally spreading or slightly drooping greyish leaves with
stiff petioles and downward-curving leaf-blades with arching to pendulous segment
lobes. Ligules moderately prominent, pale straw-coloured ageing to dirty white, with
inconspicuous but fairly dense whitish scales.

Petiole 150-200 cm long, 30-35 mm wide; triangular-flattened in t.s. with rounded
keel, concave above near lamina but with slight median keel in mid-region. Margins
mostly unarmed for upper % or more, toward base armed with mostly patent (some
antrorse or retrorse), broad-based blackish prickles to 3 mm long; surfaces
yellowish-green right to base, upper obscurely striate, hardly pitted, at first densely
whitish-wooLly over most of length, with deeply laciniate, antrorsely appressed scales

Rodd, Revision of Livistona (Arecaceae)

to 5 mm long, the laciniae straight or sharply bent, spirally twisted, becoming tangled
with age, deciduous on older petioles leaving numerous minute, oval to elliptic,
pustular, hardly raised pale grey-brown basal-masses; lower surface almost glabrous,
hardly striate, with dark brown streaky markings and irregular, shallow, elongated
pits, few inconspicuous appressed white scales sometimes present in long narrow
rows either side of keel. Hastuln base V-shaped, slightly cuspidate; rim at 40-50° to
costa, to c. 12 mm wide, fairly straight and even, with narrow to broad necrotic margin.
Lamina 130-190 cm long, 0.3 mm thick, strongly costapalmate, ± truncate at base,
rigidly coriaceous, not readily splitting, strongly contorted with 3 adaxial undulations
either side of a deep median crease along the strongly decurved costa, sharply
involute either side at base with 3-4 lowermost segments resupinate. Segments 42-46
either side of costa; largest segments 36-42 mm wide, ± parallel-sided, free for 60-70%
of their length, bifurcated for 50-75% of free length, the lobes moderately diverging
and arching with pendulous tips, evenly tapering but drawn out into fine, flexible
apices which on older leaves are shortly necrotic but remain entire. Intersegmental
appendages ± persistent, threadlike but moderately tough, to 20 cm long, dark
greyish, densely white-woolly at first, glabrescent. Ribs: abaxial ribs in t.s. oblong,
narrowing slightly to lamina, square-edged, 0.9-1.3 mm thick, up to 3.0 mm deep;
adaxial ribs similar, 0.8-1.2 mm thick, 2.0-3.0 mm deep. Venation: major longitudinal
veins 7-9 either side of abaxial rib, moderately conspicuous above, evident but not
prominent beneath; transverse veins barely discernible above, moderately prominent
beneath, mostly near-orthogonal with respect to longitudinal veins, mostly arcuate or
sinuous, mostly discontinuous. Surfaces both somewhat yellowish pale green but
appearing pale bluish-grey, strongly pruinose, especially the lower which has a
persistent thick coating of white wax readily rubbing off onto fingers; glabrous except
for ribs on both sides toward base clothed in dense, moderately persistent, whitish
matted scales as on adaxial petiole surface.

Inflorescences slightly longer than petioles, ± erect but with strongly deflexed
branches, rachis stout and fairly straight. Partial inflorescences 9-12, subequal, the
largest approaching 'A of rachis in length, branched to 4 further orders; rachillae 3-12 cm
long, c. 0.8 mm thick, creamy-white, finely wrinkled-striate, glabrous; larger axes
toward base of each partial inflorescence densely white-woolly, soon glabrescent
post-anthesis, with sinuous, dendriticallv-branched hair-like scales c. 3 mm long,
leaving minute, numerous granular-papillose remnants after shedding. Rachis bracts
somewhat flattened, the largest 3.5-4 cm diameter, rigidly coriaceous, tightly
sheathing, smooth or ± buckled with transverse wrinkles, pale green with short
red-brown necrotic apical zone, conspicuously striate with prominent raised veins,
densely and persistently lanuginose, most densely toward apices, with matted
tomentum c. 2 mm thick, straw-coloured ageing greyish-white, consisting of scales
c. 5 mm long, divided right to base into dense tufts of plumose, hair-like, straight or
slightly sinuous laciniae which become curled and matted with age, the shed scales
leaving inconspicuous but dense, hardly raised, elongated basal-masses concolorous
with bract surface; bract-apices triangular-ovate, somewhat abruptly cuspidate or
acuminate with fine, soft points which soon shrivel and break off. Flower-clusters 1-3 mm
apart, 1-flowered; cluster axis vestigial, 0.2-0.3 mm long, conical, projecting from a
larger chin-like protuberance from rachilla. Cluster-bract broad-based with
narrow-triangular, acute central portion, marcescent post-anthesis. Bracteole 1,
immediately above and laterally overlapping cluster-bract.

Flowers c. 3.0 mm long, funnel-shaped. Anthopodium 0.5 mm or less long, c. 0.9 mm
wide, tapering, deeply recessed at base with a sharp rim and 3 obscure downward
bulges, completely concealing cluster axis. Sepals closely appressed to petals, very
concave basally, thick and pigmented except for narrow, thin, translucent margin,
c. 1.5 mm long, connate for almost half their length with broad, shallow sinuses,
narrowly triangular, sharply acute to finely acuminate or even aristate at apex.

Telopea 8(1): 1998

Petals very thick, tough, moderately concave, 2.8-3.0 mm long, connate for c. A their
length, rather narrowly ovate-triangular; mostly sharply acute at the strongly
thickened, barely inflexed apex auriculate at base, the auricles quite large, sometimes
broadly overlapping; inner faces with 3 long, deep cavities. Stamens % to % as long as
petals, filaments broadly oblong with rounded apex to almost semicircular in outline,
not or barely shouldered, narrowed very abruptly into short, very slender apex; anthers
c. 0.4 mm long. Carpels slightly shorter than stamens; ovaries obovate to obconical,
strongly gibbous abaxially, narrowed very abruptly into slender, frequently curved styles
c. 0.5 mm long.

Fruit ± spherical, shallowly conical at base, 33-36 mm diameter. Epicarp moderately
tough, dark purplish-brown when ripe, smooth and moderately shiny, slightly
pruinose, with sparsely but evenly scattered pale brown pustules 0.3-1.0 mm
diameter, each with small circular pore or 3-cornered slit in summit. Mesocarp to c. 4
mm thick, granular-fleshy, moderately juicy when fresh, containing numerous thick
fibres much denser toward inner face, tending to be parallel in basal half and
anastomosing in apical half, prominent on inner wall at least in fallen fruits. Endocarp
0.3-0.5 mm thick, quite hard, tough and springy, pale straw-coloured, minutely
rugulose on both faces but appearing smooth and matt-surfaced except for prickle-like
projection at stalk position. Seed subspherical, very slightly flattened on ventral face,
c. 22 mm long, 21 mm wide, 19-20 mm thick, with distinct, large raphal scar at base;
surface smooth, mottled pale and dark red- brown; seed-coat mostly c. 1.0 mm thick.
Intrusion very large, occupying most of seed volume, penetrating % or more of seed
width from a lateral position, in l.s. very broad-based but broadening into irregular,
vaguely circular outline, divided somewhat into 2 lobes, occupying % of seed length;
in t.s. similar in outline but slightly narrower, occupying slightly over 'A seed width.
Embryo large, sub-basal (4.30-5.00 o'clock relative to stalk). (Fig. 3d, 7, 9a)

Distribution: restricted to a small area of the Burdekin River basin, tropical east
Queensland, 100-200 km from the coast, between latitudes 20°30' and 21°, and
altitudes 150-300 m.

Ecology: known only along sandy river and creek channels that flow only for part of
the year but with permanent pools or soaks; associated trees are Eucalyptus
camaldulensis, Melaleuca leucadendra, and Pandanus Icookii.

Conservation status: 2V (Briggs & Leigh 1996).

Specimens examined: Queensland: North Kennedy: Glenroy Creek 75 km SE of Ravenswood,
Irvine 1913, 25 Aug 1978 (QRS, NSW); Glenroy Creek, c. 11 km N of Burdekin Falls, Rodd 3759 &
Jackes, 25 Nov 1981 (NSW); Glenroy Creek c. 3km from Glenroy Station homestead, McLain 2, Feb
1978 (QRS); Middle Burdekin Valley somewhere near Burdekin Falls, McLain s.n., 1978 (or possibly
1977) (NSW).

Cultivated: Queensland: Townsville Palmetum, Dome 111, 18 June 1993 [ex seedlings: upper
reaches of Burdekin River in vicinity of Burdekin Dam wall, R. Tucker s.n., 1986] (NSW).

Notes

1. Bailey (1902) included ‘L. marine' in his Flora, with description probably drawn
largely from those of Mueller and Bentham. Instead of citing a region or locality for its
occurrence, however, he only notes: 'I have received a portion of a leaf from F.L. Berney
of Hellenslie, Campaspe River, which in all probability belongs to this inland palm.'

2. The above record aroused my interest in the early stages of this project, in that there
had not been any other report of palms from this region of Queensland except those
much closer to the coast (L. decip'iens, L. drudei, L. australis). In 1976 a visit was made to
'Helenslee' (as it is now spelt on the map), which is near the head of the Campaspe
about 30 km east of Pentland. The manager of the property knew of no palms in the
vicinity at all, and the river bed there appeared an unlikely habitat for palms.

Rodd, Revision of Livistona (Arecaceae)

It was concluded that the specimen sent to Bailey (which has apparently not been
preserved) must have been collected from some more distant locality.

3. Quite coincidentally, in 1977, word reached palm collectors in Sydney that an
apparently quite new Livistona had been discovered in the middle-lower area of the
Burdekin, by Peter McLain, an enthusiastic young horticulturist and plant collector
from Ayr near Townsville. Seeds of the palm distributed by him were seen to be much
larger than those of any Australian Livistona except L. alfredii.

4. Tony Irvine, CSIRO botanist from Atherton with a specialist interest in palms,
followed up Peter McLain's discovery with an investigation of his own, making
collections and field observations which he very generously communicated promptly
to me. He revisited the stand known to McLain on Glenroy Creek, south of
Ravenswood not far from Burdekin Falls.

5. Tony Irvine also, following a chance remark made to him, called on Mr Jim
Rollinson of 'Nosnillor', a large property situated at the junction of the Cape and
Campaspe Rivers. Around the homestead he found a number of mature plants of this
same species, and was informed that these had been transplanted as seedlings by Mr
Rollinson from a distant part of his property on the Cape River over thirty years
previously. In September 1978 I visited 'Nosnillor' and photographed these same
plants, but was told that the area of river where they occur wild was not accessible by
vehicle. Tony Irvine was also informed by Mr Rollinson that there is another
occurrence of these palms on the property 'Harvest Home’, lower down the Cape River.

6. Tire species clearly occupies only a very limited area of the Burdekin Basin. At least
some of the known stands exhibit vigorous regeneration with most age-classes
represented, but it is possible that, in a climate marginal for palm survival, there are
periodic droughts resulting in death of all stands other than those on the most
permanent soaks in stream beds; otherwise it is difficult to understand why an
apparently so vigorous and hardy species should not be more widespread, as its seeds
are well adapted to dispersal by floods. This reasoning applies equally to the similarly
restricted L. marine and L. alfredii.

7. Dowe (1990) says of Livistona sp. 'Cape River': 'Distribution of this species is
relatively widespread, being found as far north as the tributaries [of the Burdekin]
which have their source west of Ingham; near to Greenvale in the west; to the Cape
River in the south and to Ravenswood in the south-east. The distance from north to
south is about 200 km, but this may be misleading as groups are usually very small
(this species does not form colonies as do other Livistona spp.) and are widely located.'
Dowe does not state whether this account is based on his own observation. He
concludes that its endangered status falls into the category of 'Restricted distribution
and uncommon: presently threatened', one of only two Australian palms placed by
him in this category.

8. L. lanuginosa is a very distinct species, instantly recognisable by the abundant woolly
scales on petioles and rachis bracts and by the very large, brownish fruits. Its closest
affinity is clearly with L. mariae, there being in particular a striking similarity in
dimensions and other characteristics of the trunk, most strikingly at the base. Also, the
thickly waxy undersides of the leaves, their broad, thick-textured segments, and the
large white scales on the rachis bracts, together with overall size and architecture of
tire inflorescences, are all features shared by both species.

3. L. benthamii F.M. Bailey

Bailey (1902:1683); Beccari (1921:14,18; 1931: 84, t.6iv); Burret (1941: 325); Blake (1954:
127, t.6, f.3-4); Specht (1958: 208, t.2C-D); Covacevich & Covacevich (1978: 92);
Johnson (1979: 13); Tucker (1980:101); Jones (1984: 130).

Telopea 8(1): 1998

Type citation: 'Somerset, Cape York Peninsula, Frank L. Jardine' Dec 1897; holo BRI n.v.

L. humilis R. Br., var. [nov.] F.M. Bailey, Queensland Agric. J. 2: 130 (1898) [same
collection cited as for L. benthamii.]

L. holtzei Becc., Webbia 5(1): 18 (1921); Ann. Roy. Bot. Gard. Calcutta 13: 87 (1933).
Type citation: 'Australia boreale, presso Port Darwin (Holtze)'

Type: Northern Territory: Port Darwin, Holtze 1892?; FI, photos NSW, 4 sheets.

L. melanocarpa Burret, J. Arnold Arbor. 20:190 (1939).

Type: Papua: Daru Island, Western Division, L.J. Brass 6310, 9 Mar 1936; iso BRI.

Trunk solitary, straight, to 17 or possibly 20 m high, 12-14 cm diameter in upper part,
broadening gradually from about 5 m above ground in a long cone to 30 cm or more
diameter just above roots. Surface becoming fairly smooth with age, obscurely
patterned with sheath scars and petiole scars; vertical fissures numerous and regular
but shallow; basal 5 m or so usually bearing persistent, patent or slightly deflexed
petiole-stubs, these mostly 10-40 cm long.

Crown densely globose in adult state but greatly elongated in pre-flowering
(frequently quite tall) plants, consisting of 30-50 ascending to spreading leaves with
pendulous segment-lobes, the lower leaves slightly drooping. Ligules not seen but
remnants on petioles pale grey-brown on both sides, abaxial side with dense but
inconspicuous, closely appressed scales in deep pits.

Petiole (120-)160-180 (-210) cm long, 10-18 mm wide, roughly semi-circular in t.s.
with slightly flared margins and 2 concave grooves either side of a median ridge
above. Margins virtually unarmed on apical half except for few scattered blunt, very
small prickles and scattered, minute brown calli; toward base armed with ± closely
spaced pungent, narrowly triangular-conical, patent to retrorse shiny brownish-black
prickles to 6 mm long. Surfaces pale green, obscurely striate, clothed at first with
whitish scales toward apex, these densely crowded above, to 1.5 mm long, very
contorted and irregular, forming a deep spongy layer but soon marcescent, on lower
side more appressed, crustose, inconspicuous; basal-masses persisting as minute
conical red-brown pustules, hardly elongated. Hastula flat with upturned edges to
semicylindrical; base broadly 2-lobed with shallow median notch or 3-lobed with
lateral lobes wider than central; rim at 30-90° to costa, variable in width, its margin
early-necrotic and disintegrating.

Lamina moderately to strongly costapalmate with broad basal sinus, 70-115 cm long,
0.20-0.25 mm thick, chartaceous-coriaceous, moderately tough, moderately contorted
with 2 adaxial undulations either side of the slightly deflexed costa and lowermost
segments inflexed and erect, standing almost at right angles to petiole. Segments 29-39
either side of costa; largest segments 23-32 mm wide, broadening slightly toward
point of bifurcation, free for 60-75% of their length, bifurcated for 50-65% of free
length, the lobes slightly diverging, evenly tapering into soft, early-necrotic, acute but
never aristate or threadlike apices, lntersegmental appendages lacking, or present
only as very short remnants. Ribs: abaxial ribs in t.s. oblong with parallel faces, bluntly
squared to somewhat rounded, 0.5-0.8 mm thick, c. 1 mm deep; adaxial ribs slightly
smaller often tapering and rounded. Venation: major longitudinal veins 6-9 either side
of abaxial rib, prominent above and below; transverse veins evident both sides but
more prominent below, mostly at low angles to longitudinal veins, usually very
crooked and minutely sinuous, frequently continuous across 3—4 major longitudinal
veins. Surfaces olive-green, duller beneath, drying to pinkish-brown above contrasting
strongly with greenish-brown beneath, glabrous except for few white chaffy scales on
bases of ribs.

Rodd, Revision of Livistona (Arecaceae)

Inflorescences shorter than petioles. Partial inflorescences c. 9, subequal, short relative
to rachis; each branched to 2 further orders; rachillae (1—)5—12 cm long, 0.6-0.7 mm
thick, reddish-brown (fruiting stage), finely and closely but shallowly striate, minutely
but patchily strigose-pubescent with twisted, semi-appressed hairs, glabrescent
post-anthesis but remaining minutely scurfy-papillose; larger axes similar but not
pubescent. Rachis bracts flattened-cylindrical, the largest 20-22 mm diameter, smooth
and tightly sheathing, very stiffly coriaceous and rather hard, pinkish-brown ageing to
red-brown or greyish, finely but shallowly striate, patchily clothed with closely
appressed silvery scales, these wearing off after anthesis leaving minute pustules as on
petioles; bract apices triangular-ovate, smoothly acuminate. Flower-dusters 1-4 mm
apart, l-2(-3)-flowered; cluster axis 0.5-1.2 mm long, cylindrical or somewhat
bulging. Cluster-bract caducous, evident (on specimens available) only as a narrow
reflexed rim, if at all. Bracteoles not discernible.

Flowers (only seen post-anthesis, in early fruit stage) c. 1.5 mm long, possibly
cup-shaped. Anthopodium 0.3-0.5(-0.8)mm long, wider than long, recessed at base
and bluntly trigonous. Sepals very concave, membranous, thickening only close to
base, 0.8-2.0 mm long, basally connate for /-A their length; free portion triangular,
acute. Petals slightly concave, thick and leathery, c. 1.3 mm long, connate for %(?) their
length, broadly triangular-ovate, acute or subacute, slightly mucronate, auriculate at
base with relatively large, diverging auricles, inner faces with 2 deep narrow cavities
close and parallel to margins. Stamens c. %> as long as petals, connate for Z-'A their
length; filaments thick, broadly triangular-ovate, gently rounded either side toward
base, tapering smoothly to a slender apex; anthers (only 1 found) 0.33 mm long.
Carpels almost as long as stamens; ovaries cylindrical, abruptly contracted into short
slender styles.

Fruit obovoid-pyriform narrowing into prominent slender nipple at base, slightly
flattened ventrally, 9-13 mm long, 9—11 mm diameter; usually with floral remains
attached, the enlarged anthopodium and torus forming an extension c. 1 mm long.
Epicarp blackish when ripe with thin glaucous bloom, quite smooth, thin and weak.
Mesocarp c. 0.7 mm thick (dry), greenish and juicy when fresh, granular when dry.
Endocarp 0.2 mm thick, horny but rather brittle, smooth and dark red-brown on inner
face, outer surface with widely but irregularly spaced narrow longitudinal ridges. Seed
subspherical or slightly ovoid, flattened ventrally, 8-9 mm long, 7-8 mm diameter;
surface smooth and dark brown. Intrusion penetrating from lateral position for c. A.
seed diameter, in l.s. slightly lobed, occupying c. 'A seed length; in t.s. more regularly
2-lobed with somewhat diverging lobes, occupying slightly over 'A seed width.
Embryo lateral (about 3 o'clock relative to stalk), showing as prominent dimple on
seed surface. (Fig. la, 3e, 8a)

Distribution: Northern Territory, North Queensland, and southern Papua New
Guinea: in the Northern Territory restricted to lower reaches of rivers and creeks
draining north into the Arafura Sea and Van Diemen and Beagle Gulfs, also larger
islands off the north coast; in North Queensland to Cape York Peninsula north of Iron
Range, on lower reaches of coastal streams, mainly on east coast; in Papua New
Guinea (as far as is known) to coastal lowlands on west shore of Gulf of Papua.

Ecology: occurs mostly in gallery-rainforest along sluggishly-flowing streams usually
not far from tidal influence. In northwest Cape York Peninsula Tucker (1980) says it is
'Apparently confined to lagoon margins, moist gallery forests, and moister vine
scrubs. It colonises the mangrove-vine scrub ecotone with much success and attains
pure stand populations in several places. It extends inland where conditions permit.'
My own limited observations suggest that it occupies similar habitats in the Northern
Territory. However, on Elcho Island, NT, collectors' notes indicate that it grows in
swamp forests dominated by Melaleuca. Near Oenpelli it grows on river flats

Telopea 8(1): 1998

apparently among eucalypts. It appears mainly to be found on depositional sands,
silts and clays.

Conservation status: not considered at risk.

Specimens examined: Northern Territory: Darwin & Gulf: Howard River, c. 3 km N of Howard
Springs (23 km E of Darwin), Rodd 2904, 2905, 29 Oct 1974 (NSW, DNA, K); Howard River, Must
1150, 28 Aug 1973 (DNA, CANB); About S of Koolpinyah, Blake 16967, 10 Sep 1946 (BR1); Black
Jungle, Darwin, Trapnell 105, 13 July 1958 (BR1); Oenpelli, Specht 1187, 14 Oct 1948 (BR1, MEL, AD,
CANB); Jim Jim Ck, Jacobs 1790, 31 May 1974 (NSW); Obiri Rock, East Alligator River, Maloney 1978
(NSW 108859); Warangaiyu Lagoon, Elcho Is., Latz 6273, 19 July 1975 (DNA, CANB, NSW, BRI, K,
L); Elcho Island, Macoliochie 2119A&B, 2234, 5 July 1975 (DNA, NSW, CANB).

Queensland: Cook: Bamaga District, Cape York, 'Long Scrub’ 7 miles [11.3 km] NE of Bamaga on
Jacky Jacky Rd, Webb & Tracey 6122, 15 May 1962 (BRI); Jardine River, N bank, near crossing of
Peninsula Development Rd, Hind 377, Oct 1974 (NSW).

Papua New Guinea: Western Division: Daru Island, Brass 6406, Mar 1936 (BRI); Upper Wassi Kussa
River, Brass 8645, June 1936 (BRI). Central District: Brown River, alt. 100 ft, Millar NGF 38385,
20 Nov 1969 (BRI); 7 miles W of Kanosia Plantation, Darbyshire 685, 18 July 1962 (BRI); c. 6 miles
SW of Karema, Brown River, Schodde 2677, 30 July 1962 (BRI); Waigani Swamp, alt. 200 ft, Gillison
NGF 22018, 8 Dec 1964 (BRI).

Notes

1. This could be regarded as the least specialised Australian member of the genus,
though possibly the apparent lack of specialisation of some characters is due to
paedomorphy. Its leaves are thin-textured, quite non-glaucous, apparently
mesomorphic; its slender inflorescences are less richly branched than other species of
comparable stature, even down to the flower-clusters which are reduced to 1-2
flowers. Its preferred habitats are certainly more mesic than those of any other
Australian species except perhaps for L. drudci. It shares with the latter (and with the
sympatric L. muelleri) a fruit size at the bottom end of the range for Livistona, a feature
doubtless related to the preferences of the fructivorous birds in such habitats.

2. An interesting gross feature of this species is its propensity to retain long,
horizontally spreading petiole-stubs on the lower trunk. Although seemingly a
specific character, it is possible that this is developed only in closed-forest habitats
where there is protection from both fires and strong winds, allowing gradual decay of
the persistent leaves. There is evidence for this in a large plant at Obiri Rock
(photograph Maloney NSW 108859), where it is growing out in the open, probably in a
fire-prone habitat, and showing a smooth, bare trunk.

3. Other distinctive morphological features are: (1) pubescent rachillae (shared only by
L. humilis); (2) basal-masses of petiole, rib and lamina scales raised rather than sunken
as in most other species; (3) the noticeably discolorous surfaces of dried leaves which
become crinkled, and with a satiny sheen on both surfaces.

4. Beccari (1931) amplifies his 1921 protologue of L. holtzei: 'Discovered by Mr.
Nicholas Holtz, at Glencoe Station which is about 100 miles from Port Darwin'. This
was presumably based on his reading of a letter from Holtze to Mueller (28 November
1892), now held in FI, but a careful reading of the letter suggests that this was a
misinterpretation. The Livistona specimen appears to have been, in fact, collected in the
vicinity of Port Darwin. Blake's (1954) assignment of L. holtzei to the synonymy of
L. humilis was based on his observation that only the latter species occurred at Glencoe.

5. Beccari did not allow for the occurrence of L. benthamii in the Northern Territory,
though he believed his L. holtzei was closely related. The conspecificity of the Cape
York and Northern Territory palms is perhaps open to slight doubt, since the material
available from both areas is so fragmentary and largely juvenile and vegetative.

Rodd, Revision of Livistona (Arecaceae)

However, there is a striking correspondence between them in growth-habit, in
particular in the persistent petiole-stubs on the lower trunk, projecting horizontally.
I have not been able to locate the type material of L. benthamii from Somerset, but
Beccari's description of it corresponds well with my description above, except that the
fruits are described as being only 8-10 mm long and 6-7 mm 'through', compared with
11-13 x 9-11 as measured by me on the one fiuiting collection available (Rodd 2904
from near Darwin).

6. Burret (1941) appears to have thought his L. melanocarpa almost certainly conspecific
with L. benthamii, though regretting his lack of material for comparison. An isotype of
L. melanocarpa held in BRI is identical in nearly all characters with material of
L. benthamii from the Northern Territory. Hie only difference that might be regarded as
significant is in fruit size. Fruits in a package attached to the sheet are quite small
(about 7x6 mm) though apparently ripe. These dimensions are somewhat at odds
with Brass's field notes, in which the fruits are described as '± 1 cm long'. However, in
shape as well as mesocarp and embryo characters, the PNG and Australian fruits agree
well enough. Brass's notes on growth habit and habitat are also consistent with
L. benthamii in Australia, particularly his note 'lower half of trunk bristling with
persistent horizontal leaf-bases ± 20 cm in length'.

7. The New Guinea specimen, Millar NGF 38385, has the label note 'fruit red when
ripe'. However, the fruits on the specimen are small and very immature, being hollow
when broken. The leaf (in BRI) is also small and very immature, barely hardened, but
in characters of rachilla and floral remains, this material matches L. benthamii well.
Some New Guinea specimens have longer leaves than have as yet been encountered
in the relatively few adult collections from Australia, e.g. Brass 6310 has leaves 93 cm
long, widest segment 28 mm; Millar NGF 38385 has leaves 95 cm long, widest segment
27 mm. Schodde 2677 has leaves 114 cm long, widest segment 27 mm.

4. L. drudei F. Muell. ex Becc.

Beccari (1921: 15,19; 1931: 89-90, t.6vii); Johnson (1980:13); Jones (1984:132); L. drudei
F. Muell. nomen provisorium et subnudum Mueller (1878: 55, 147); Drude (1893: 55,147
other notes give 1893: 11,12).

Lectotype (here chosen): Queensland: Rockingham Bay, Dallaclnj (?), Sep 1865. (MEL)
[The lectotype is cited by Wendland & Drude (1875) under 'L. inermis', together with a
Thozet specimen from Rockhampton which is L. decipiens. Labelling of Dallachy's
collection in MEL is very confused, with a slip in Dallachy's (?) hand saying 'Valley of
the Mackay September 1865' plus field notes, whereas 'Rockingham's Bay' appears
only on the seed packet, also with 'sept. 1865'. The material is clearly L. drudei which
to the best of my knowledge does not occur anywhere near the Mackay River.]

[Livistona sp. 'Cooktown' (Jones 1984)]

Trunk solitary, to c. 20 m high, 15-20 cm diameter at 1 m above ground, broadening
from a point c. 1.5 m above ground into a flared cone to c. 30 cm diameter sitting on a
large root-mass up to 80 cm diameter and 40 cm high, reducing upward to a final
constant diameter of c. 17-23 cm. Surface of old trunk pale grey, very smooth,
waxy-whitish in patches, sheath scars barely discernible (but prominent on upper
trunk); vertical fissures close, shallow, fine, not very prominent; petiole-stubs absent.
Crown rather open, ± globose, consisting of 30-60 leaves with long, weak, arching
petioles, the lowest leaves radiating ± horizontally or moderately pendulous but with
petiole bases strongly ascending and imbricated, and pendulous segment-lobes.
Ligules prominent, pale straw-coloured ageing dirty white, persisting as papery plates
in upper half of crown, woolly-tomentose with narrow, deeply laciniate, contorted
whitish scales.

Telopea 8(1): 1998

Petiole 160-230 cm long, 15-25 mm wide, flattened-triangular in t.s. with very
rounded keel to unequally biconvex, with thin upturned edges. Margins virtually
unarmed on upper half except for closely but irregularly spaced minute dark brown
calli, grading randomly into pricklets; toward base armed with mostly retrorse,
narrow, slightly falcate or sigmoid, pungent, shiny dark brown prickles to c. 7 mm
long. Surfaces green, deep dull purplish toward base, ± smooth, hardly striate, the
upper with distinct median zone of fairly deep, narrow pits abruptly incised into
surface, often obliquely oriented, closely crowded and frequently confluent, mostly
empty of scales; the lower more closely and evenly pitted but pits shorter, broader,
frequently confluent; scales when present whitish, very irregular, appressed,
elongated, moderately thick, subentire, to c. 2 mm long, those on lower surface
shorter. Hastuln ± flat or shallowly channelled, somewhat warped; base V-shaped and
cuspidate to shallowly 3-lobed; rim at 25-35° to costa, to 40 mm wide, subacute, entire
except for (commonly) 1-2 deep clefts, up to % of width necrotic.

Lamina 100-140 cm long, c. 0.2 mm thick, strongly costapalmate, somewhat elongated
with the undissected portion rather long and narrow, the base ± truncate or with
broad sinus, fairly tough, not splitting readily; rather strongly contorted with 1 major
and 1 smaller adaxial undulation either side of the almost straight costa and lower
edges strongly involute, with lowermost few segments resupinate. Segments 33-35
either side of costa; largest segments 20-27 mm wide, free for 60-70% of their length,
bifurcated for c. 60% of free length, the lobes moderately diverging, evenly tapering
into finely attenuate but not threadlike, soft, flexuous, necrotic apices; in some
specimens secondarily bifurcated to a depth of 8-15 cm, the two parts of unequal
width. Intersegmental appendages 5-10 mm long, somewhat stiff and brittle, white.
Ribs: abaxial ribs in t.s. oblong or narrowing to lamina, bluntly square-edged to quite
rounded, to 0.9 mm thick, 1.5 mm deep; adaxial ribs semi-elliptic to triangular, to 0.8
mm thick, 1.0 mm deep. Venation: major longitudinal veins 8-9 either side of abaxial
rib, prominent above, less prominent below; transverse veins evident above,
moderately prominent below, orthogonal to quite steeply angled, frequently arcuate
or sinuous, mostly short, seldom crossing more than 1 longitudinal vein. Surfaces
deep green, upper glossy, lower rather dull, quite glabrous except for inconspicuous
pale brownish narrow scales on rib bases on underside.

Inflorescences slightly shorter than to longer than petioles, at most equalling petiole
plus half lamina. Partial inflorescences 7-8, short relative to rachis, each branched to
3 further orders; rachillae 0.5-8 cm long, 0.5-0.8 mm thick, minutely pubescent with
stiff antrorse hairs only c. 0.2 mm long, glabrescent post-anthesis, closely striate.
Rachis bracts flattened-cylindrical, to c. 2 cm diameter, smooth but rather loose,
coriaceous, rather tough, when dry pale yellowish-brown to darker reddish-brown,
closely striate, sparsely sprinkled with minute white appressed scales, glabrescent
post-anthesis; bract apices narrowly triangular, acute to slightly acuminate, variably
fringed with narrow white scales. Flower-clusters 0.5-4 mm apart, 2-5-flowered; cluster
axis 0.3-1.0 mm long, cylindrical to flattened, rather slender. Cluster-bract 0.5-1.0 mm
long, triangular-acuminate or sparsely fimbriate, very delicately membranous except
for fleshy base. Bracteoles approx. 1 per flower, similar to cluster-bract but shorter and
often rounded.

Flowers c. 2.2 mm long, funnel-shaped. Anthopodium 0.2-0.3 mm long, cylindrical,
rather slender, only slightly recessed at base. Sepals c. 1.5 mm long, connate for 'A their
length, appressed to petals, concave, membranous toward margins, free portion
narrowly triangular-ovate, subacute. Petals 1.7-2.0 mm long, connate for !4 their
length, slightly concave, fleshy-textured; broad-ovate, acute to obtusish with very
thickened apex, base with minute retrorse, rounded auricles; inner face with 2 deep,
narrow cavities close to margins and a less distinct median cavity. Stamens c. X as long
as petals, connate for almost 'A their length; filaments very fleshy, broadly ovate,
hardly shouldered, abruptly narrowed into a quite short, slender apex; anthers

Rodd, Revision of Livistona (Arecaceae)

0.35 mm long. Carpels c. % as long as stamens; ovaries obovoid-cylindrical, abaxially
gibbous apically, narrowing rather abruptly into styles only c. 0.2 mm long.

Fruit ± spherical or slightly pyriform, somewhat flattened ventrally, 10-12 mm long,
10-11 mm diameter (9-10 mm dorsiventrally). Epicarp dull purplish-black, finely
rugulose. Mesocarp c. 0.4 mm thick, granular, rather dry and oily when fresh.
Endocarp bony, rather brittle, straw-coloured, both surfaces smooth but dull.
Seed ellipsoid-globose, flattened ventrally, 8-9 mm long, 7-8 mm diameter, surface
dark brown. Intrusion penetrating from lateral position for slightly over A seed
diameter, in l.s. very slightly lobed but not apically dilated, occupying a little over 'A
seed length; in t.s. obscurely 2-lobed and slightly dilated, occupying c. / seed width.
Embryo lateral (c. 3 o'clock relative to stalk). (Fig. 3f, 8b)

Distribution: coastal northeast Queensland, scattered and discontinuous in coastal
lowlands from the Kennedy River north-west of Cooktown south to Rollingstone
Creek a short way northwest of Townsville, also on Hinchinbrook Island; additionally,
I have seen and photographed palms that appear to be of this species around the
estuary of the Proserpine River east-south-east of Proserpine, an occurrence separated
by VA degrees of latitude and 2 degrees of longitude from what is otherwise its
southern limit at Rollingstone Creek.

Ecology: chiefly a species of stream banks on flat coastal plains, mainly in bouldery
alluvium, apparently confined to streams with a continuous dry-season flow or at least
persisting as soaks. It occupies similar habitats above upper limit of mangroves beside
sheltered estuaries. It occurs less commonly on hill-slopes, in clay soils on shale or
siltstone. The vegetation types in which it occurs range from swamp-forest dominated
by Melaleuca leucadendra to fringes of gallery-rainforest or tropical rainforest bordering
on eucalypt forest dominated by species such as Eucalyptus tereticornis and Corymbia
tessellaris.

Conservation status: 3RC-. This coding is recommended in place of the code 3VC-
assigned by Briggs and Leigh (1996) who treated the Cooktown populations as a
separate undescribed species with a code 2R (Briggs & Leigh 1996).

Specimens examined: Queensland: Cook: Kennedy River 5.5 km N of New Laura Station, Irvine
2204, 26 Nov 1981 (QRS); Kennedy River, Irvine 2205, 26 Nov 1981 (QRS); 3.3 km off main road
along road at north end of Cooktown Airport, Hind 4594, Hill & Healy, 3 Aug 1986 (NSW). North
Kennedy: North side of Herbert River, at SW base of Mt Leach Range, 14.5 km NW of Ingham, Rodd
3135 & Jacobs, 17 May 1976 (NSW, BRI, K); at mouth of Herbert River, Webb & Tracey 11176, (no date)
(BRI); W.J. Huston's property 'Braemeadows' near Ingham, Thomas s.n., T7 Nov 1959 (BRI); Halifax,
approx. 2 km along road to Lucinda, Irvine 1832, 9 May 1977 (QRS, NSW); Hen Camp Ck, Kinduro,
near Bruce Highway, Irvine 1833,1834, 9 May 1977 (QRS, NSW).

Notes

1. This species was known only as a somewhat shadowy entity until Beccari (1931)
published an adequate description. In Wendland and Drude's (1875) account of
Australian palms a broad concept of L. inermis was adopted, and this taxon appears to
be the major element in their description. Mueller (1878) recognised this but chose to
follow Bentham (1878) who placed tropical Queensland plants of this taxon under
L. australis without comment. Bailey (1902) followed Bentham's treatment.

2. Mueller's epithet appeared only in the index of Fragmcnta vol. 11, being only
implied in the text and associated with L. australis, tire question of its specific status
being left open. Following upon some brief but arguably diagnostic descriptive and
distributional comments, Mueller concluded: 'Huic palmae nunc pro speciei vel
varietatis distinctione nomen Doctoris Oscaris Drude impono.' It is concluded that this
name cannot be regarded as validly published by Mueller.

Telopea 8(1): 1998

3. The first subsequent mention of L. drudei is in Drude's long but disorganised paper
on Australian Livistona, in which Mueller's assessment of its status is repeated
('Unterart oder Art?') but without any clear-cut pronouncement based on his own
opinion. In his indented key to species Drude relates L. drudei to L. australis.
His nesting it under L. australis without a number of its own suggests that it is given
some sort of inferior status; on the other hand non-overlapping distributions are
attributed to these taxa, implying separation. I conclude that Drude's (1893) treatment
of L. drudei is still as a nomen provisorium at species rank.

4. The next use of this name is by Beccari, in his synoptical paper of 1921. Here, though
the name is attributed to Mueller, it is clearly treated as a species, with an adequate
diagnosis in the key. No specimens are designated, but there is a precise reference to
Wendland and Drude's misapplied L. inermis ('non Rob. Br') as synonym, thus
indirectly citing specimens from which the lectotype is here chosen.

5. Jones (1984) draws attention to populations of Livistona in Cape York Peninsula, as
Livistona sp. 'Cooktown'; he cites the localities 'Flinders Island near Bathurst Bay,
Kennedy River near Lakefield, tributaries of the Endeavour River and south towards
Gap Creek near the Bloomfield River.' The only adequate collections I have seen from
any of these localities are Irvine 2204 and 2205, both from the Kennedy River. I have
examined these carefully and have concluded that these collections fall within the
range of variation of L. drudei. Hind 4594 from near Cooktown consists only of a fallen
dead leaf and old infructescence branch but the characters it exhibits all point to it
belonging in L. drudei.

5. L. australis (R. Br.) Mart.

Martius (1838: 241-2); Hooker (1877); Mueller (1878: 55); Bentham (1878: 146); Moore
& Betche (1893: 427); Bailey (1902: 1684); Domin (1915: 501); Maiden (1921: 353, t.256);
Beccari (1921:18; 1931: 79-80); Evans & Johnson (1962: 2); Moore (1963:149; 1965:148).

Corypha australis R. Br., Prodr.: 267 (1810).

Type: New South Wales: Port Jackson, Brown-, holo BM; iso FI, photo NSW.

Trunk solitary, to c. 25 m high, 25-35 (-40?) cm diameter at 1 m above ground,
broadening close to ground into a gently flared cone to c. 50 cm diameter, reducing
upward for 5-7 m to a final constant diameter of c. 20-25 cm. Surface rough but very
regularly patterned, prickly with protruding fibres, with strongly stepped sheath
scars; vertical fissures usually closely spaced, often brandling, deep and gaping, often
with criss-crossing fibres showing within; petiole-stubs absent from adult trunks.
Crown moderately compact to rather open, ± globose, consisting of 35-60 leaves with
straight to gently arching or moderately drooping petioles and pendulous
segment-lobes. Ligules moderately prominent, pale brown, patchily floccose with ±
appressed scales, margins irregularly fimbriate with fine, soft, straight, tapering scales
to 8-10 mm long.

Petiole 140-250 cm long, 10-20 mm wide; t.s. concave-convex to roughly semi-circular,
sometimes convex above and often with median ridge (sometimes 2 ridges). Margins
unarmed except at base or with a few small, pungent, hooked prickles near apex and
scattered minute, blunt teeth elsewhere; toward base armed with ± closely and
regularly spaced pungent, mostly retrorsely hooked blackish prickles to 2.5 mm long.
Surfaces green, purplish-brown toward base, very finely and shallowly
rugulose-striate; upper quite glabrous, sometimes irregularly pitted with narrow,
elongated pits, lower rather closely pitted with obscure, shallow, short and broad to
long and narrow pits, on young leaves bearing floccose scales to 3-4 mm long, usually
pale brown, divided to base into several straplike but usually tightly twisted,
branched laciniae; on older leaves usually only numerous minute, whitish to dark

Rodd, Revision of Livistona (Arecaceae)

brown pustular, basal-masses with small scale-remnants persisting. Hastula cylindrical
to flattened; base shortly but broadly 3-lobed, lateral lobes wider than central; rim at
10-90° to costa, 5-30 mm wide usually with blunt peak at centre, with very narrow to
broad necrotic margin.

Lamina 110-130 cm long, 0.15-0.25 (mostly 0.18-0.20) mm thick, moderately
costapalmate, usually with narrow basal sinus or even with lower edges overlapping,
chartaceous-coriaceous, moderately tough to rather brittle and readily split,
moderately to strongly contorted with mostly 2 adaxial undulations either side of the
moderately deflexed costa and lower edges sometimes involute with lowermost few
segments resupinate. Segments (35-) 40-46 either side of costa; largest segments 28-49 mm
wide, tapering slightly toward point of bifurcation, free for 54-62% of their length,
bifurcated for 50-63% of free length, the lobes parallel or slightly diverging, evenly
tapering into soft, acute but never aristate or threadlike apices, early-necrotic and
frequently broken or split. Intersegmental appendages lacking even on the youngest
expanded leaves, or rarely present as remnants to 3 mm long. Ribs: abaxial ribs in t.s.
elliptic to trapeziform (narrowing to lamina), quite rounded to sharp-edged on one
side only, rarely 2-angled though slightly rounded, 0.8-1.3 mm thick, 1.2-2.5 mm deep;
adaxial ribs mostly rounded or sharp-edged on one side only, 0.6-0.8 mm thick,
1.4—3.0 mm deep. Venation: major longitudinal veins 7-10 either side of abaxial rib,
moderately prominent above, slightly less prominent below; transverse veins evident
to moderately prominent above, prominent below, fairly crowded, rarely steeply
angled, finely sinuous and often also coarsely zigzagging or arcuate, frequently
branched, most continuous across of each half-segment. Surfaces green, glossy,
often with strong brownish tinge above, deep green beneath, closely striate between
veins, quite glabrous except for ribs minutely punctate with brownish, pustular
basal-masses, these stipitate on minute prickle-like stalks on adaxial ribs close to
hastula; on young leaves bearing dense, floccose, pale brownish scales toward bases
of ribs.

Inflorescences mostly shorter than petioles, occasionally equalling them. Partial
inflorescences 6-9, subequal, the largest slightly over half total inflorescence length but
with strongly recurving branches; each branched to 4 (-5) further orders; rachillae
5-25 cm long, 0.7-0.9 mm thick, white when fresh, glabrous, minutely warty,
wrinkled-striate when dry. Rachis bracts flattened-cylindrical, 4—5 cm diameter, often
rather loose, warped, often wrinkled, very tough, whitish to pale purplish-brown
when fresh, drying to deeper chestnut brown; ± striate, sparsely to densely
floccose-scaly, especially toward apex, scales usually pale brown, divided into several
twisted, branched laciniae to 7 mm long; bract apices triangular, acute or slightly
acuminate. Flower-clusters 0.5-3 mm apart, 1-4-flowered; cluster axis 0.5-2.0 (-3.0) mm
long, cylindrical, moderately slender; longer cluster axes often with lateral flowers
closer to base than apex. Cluster-bract 0.5-1.0 mm long, triangular-ovate, acute or
sometimes rather jagged, appressed, membranous except for fleshy base. Bracteoles 1
per flower or sometimes an extra one present on 1-flowered clusters, similar to
cluster-bract but shorter, mucronate.

Flowers 2.5-3.5 mm long, broadly funnel-shaped to rotate. Anthopodium 0.2-0.5 mm
long, conical to cylindrical sometimes with slight constriction below calyx, minutely
recessed at base with 3 obscure basal swellings. Sepals 1.2-2.1 mm long, connate for
'A-'A their length with very broad shallow sinuses, closely appressed to petals,
moderately concave, membranous apically, fairly tough, broadly to narrowly
triangular, subacute to acute. Petals 1.8-3.3 mm long, connate for 14-14 their length,
thick-textured, narrowly triangular to broadly oblong-ovate, bluntly acute to
mucronate or almost rounded with thickened, sometimes hooded apex; auriculate at
base with distinct retrorse, rounded to angular auricles; inner faces with 3 distinct,
broad but rather shallow cavities. Stamens %-A as long as petals, connate for /-/ their
length; filaments thick and fleshy, broadly to narrowly triangular-ovate, hardly

Telopea 8(1): 1998

shouldered, tapering smoothly to somewhat abruptly into a slender subulate apex;
anthers 0.5-0.7 mm long, the locules narrowly oblong (sterile?) to more broadly oval,
often slightly displaced. Carpels %- 3 / as long as stamens; ovaries cylindrical to ovoid
to obovoid and strongly gibbous apically, gradually to abruptly contracting into styles
0.2-0.4 mm long.

Fruit spherical, often slightly flattened ventrally, 13-22 mm diameter; suture often
faintly visible; base sometimes extended into a very shallow cone, on stalk to c. 3.4 mm
long consisting of thickened cluster axis topped by swollen anthopodium 0.9-1.5 mm
long and 1.4-1.8 mm diameter. Epicarp deep reddish-brown to black, or black with
brownish 'cheek', dull or with subdued satiny sheen, minutely verruculose and with
sparsely and irregularly scattered pale brown circular spots 0.2-0.8 mm diameter often
with central pore. Mesocarp 2-3 mm thick, olive green, firmly fleshy and hardly juicy
when fresh; when dry granular with reddish-brown tannin-bodies, sparse thin fibres
present, sometimes only adjacent to endocarp. Endocarp 0.2-0.3 mm thick,
straw-coloured to pale brown, flexible but rather weak, mostly adhering to mesocarp;
both inner and outer surfaces smooth but very minutely rough with loose spicular or
papillose surface cells. Seed 11-16 mm diameter; surface pale brown, minutely rough
and porous. Intrusion penetrating diagonally from sub-basal position for 'A-Z seed
diameter, in l.s. ± oblong with broadly flared base, and rounded to shallowly 2-lobed
summit, occupying 'A-A seed length; in t.s. similar but less flared at base, occupying
c. 2 A seed width. Embryo sub-basal, (4 to 5 o'clock relative to stalk). (Fig. 3g, 8c, 10)

Distribution: coastal eastern Australia south of 19° lat. It extends southward from
about the upper Noosa River (E of Gympie) in Queensland more or less continuously
through New South Wales to near Batemans Bay but thereafter shows increasingly
wide disjunctions until its last outpost in far eastern Victoria, close to the Snowy River
mouth (see Note 6 below). Furthest inland occurrences observed are about 100 km
from the sea in the upper Richmond and Clarence River basins, this distance
diminishing southward, and at altitudes not exceeding about 500 metres. North of the
Noosa River it is known from one small stand on Fraser Island (the common species
on Fraser Island is L. decipietts), and thereafter only as widely disjunct occurrences in
the Eungella-Crediton area near Mackay, and the southern Paluma
Range-Keelbottom Creek area northwest of Townsville. Stands of palms which may
well belong to this species have been sighted near the summit of Mt Elliot just south
of Townsville, on Mt Dryander north of Proserpine, and on Mt Aberdeen west of
Bowen. All these occurrences north of the Tropic are at altitudes between 500 and
1000 metres.

Ecology: occurs in a wide range of forest types, but most commonly in swamp-forest
associated with species such as Melaleuca quincjuenervia and Eucalyptus robusta ,
rainforest margins, or gallery-rainforest. In such wet situations it appears to be
indifferent to soil type. Also found frequently on exposed slopes above the sea,
principally on exposures of softer sediments, e.g. the Narrabeen Series close to Sydney,
sometimes locally dominant on such sites. In some near-coastal areas it is common on
valley-slopes in tall wet sclerophyll forest dominated by, e.g.. Eucalyptus saligna and
Syncarpia glomulifera, but is absent from such forests over wide regions. It is absent
from all the 'rain shadow' reaches of coastal valleys such as those of the blunter and
Hawkesbury Rivers, though this may be due to incidence of destructive floods rather
than insufficient moisture; isolated stands often occur in nearby tributary valleys in
the mountain escarpment.

Conservation status: not considered at risk. Eungella and Paluma populations were
listed as Livistona sp. 5 and sp. 6 respectively by Briggs and Leigh (1996) and given
conservation codes of 2RC- but I consider these to be part of this widespread and
abundant species (see note 4 below).

Rodd, Revision of Livistona (Arecaceae)

Specimens examined: Queensland: North Kennedy: Paluma Range, Dotswood Holdings, 22 km
west of Bluewater Creek on Forestry Road, Irvine 1928, 1929, 1930, 28 Jun 1979 (QRS, NSW);
Bluewater Creek Forest Road about 22 km SW of Bruce Highway, Rodd 3758 & Lavarack, 24 Nov
1981 (NSW). South Kennedy: c. 2 km SE of Eungella on Broken River Road, Rodd 3073 & Jacobs,
13 May 1976 (NSW). Wide Bay: Fraser Island, on margins of rainforest inland from Happy' Valley
on eastern side of island, Baxter 901, 4 May 1967 (BR1); 13 km E of Cooroy on Tewantin road, Rodd
3030 & Jacobs, 6 May 1976 (NSW, BRI); about 4.8 km N of Nambour, Moriarty 671, 672, Aug 1971
(BRt). Moreton: Mooloolaba, c. 70 miles [ 113 km] N of Brisbane, Moore 9229 & Everist, 9 Feb 1964
(BRI, BI I?); between Mt Glorious and Mt Nebo, Blake 21484, 15 Apr 1961 (BRI).

New South Wales: North Coast: Seal Rocks, 20 miles [32 km] E of Bulahdelah, Briggs s.n., 1 Aug
1964 (NSW); N of Ferny Creek, NW of 'Hut', W of Wallis Creek, Salasoo 3322, 6 Jan 1967 (NSW).
Central Coast: 'Warra' (Univ. of Sy'dney Biological Field Station), Pearl Beach, Rodd 2672, 17 Jul 1974
(NSW); Bayview, Cabbage Tree Road, just inside W edge of golf course, Rodd 3615, 12 Dec 1980
(NSW); McCarrs Creek, Kuring-Gai National Park, Rodd 2671, 17 Jun 1974 (NSW); Kumell, Botany
Bay', Boorman s.n., May 1906 (NSW); above Garie Beach, Royal National Park, Rodd s.n., 28 Aug 1975
(NSW, QRS); Upper Kangaroo Valley, Rodd 2398, 27 Aug 1973 (NSW); Mt Cambewarra NW of
Nowra, Constable s.n., 7 Dec 1950 (NSW). South Coast: 13 km from West Bodalla, Evans 96, 23 Jul
1976 (NSW); Tanja, on coast ENE of Bega, per Glcnfield Vet. Res. Station, s.n., Jul 1954 (NSW); New
Holland, Banks & Solander 1770 FI n.v. photo NSW.

Victoria: Cabbage Tree Creek, Webb CBG 010319, 9 Mar 1965 (NSW).

Notes

1. This is the only Livistona occurring naturally south of 26° latitude (except perhaps
for L. decipiens and L. nitida whose southern limits are both very close to this parallel).
Its southern limit at lat. 37° 45’ in eastern Victoria makes it the second most southerly
palm in the world after the New Zealand Rhopalostylis sapida.

2. A common species in the Sydney area, it is abundant on the south shore of Botany
Bay, and was collected by Banks and Solander when Cook's first voyage reached this,
its first Australian landfall, in 1770. No palm, though, is illustrated or even mentioned
in the 'Illustrations of the Botany of Cook's Voyage Round the World', which suggests
that even these intrepid botanists and their artist, Parkinson, gave little attention to
such massive plants. However, there are specimens in BM and FI and a manuscript of
Solander's in the British Museum (a copy of a page of which was kindly shown me by
Mr D. McGillivray), summarising the species to be included in his never-completed
manuscript flora. The names in it are mere provisional tags; four apparent palm names
are listed, one being Corypha nuda from Botany Bay. It is likewise strange that no
subsequent mention of this palm appeared in botanical or horticultural literature until
Brown's 'Prodromus' was published in 1810.

3. Another puzzle is Brown's attribution of this species to tire genus Corypha L., while
at the same time erecting the new genus Livistona for L. inermis and L. humilis. The
characters by which he differentiates these genera are very slight, amounting to little
more than a small shift in embryo position. Brown was surely aware of Linnaeus's
Corypha umbraculifera being typified (primarily) by Rheede's Codda-pana with its
striking terminal inflorescence, but he may have preferred to characterise it by its other
element, Rumphius's Saribus, which is the species now called Livistona rotundifolia.
Against the latter interpretation is the separation by Lamarck in 1786 of the latter
element as a distinct species, C. rotundifolia, in effect narrowing the application of
Linnaeus's name to 'Codda-pana'. Brown's separation of the two tropical Australian
species under a new genus ( Livistona) was probably influenced by their different
growth-habit and ellipsoid fruits, but these characters are not part of his generic
diagnosis.

4. Jones (1984), quoting the observations and collections of A.K. Irvine, distinguished
L. sp. 'Eungella Range' and L. sp. 'Paluma Range'. These were characterised as follows:

Telopea 8(1): 1998

'Eungella': 'The leaves of similar light green colouration on both surfaces, the broad
central area of fused segments in the lamina'; 'Paluma': 'The leaf has a circular profile
with a distinct sector at the base of the circle lacking segments; the lowest segments
near the petiole are compound; the flowers are distinct with blunt petals which are
held flat'. I have not as yet seen any convincing evidence that these populations differ
in any significant respect from L. australis, but better flowering and fruiting collections
may show characters by which they can be separated Tire foliage features noted by
Jones may be associated with highly mesic or shady environments.

5. Orscheg and Parsons (1996a & b), in an account of this species at its southern limit,
present one of tire most comprehensive ecological and phenological studies hitherto
published of an Australian palm species. They point out, inter alia, the large
disjimction of about 180 km between the only known Victorian stand (Cabbage Tree
Creek near Orbost) and the next most southern stand a little north of Bega, N.S.W. and
its very sparse occurrence from there north to Batemans Bay. A noteworthy
observation was that fruiting trees flower at intervals of at least 2 years, with fruit
ripening in the intervening year. The Victorian stand occurs in riparian forest
dominated by Eucalyptus botryoides, a vicariant of the E. saligna and E. grandis that often
accompany this palm further north.

6. Burret (1941) tentatively identified a palm collected by him in 1937 in the botanical
garden at Rio de Janeiro as a hybrid between L. australis and L. chinensis var. subglobosa,
but the evidence he presents for this conclusion is rather slender.

7. See '7. Evidence of dioecy (or androdioecy)' in the section 'Morphology and
Terminology' (p. 70), for observations of sexual behaviour in this species.

6. L. nitida AN. Rodd, sp. nov.

Palma magna L. australis affinis sed folia petiolis latioribus, partibus apicalibus
vaginarum longioribus truncatioribusque, hastulis planioribus ad laminam
appressioribusque; fructus magnitudines similes sed atro-nitidissimi, fibris mesocarpii
densioribus crassioribusque, pagina interiore endocarpii ordinatione reticulata.

Type: Queensland: Junction of Dawson River and Delusion Creek, NW of Cracow, 25°
ll'S, 150°11’E, A.N. Rodd 3055 & S. Jacobs, 9 May 1976; holo NSW; iso BRI, K, BH.

[Livistona sp. 'Carnarvon' (Jones 1984)]

Trunk solitary, to 25 or possibly 30 m tall, c. 30-35 cm diameter at 1.5 m above ground,
gradually narrowing upward to c. 20 cm near crown on old plants, broadening below
to a conical base c. 45-50 cm diameter. Surface ringed with sheath scars, obscure
toward base but prominent and stepped higher up; vertical fissures numerous,
regularly spaced, long and deep.

Crown globose, moderately dense, consisting of 35-50 strongly ascending to
horizontally spreading to deflexed leaves with segment-lobes arching smoothly to
abruptly with vertically pendulous tips. Ligules moderately prominent, pale
straw-coloured, their margins patchily fimbriate with closely crowded, soft, hairlike
brownish scales 10-15 mm long.

Petiole 170-190 cm long, 20-26 mm wide; flattened-triangular with smoothly rounded
keel to almost semicircular in t.s, slightly convex to slightly concave above, usually
with slight median keel. Margins virtually unarmed for most of their length, minutely
crenulate with closely spaced, slightly raised brown calli and at wider intervals
obscure blunt pricklets; at very base armed with ± conical dark brown pungent
prickles to 2 mm long, most antrorsely or retrorsely curved. Surfaces green, finely but
shallowly striate; upper virtually glabrous, obscurely and sparsely pitted, with few
appressed scales or scale remnants close to lamina; lower with scattered shallow,
somewhat elongated pits, deeper and more crowded close to margins, bearing

Rodd, Revision of Livistona (Arecaceae)

appressed, flat, ± oval white scales under 1 mm long, their margins irregular but not
fimbriate, soon deciduous leaving small but moderately conspicuous orange-brown
basal-masses flush with petiole surface. Hastula base ± truncate, shallowly 3-lobed, the
central lobe shorter and narrower, acute; rim flatfish, up to 6 cm long, ± appressed to
lamina, short- to long-triangular with broad, ± entire necrotic margin
Lamina 160-190 cm long, 0.20-0.30 (mostly 0.25-0.28) mm thick, moderately
costapalmate, truncate at base, coriaceous, moderately tough but fairly easily split,
slightly to moderately contorted with 1 adaxial undulation either side of the slightly
decurved costa, base on either side involute with lowermost few segments resupinate,
or involute and then shortly revolute (S-shaped in l.s., sometimes only on one side).
Segments 34-38 either side of costa; largest segments 30-42 mm wide, narrowing
slightly towards both point of bifurcation and basal sinuses, free for 63-70% of their
length, bifurcated for 60-73% of free length, the lobes slightly diverging, evenly
tapering, gradually attenuated into fine, soft but not threadlike, aristate apices,
early-necrotic for final 10-20 cm but mostly remaining entire. Intersegmental
appendages up to 60 cm long, ± persistent, pale brown, glabrous, threadlike. Ribs:
abaxial ribs in t.s. trapeziform (narrowing to lamina), sharply square-edged, 1.0-1.2 mm
thick, 2.0-3.0 mm deep; adaxial ribs obliquely trapeziform, with one sharp edge,
0.7-0.9 mm thick, 1.2-2.0 mm deep. Venation: major longitudinal veins 7-9 either side
of abaxial rib, moderately prominent above, slightly more prominent beneath;
transverse veins evident above, very prominent beneath, closely crowded, moderately
to steeply angled, most strongly arcuate and/or sinuous, mostly continuous across
whole half-segment and dendritically branched and/or anastomosing. Surfaces glossy
bright green, somewhat yellowish above, beneath with suggestion of glaucousness on
dried leaves (but waxiness not readily rubbed off), glabrous except for scattered
whitish scales on ribs.

Inflorescences equalling or slightly exceeding petioles, strongly arching. Partial
inflorescences 8-12, subequal, the longest slightly under half total inflorescence length,
each branched to 4 further orders; rachillae 5-20 cm long, c. 0.8 mm thick, glabrous but
whitish granular-papillose with age, obscurely wrinkled-striate; larger axes more
prominently striate. Rachis bracts flattened-cylindrical, to 5-6 cm in diameter, tightly
to rather loosely sheathing, frequently wrinkled, warped and tom, rich red-brown,
shallowly striate with raised veins at regular intervals, sparsely to densely
lanuginose-scaly with dirty-white tangled scales to 4 mm long, these stellately
branched to somewhat plumose, the branches hairlike, contorted and twisted, each
scale borne on a minute, forward-pointing, prickle-like stalk protruding from bract
surface; bract apices rather narrow-triangular, acute to moderately acuminate,
frequently split. Floiuer-clusters 0.8-3 mm apart, 2-4(-5)-flowered; cluster axis
0.5-1.5(-2.5) mm long, rather slender, longer ones with flowers scattered over their
apical two-thirds. Cluster-bract c. 0.7 mm long, appressed to cluster axis, concave,
oblong-ovate, acute, base persistent. Bracteoles 1 per flower with sometimes 1 empty
one in addition, broad-ovate, concave, ± fleshy, persistent at least as narrow rim.
Flowers (only 1 collection seen: Rodd 3055, with dead, semi-decayed flowers in
advanced bud) c. 2.2 mm long, shape at anthesis not known. Anthopodium c. 0.5 mm
long, 0.6-0.8 mm diameter, somewhat trigonous with 3 prominent downward bulges
around the recessed base, usually constricted at junction with sepals. Sepals loosely
appressed to petals, strongly concave, inflated, ± membranous except along median
line, c. 1.5 mm long, connate for 'A to almost 14 their length with very broad rounded
sinuses, triangular-ovate, subacute.. Petals thick-textured and leathery, strongly
concave, 2.0-2.2 mm long, connate for c. % their length, broadly obovate with bluntly
acute, slightly mucronate apex, auriculate at base with small(?), rounded(?) auricles;
inner faces with 3 large, distinct cavities. Stamens c. % as long as petals, connate for
c. %(?) their length; filaments rather thin(?) and flat, ovate, not shouldered, smoothly
tapering into a relatively long, slenderly aristate apex; anthers 0.42 mm long.

Telopea 8 ( 1 ): 1998

Carpels c. % as long as stamens; ovaries obovoid-cylindrical, strongly gibbous apically,
very abruptly constricted into rather thick, straight styles c. 0.25 mm long.

Fruit spherical, to c. 17 mm diameter, often with slight basal protrusion, on stalk to
5 mm long consisting of greatly thickened cluster axis topped by swollen
anthopodium to 2 mm long and almost 2.5 mm diameter. Epicarp jet-black, very
smooth and glossy when fresh, drying to irregularly wrinkled with bluish waxy
bloom. Mesocarp c. 2.5 mm thick fresh, greenish, fleshy, drying to c. 1 mm thick,
containing numerous fairly crowded, moderately thick longitudinal fibres,
concentrated mainly toward endocarp. Endocarp 0.20-0.25 mm thick, straw-coloured,
rather hard and brittle, adhering closely to mesocarp in dried ripe fruit, its inner
surface with a distinct regular pattern appearing reticulate to the naked eye, when
magnified seen as small circles of radiating fine striations or minute, loosely attached
fibres. Seed 10-12 mm in diameter, not flattened ventrally, pale brown finely speckled
whitish except for deeper brown hilum. Intrusion penetrating ± diagonally from well
below median position for c. % seed diameter, in Ls. half-dumbbell shaped, occupying
less than 'A seed length, in t.s. very similar. Embryo at or slightly above median
position (2 to 3 o'clock relative to stalk). (Fig. 2a-b, 3h, 8d, 9b-c)

Distribution: subcoastal southeast Queensland, confined to the upper Dawson River
basin and the uppermost Comet River basin, within an area bounded approximately
by Moura and Rolleston to the north, Wandoan and Injune to the south, the Great
Dividing Range to the west, and Auburn Range to the east. This region is
approximately 350 km WNW of Brisbane and covers approximately 2 degrees of
latitude and 2 of longitude; the downstream limit on the Dawson is not known to me
with certainty. The species is abundant at Theodore but is not evident where the
Dawson Highway crosses the river west of Moura. I am informed by Peter Hind that
there is a population of Livistom in Cania Gorge, about 30 km NW of Monto, which
may be this species and which would extend its range into the Burnett River catchment.

Ecology: confined to banks and channels of larger, semi-permanent streams, in sandy,
silty or gravelly alluvium; also at cliff-bases and on cliff-ledges in sheltered sandstone
gorges, most notably in Carnarvon Gorge. In the eastern part of its range it is mostly
found associated with Eucalyptus tereticomis and E. microtheca s. lat., but to the west
(Carnarvon Range, Great Dividing Range) it is frequently associated with
E. tereticomis, E. melanophloia, Corymbia maculata, C. citriodora, Angophorafloribunda and
Macrozamia moorei.

Conservation status: 2RC- (Briggs & Leigh 1996, as Livistona sp. 3: Carnarvon & Taroom)

Specimens examined: Queensland: Leichhardt: Cracow-Theodore road on Delusion Creek about
12 miles [19 km] N of Cracow, Johnson 847, 28 June 1959 (BRI); Junction of Dawson River and
Delusion Creek, NW of Cracow, Rodd 3051, 3052, 3054, 3055 & Jacobs, 9 May 1976 (NSW, BRI, K,
BH); Palm Tree Creek 19 km N of Taroom, Telford 5648, 7 June 1977 (CBG, NSW); Palm Tree Creek
c. 12 km from Taroom on Taroom-Theodore road, Moriarly 1280, 15 May 1973 (BRI); Robinson Creek
and Palm Tree Creek, Taroom, Stiller s.n., 16 Sep 1960 (BRI); Robinson Creek 10 miles [16 km] NW
of Taroom, Leichhardt District, Speck 1986, 3 May 1964 (CANB, BRI); Carnarvon Creek, near
entrance to Carnarvon Gorge National Park, Rodd 1112, 12 Aug 1970 (NSW).

Notes

1. This species is distinguished from L. australis on rather slight characters, but these
appear to be fairly constant and reliable over the whole area of its occurrence.

2. The specific epithet, meaning 'shiny', refers to its most distinctive feature, viz. the
highly glossy surface of the ripe fruits, in contrast to the dull, minutely rugulose fruits
of L. australis. Internal features of the fruit also serve to distinguish it: the mesocarp has
more numerous, thicker fibres, and the endocarp is thicker, its inner face displaying an
unusual reticulate pattern. In vegetative characters the only striking differences from

Rodd, Revision of Livistona (Arecaceae)

L. australis are the shape and length of the ligules; good specimens of these organs,
however, are represented in only one collection each of these two species, both
obtained by felling an adult tree and dissecting the crown. Other foliar differences are
more subtle, the most apparent being the slightly wider petiole and flatter, more
appressed hastula in L. nitida ; the undulation of the lamina is possibly also less
pronounced than in L. australis.

3. Growth-habit is strikingly similar to that of L. australis, also trunk dimensions and
surface characters. It is possible, though, that L. nitida attains a greater height than any
other Australian species: the magnificent stand at the junction of Delusion Creek and
the Dawson River includes individuals which, by a visual estimate, possibly exceed
30 metres.

4. For approximately the last 30-40 years this has been recognised as a distinct but
unnamed species in the Queensland Herbarium; the late Dr S.T. Blake was the first to
treat it as such. Previously its populations were doubtless treated as L. australis (just as
were those of the far more distinctive L. decipiens), though 1 have not come across any
record even of the existence of palms in the Dawson River basin from earlier than
the 1960s.

7. L. decipiens Becc.

Beccari (1910: 301, fig. 1, t. II; 1921:18; 1931: 81, t.6II); Moore (1963:150); Johnson (1980:
13); Blombery & Rodd (1982: 123); Jones (1984: 131).

Lectotype (chosen here): [cultivated] A. Robertson-Proschoivsky s.n., May 1908 (sheet
with older inflorescence) (FI, photo NSW). Syntypes: cultivated Nice, France,
A. Robertson-Proschoivsky s.n., Sep 1908 (leaf) (FI, photo NSW); A. Robertson -
Proschowsky s.n., Sep 1907 (fruits) (FI, photos NSW).

L. decipiens var. polyantha Becc.

Beccari (1921: 15,18; 1931: 82).

Type: cultivated Bot. Garden Buitenzorg (now Bogor), Java; holo FI (two sheets), photo
NSW; ?iso BO.

Trunk to c. 10 m tall, c. 25 cm diameter at 1 m above ground, reducing to c. 20 cm
diameter toward top, widening below (from c. 0.5 m above ground) into a gently flared
cone up to c. 35 cm diameter at base. Surface moderately smooth but with prominent
annular sheath scars; vertical fissures numerous, fine, scattered; petiole-stubs
frequently present on lower 1-2 m of trunk, closely appressed, to c. 10 cm long.
Crown globose or frequently broader than deep, moderately dense, consisting of
30-60 leaves with fairly straight, rigid petioles, the leaf-blades spreading rigidly to the
point of bifurcation of their segments but with lobes vertically pendulous producing a
curtain-like effect. Ligules not very prominent, pale reddish-brown aging to dirty
white; shredding into fibres at early stage leaving only remnants of unshredded apex,
its margin sparsely fimbriate with long slender white scales, its outer surface with
close network of strongly raised fibre-bundles and scattered, inconspicuous,
contorted, appressed translucent scales.

Petiole 160-280 cm long, 18-24 mm wide; flattened-triangular in t.s. with bluntly
angular to rounded keel, slightly convex to slightly concave above sometimes with
slight median ridge. Margins virtually unarmed over most of petiole length or with
scattered, small, mostly antrorsely curved, pungent prickles; toward base with closely
and rather evenly spaced, slender, shiny dark brown pungent prickles c. 4 mm long,
retrorse or patent or mixed antrorse-retrorse. Surfaces finely rugulose-striate, upper
with sparse, shallow pits, in 2 rows either side of centre, containing inconspicuous,
early-deciduous scales; lower with more numerous, poorly defined, often confluent

100

Telopea 8(1): 1998

pits, densely but not completely covered in crustose, dirty white scales, these roughly
circular, c. 1 mm long, quite thick but with flat, shaggy surface, dotted with minute
reddish tannin-bodies, margins shortly fimbriate; persistent basal-masses very
minute, slightly elongated, flush with surface or very slightly raised. Hastula base
broadly 3-lobed, the centre lobe longest, acute; rim almost parallel to costa, very
ragged on fully expanded leaf, 3-lobed, the centre lobe 8-15 cm long, lateral lobes
much shorter, their outer edges adnate to lowermost segments of lamina; necrotic
margin conspicuous, brittle, fragmenting.

Lamina 13CM80 cm long, 0.20-0.25 mm thick, strongly costapalmate, elongated,
dissected virtually to base into slender radiating segments, (palman very elongated
and extremely narrow in upper part, forming a mere narrow wing on either side of
costa), coriaceous, moderately tough, splitting readily, the whole lamina very strongly
contorted with 2 adaxial undulations either side of the slightly recurved costa and
lower edges inflexed with lowermost 8-11 segments resupinate and again reflexed to
give an S-shaped section. Segments 35-42 either side of costa; largest segments 26-34 mm
wide just below point of bifurcation, narrowing toward bases to 18-25 mm, free for
82-88% of their length, bifurcated for 44-54% of free length, the lobes slightly
diverging and drooping from base to a vertically pendulous attitude, tapering evenly
to soft, shredded, early-necrotic apices. Intersegmental appendages up to 8 cm long,
fine, somewhat brittle, threadlike, dull purplish at tips, few persisting on older leaves.
Ribs: abaxial ribs in t.s. oblong-elliptic to trapeziform-obovate (narrowing to lamina),
bluntly square-edged to rounded, 0.8-1.5 mm thick, 1.5-3.0 mm deep; adaxial ribs
trapeziform, broadest at lamina, bluntly to somewhat sharply squared at summit,
c. 1.3 mm thick, 1.3 mm deep. Venation: major longitudinal veins 7-8 either side of
abaxial rib, prominent above, less prominent beneath; transverse veins evident but not
prominent above, moderately prominent beneath, rather crowded, nearly all angled
but most not steeply, most arcuate or sinuous, most continuous across at least
1 longitudinal vein, many across 2-5. Surfaces green, upper glossy and slightly
yellowish, quite glabrous, lower duller green with very thin, virtually invisible waxy
bloom, glabrous except for scattered whitish scales on ribs.

Inflorescences from shorter than petioles to exceeding petioles by half lamina length,
erect and arching or arching to below horizontal even in flower; rachis very stout.
Partial inflorescences 8-12, markedly distichous, all ± equal in size, the larger axes
sinuous in a horizontal plane; branched to 4 further orders; rachillae 5-20 cm long,
0.5-0.7 mm thick, wrinkled-striate, glabrous but minutely granular-papillose, pale to
dark red-brown by fruiting stage; larger axes similar, sometimes with bands of woolly,
slender scales on inner faces of lst-order branches, otherwise glabrous. Rachis bracts
flattened-cylindrical to obscurely trigonous, broadening toward their apices, up to
4 cm diameter, somewhat loose, transversely wrinkled and warped, larger ones often
stretched and rent into fibres near apex, very stiff and hard-textured, finely
rugulose-striate toward apex, glabrous except for dusting of fine whitish granular
papillae toward apex, greenish right up to fruiting; bract apices ovate, shortly
acuminate, necrotic, greyish to red-brown. Flower-clusters 0.7-5 mm apart, racemosely
(2-)3-6-flowered; cluster axis 0.3-5(-6) mm long, c. 0.25 mm thick at flowering,
bearing flowers most of its length. Cluster-bract c. 0.5 mm long, narrow, acuminate or
fimbriate with very delicate, early-deciduous apical portion, thickened across base.
Bracteoles 1 per flower, short, broad, appearing as narrow rim even at early
flowering stage.

Flowers c. 2.0 mm long, probably funnel-shaped (only seen at very early anthesis).
Anthopodium slender, c. 0.5 mm long with base oblique, junction with cluster axis
slightly recessed, with narrow sharp rim. Sepals forming a loose, somewhat inflated
cup around petals, membranous, thickening gradually to base, 1.3 mm long, connate
for at least % their length, ovate-triangular, obtuse. Petals thin-textured but tough,
slightly concave, 1.8 mm long, connate for c. K their length, broadly ovate, broadly

Rodd, Revision of Livistona (Arecaceae)

101

acute, slightly thickened and indexed at apex, auriculate at base with rounded,
broadly overlapping auricles; inner faces with two large, shallow cavities adjacent to
margins and a smaller cavity beneath thickened apex. Stamens c. % as long as petals,
connate for c. 'A their length; filaments thin and flat, strongly shouldered, tapering
smoothly but at wide angle into very short subulate apex; anthers 0.35 mm long.
Carpels almost as long as stamens; ovaries obovoid, strongly gibbous abaxially,
narrowing fairly abruptly into styles c. 0.35 mm long.

Fruit spherical, 13-15 mm diameter, perfectly rounded at base and apex, on stalk up
to 9 mm long consisting of crooked, enlarged cluster axis of variable length, plus
enlarged conical anthopodium to c. 2 mm long, plus greatly swollen floral torus
(between sepals and petals) c. 1 mm long and 2 mm diameter. Epicarp black, smooth
and shiny when fresh, closely wrinkled when dry. Mesocarp 1-1.5 mm thick, soft and
fleshy but toward endocarp containing a close reticulum of flattened fibres, very
conspicuous in old fallen fruits. Endocarp c. 0.15 mm thick, closely adhering to seed
where it persists after fruit decays, outer surface smooth, pale brown. Seed spherical,
c. 10 mm diameter, shiny dark brown. Intrusion penetrating from lateral position for
% width of seed, in l.s. shallowly 2-lobed, slightly dilated apically, occupying 'A seed
length, in t.s. similar but occupying less than 'A seed width. Embryo sub-lateral
(c. 4 o'clock relative to stalk). (Fig. 2d, 3i, 9d, 11)

Distribution: east-coastal Queensland from latitude 19°30’ (approx.) to 26°, i.e. from
just south of Townsville to the northern end of Cooloola National Park, south-east of
Maryborough. Distribution is semi-continuous over much of this range, with only
minor gaps; it is apparently restricted to a narrow lowland zone, extending inland no
further than about 50 km and onto bases of hills up to no more than 100-200 m altitude.

Ecology: most commonly found along banks of rivers and smaller streams, on alluvial
deposits; in some areas found also on gentle to steep hillsides, on clay soils derived
from shales or skeletal soils over igneous rock; also near seashore in sheltered
hind-dune hollows and swampy depressions. Mostly associated with margins of
gallery-rainforest or dry rainforest or extending a little into open-forest dominated by
Corymbia tessellaris or Eucalyptus tereticornis, sometimes in swamp-forest with
Melaleuca leucadendra or M. quinquenervia and Eucalyptus alba.

Conservation status: not considered at risk.

Selected specimens examined: Queensland: North Kennedy: West Baratta Creek, Bruce Highway,
between Ayr and Giru, Rodd 3117, 14 May 1976 (NSW); 51 miles [82 km] S of Townsville, Trapnell
305, 20 July 1960 (BRI); Bruce Highway, a few km N of St Margaret Creek between Ayr and
Townsville, Moriarty 767, 21 May 1971 (CANB, BRI); Haughton River near Bruce Highway, Irvine
1818, 21.1.1977 (QRS). South Kennedy: Palm Creek, Bruce Highway, North Kennedy District, Briggs
2071, 8 Aug 1968 (NSW); 10 km W of Mackay on bypass road, Rodd 3075, 13 May 1976 (NSW);
E of Dingo Beach road on 'Earlando' Beach turn-off, c. 35 km NNE of Proserpine, Rodd 3077 &
Jacobs, 13 May 1976 (NSW, BRI). Port Curtis: between Byfield and Mary vale in forest. White 8197,
25 Sep 1937 (BRI); Yeppoon-Emu Park road, c. 11 km S of Yeppoon, Rodd 3069 & Jacobs, 12 May 1976
(NSW, BRI, K, BH); Joskeleigh, about 15 miles [24 km] E of Rockhampton in sandy, swampy soil
c. 'A mile [0.8 km] from beach, Everist 7634, 17 Mar 1964 (BRI, K); Moore's Creek Range near
Rockhampton, Thozct s.n„ (before 1875) (MEL); near Bustard Bay (S.F. 86 Eurimbula), Cooper s.n.,
July 1966 (BRI); between Agnes Water and Round Hill Head, S of Bustard Bay on coastal dunes,
Everist s.u., 28 Mar 1970 (BRI); Agnes Water, 40 km E of Miriam Vale, McDonald 1224X & Batianoff,
4 Apr 1975 (BRI); Palm Creek, 8 km S of Miriam Vale, Bruce Highway, Hind 257, Aug 1973 (NSW);
Reedy Creek, 8 km SE of Lowmead on Bundaberg road, Rodd 3045 & Jacobs, 8 May 1976 (NSW, BRI).
Wide Bay: Cooloola State Forest ENE of Gympie, Moriarty 668, 668A, 669, Apr 1971 (BRI).

Cultivated: Bogor, Java, C.X. Furtado, Singapore Field No. 30948, Gardens No. XII.E.91, iv-v.1936 (BRI).

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Notes

1. In his protologue paper (in Italian) Beccari recounts how this palm was known in a
number of gardens in the Riviera, under the name 'Copernicia cerifera' and that some
palm collectors realised that this name could not be correct. He quotes at length from
a letter written to him by B. Chabaud di Tolone, who reported that this identification
had originated from (the late) Nabonnand, 'rosieriste au Golfe Juan', who had two
healthy specimens 2.5 m high in his garden in 1883. Chabaud, however, had the same
palm growing, but his originated from seed sent to him by von Mueller under the
name Livistona humilis. He mentions another grower, the Comte d'Empremesnil, who
likewise had it as L. humilis.

2. From the above, it seems likely that all these cultivated plants originated from the
one source. One might plausibly surmise that this source was Thozet, who sent to von
Mueller a specimen of this species he had collected at Moores Creek Range,
Rockhampton, before 1875. According to Hall (1978), Thozet, who lived in the
Rockhampton area from 1858 to his death in 1878, sent most of his specimens to von
Mueller in Melbourne, 'but was also active in making Australian plants known in
France'. It is possible, even likely, that Thozet made repeated collections of this species,
which is reasonably frequent in the Rockhampton area, for distribution overseas.

3. It is puzzling that this species should have become relatively common in the Riviera
by the late 19th century, yet unknown in cultivation in Australia right up to the 1950s.
Beccari's correspondent Robertson-Proschowsky noted that it throve in his garden at
Nice, growing faster than L. australis and very much faster than L. chinensis, though he
considered it less ornamental than either on account of its 'leaf divisions hanging like
narrow sh ips around the end of the petiole giving it the aspect somewhat of a mass of
rags' (my translation).

4. Beccari (l.c.) believed that his L. decipiens originated from temperate eastern
Australia, basing this notion largely on a specimen (consisting only of a leaf) sent to
him by Ewart, collected by E. Reader from Mt Dromedary, New South Wales. This,
Beccari considered, could not be L. australis by reason of its segments
'profondissimamente divisi in lunghe lacinie pendenti' which feature he regarded as
characterising L. decipiens. I have not seen this specimen, but present knowledge gives
no reason to doubt that L. australis is the only species present in New South Wales.

5. Var. polyantha Becc. was distinguished from typical L. decipiens by its smaller flowers
in clusters of 3-5 at apices of 'pedicels', and smaller fruits. In his 1921 protologue
Beccari gives simply 'Buitenzorg' (cultivated in botanic garden) as its origin, but in his
monographic 1931 treatment he is more specific, nominating its source as 'XII E. 91
position in the Garden; No. 408 of the Herbarium'. In BRI there is a good specimen
collected in Bogor by Furtado in 1936 from a plant 50-70 feet [c. 15-20 m] tall, 'Gardens
No. XII.E.91'; there seems little doubt that this is the type plant of var. polyantha, and
it falls comfortably within the range of variation of the wild species. In FI there are two
sheets (one with leaves and the other with an inflorescence) labelled 'No. 908 Herb.
Hort. Bot. Bog.' and 'XII E 91' plus 'Livistona decipiens Becc.' — the herbarium
number is hand-written and looks more like '908' than '408'.

6. L. decipiens is a species readily recognisable in the field by its very large but very
deeply dissected leaves; dissection is mostly to within 8 cm of the hastula and within
3 cm of the costa in its upper half. Another striking feature is best exhibited by fruiting
specimens, viz. longer cluster axes than any other Australian species, up to 9 mm in
fruit; in flower the cluster axes are considerably shorter but very slender, with flowers
racemosely arranged. Fruits are distinctive by their well-developed, crowded inner
mesocarp fibres.

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103

7. The only species it is likely to be confused with is L. drudei, which comes close to it
geographically, but this has less deeply dissected, smaller leaves, much shorter cluster
axes, and slightly pyriform, smaller fruits, apart from differences of indumentum and
growth-habit.

8. L. fulva Rodd, sp. nov.

Palma solitaria magnitudine mediocris, trunco ad 13 m alto, c. 25 cm diametro. Coma
globosa, sparsa foliis rigide radiantibus petiolis ultra 2 m longis, c. 14 mm latis
proxime laminam; laminae c. 1 m longae, dissectae ad c. 'A totarum longitudinum,
segmentis grandioribus c. 40 mm latis brevissime bifurcatis. Inflorescentiae quam
petiolis breviores, 6-9 ramis lateralibus, illis denuo 4-plo ramosis, rhachillis 5-16 cm
longis, c. 0.7 mm diametris. Flores 1-3 fasciculis sessilibus. Fructus globosus, niger,
pruinosus, c. 15 mm diametro. Semen ad 12 mm diametro, intrusioni testae laterali,
embryone laterali.

Type: Queensland: Blackdown Tableland, just below escarpment at NE end (road
pass), 23°43'S, 149°07'E, alt. 4-500 m, A.N. Rodd 3062 & S.W.l. Jacobs, 11 May 1976; holo
NSW; iso BRI, K, BH.

[Livistona sp. 'Blackdown' (Jones 1984)]

Trunk solitary, to 13 m high, 20-25 cm diameter at 1 m above ground, below usually
broadening gently at base to perhaps 30 cm diameter at ground. Surface closely ringed
with stepped sheath scars, rough-textured with prickly fibres; vertical fissures numerous,
regularly spaced, deep and gaping, some 10 cm or more long; petiole stubs absent.
Crown rather open, hemispherical to somewhat more globose, consisting of c. 35
mostly spreading, flattish leaves with long, arching petioles, the lowest drooping to
not much below horizontal or less commonly to a steeper angle. Ligules moderately
large, pale straw-brown, their margins with finely laciniate soft teeth c. 10 mm long,
outer surfaces appressed-tomentose with twisted, hair-like whitish scales.

Petiole 200-250 cm long, c. 14 mm wide; roughly semi-circular in t.s., the upper side
with a concave groove either side of a pronounced median ridge. Margins unarmed on
apical half at least, with only occasional small, dark-coloured bumps and more
frequent minute calli, these with a slightly sunken, elongated dark brown centre; neai
base armed with closely-spaced retrorse, slightly sigmoid, slender, very pungent
blackish prickles up to 10 mm long. Surfaces both yellowish-green, closely striate
especially in furrows on upper side, thinly clothed in appressed, translucent, hair-like
scales with rows of red-brown tannin-bodies giving them an orange cast, soon
glabrescent leaving persistent, densely crowded but very minute dark brown
basal-masses, sparser but slightly larger and elongated, slightly raised, on underside.
Haslula base ± truncate; rim at approx. 90° to costa, to 4 cm or more wide, flattish in
middle part but lateral parts inflexed abruptly at an acute angle, early-necrotic and
disintegrating into irregular narrow ribbons. Rudimentary abaxial hastula present, as
cuspidate to long-acuminate, narrow dark-coloured rim projecting from costa as a
slight step.

Lamina 90-100 cm long, 0.2 mm (or slightly more) thick, weakly costapalmate,
± circular in outline, chartaceous-coriaceous, moderately tough and flexible, virtually
flat to slightly contorted with 1 broad, gentle adaxial undulation either side of the
slightly deflexed costa and lowermost segments slightly involute. Segments 30-33
either side of costa; some lower segments partially fused into 2—3-ribbed compound
segments with primary sinuses displaced to near their apices; largest segments 35-40
mm wide, widest at base, free for 50-55% of their length, bifurcated for at most a few
cm, the lobes of the apical few segments usually rounded, diverging, often slightly
hooked, 0.5-1 cm long, of other segments ± parallel, acute to obtuse, 1-5 cm long.
Intersegmental appendages present only at early stages, very fine, weak, threadlike, to

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Telopea 8(1): 1998

c. 6 cm long, breaking off to leave an occasional small remnant on older leaves. Ribs:
abaxial ribs in t.s. ± oblong, bluntly square-edged to quite rounded, c. 2 mm deep, 0.9 mm
wide; adaxial ribs oblong to elliptic, mostly rounded, c. 0.7 mm wide, 1.5 mm deep.
Venation: major longitudinal veins 7-10 either side of abaxial rib, prominent on upper
surface, somewhat obscure beneath; transverse veins evident above, more prominent
beneath, most near-orthogonal with respect to longitudinal veins, most arcuate,
sigmoid or sinuate and continuous across 2-4 longitudinal veins. Surfaces: upper
glabrous and slightly greyish green with slight satiny gloss; lower paler bluish-green,
on newly expanded leaves partially masked by conspicuous orange-brown
indumentum, this finely and closely floccose-tomentose consisting of thin translucent
scales with densely tangled fimbriate margins and flecked profusely with
orange-brown tannin-bodies, their basal-masses raised, slightly elongated, also
orange-brown and persisting after scales fallen; both surfaces finely and quite
deeply striate.

Inflorescences a little shorter than petioles. Partial inflorescences 7-9, closely spaced,
subequal, the largest approximately half as long as total inflorescence, moderately
deflexed with pendulous rachillae, each branched to 4 further orders; rachillae 5-16 cm
long, c. 0.7 mm thick, cream at flowering darkening to red-brown, closely and deeply
striate, densely granular-papillose; larger axes similar, with deep grooves in addition
to fine striations. Rachis bracts cylindric-conic broadening toward apex, to c. 4 cm
diameter, loose and warped, thick-textured and stiffly coriaceous, rich chestnut-brown
but rapidly greying on exposure, finely but rather obscurely striate; patchily clothed
with floccose-tomentose orange-brown scales as on leaf undersides, these dense and
continuous near apices; bract apices apparently initially ovate-cuspidate but breaking
down at an early stage of development into a number of irregular, fibre-fringed teeth,
resulting in a ± truncate rather than 2-lobed bract. Flower-clusters 1-4 mm apart,
1-3-flowered; cluster axis 0.5-1.5 mm long, irregularly cylindrical, moderately slender.
Cluster-bract ovate-triangular, c. 0.5 mm long, membranous, whitish, laciniate at apex
but laciniae rarely surviving beyond early flowering stage. Bracteoles 1—3(—5?), similar
to cluster-bract but smaller.

Flowers c. 1.8 mm long, broadly funnel-shaped. Anthopodium c. 0.5 mm long,
moderately slender, obscurely trigonous and recessed at base. Sepals membranous,
thickening toward median line and base, concave in lower half, 1.3 mm long, connate
for M their length, narrowly triangular, acute to slightly cuspidate. Petals moderately
concave, thick and leathery, 2.0 mm long, broadly triangular-ovate, acute and strongly
hooded at apex; auriculate at base with rather broadly rounded auricles, inner faces
with 3 shallow cavities. Stamens c. % as long as petals, connate for c. % their length;
filaments thick, broadly ovate-orbicular narrowing rather abruptly into subulate apex;
anthers 0.45 mm long. Carpels almost equalling stamens; ovaries ovoid, very fat,
strongly gibbous abaxially, narrowing rather gradually into styles c. 0.3 mm long.
Fruit spherical, c. 15 mm diameter, without obvious protrusions at stalk or stigma
positions. Epicarp black, bluish-pruinose, dull and slightly dimpled, moderately thick
but easily damaged. Mesocarp c. 1.2 mm thick, rather dry, reddish. Endocarp c. 0.1 mm
thick, flexible but weak and membranous, pale red-brown. Seed, subspherical, c. 13 mm
long, 12 mm diameter, surface smooth, dull red-brown. Intrusion penetrating laterally
from slightly below median position for c. % seed diameter, in l.s. narrowly
anvil-shaped and occupying M seed length, in t.s. shallowly 2-lobed at apex and
occupying c. % seed diameter. Embryo lateral (2.30? to 3 o'clock relative to stalk).
(Fig. 4a, 12,13a)

Distribution: restricted entirely to the summit and slopes of the Blackdown Tableland,
which lies at the northern extremity of the Expedition Range in the lower Dawson
River catchment, central eastern Queensland, latitude c. 23°45’S, about 180 km from
the coast near Rockhampton.

Rodd, Revision of Livistona (Arecaceae)

105

Ecology: occurs in great abundance mainly along sandstone cliff-lines, on rocky foot-
slopes below cliffs, in shallow rocky gullies on the plateau, and in deep sandstone
gorges below major waterfalls around edge of the plateau. Most occurrences are at
altitudes between 300 and 600 m, the climate there being somewhat cooler and moister
than that of the surrounding semi-arid lowlands. Grows in moderately tall eucalypt
forest, dominated principally by a stringybark, Eucalyptus sphaerocarpa.

Conservation status: Briggs and Leigh (1996) assign the conservation code 2RC- to
'Livistona sp.l Blackdown Tableland'. However, the major part of its population
appears to be within the Blackdown Tableland National Park, so a coding of 2RCa is
recommended.

Specimens examined: Queensland: Leichhardt: Slopes below N escarpment of Blackdown
Tableland, Blaxell 92 1 & Johnson, 28 Nov 1972 (NSW); Blackdown Tableland, just below escarpment
at NE end (road pass), Rodd 3063-3065 & Jacobs, 11 May 1976 (NSW, BRI, K, BH); Blackdown
Tableland, c. 35 km SE of Blackwater (campsite at old stockyard on Mimosa Creek), c. 8.5 km NNE
of campsite (c. 0.8 km below the entrance gates) on the north facing scarps, Henderson 1180,
Durrington & Sharpe, 14 Sep 1971 (BRI, NSW).

Notes

1. The epithet fulva, Latin for 'tawny', draws attention to the striking golden-brown
indumentum of the leaf undersides, a unique feature among Australian members of
the genus, which otherwise have virtually glabrous leaves except for the ribs. The
colour is hardly evident on older leaves, but on newly unfurled leaves, especially on
semi-juvenile plants, it is immediately noticeable.

2. This species is sharply distinct from all other Australian species and shows no
obvious relationships. Apart from the leaf indumentum, other striking characters are
the presence of a slight abaxial hastula (which may be correlated with the weakly
costapalmate lamina), the very shallow bifurcation of the segments (a neotenic
character?) and the several-toothed apices of the rachis bracts.

3. The existence of this palm seems to have been quite unrecorded until the 1960s. It
was observed but not collected by Mr C.H. Gittins, who in 1961 made some of the
earliest botanical collections on the Blackdown Tableland, and in 1968 by L.A.S.
Johnson and L.J. Pryor who were searching for a new eucalypt discovered by Gittins,
and who thought it might be L. australis. I first saw it in August 1970 when driving up
the forestry road, still under construction, through the northern escarpment; I made a
leaf collection and believed at the time that it might be a new species. The first good
collections (flowering) were made by Henderson et al., collecting for the Queensland
Herbarium in 1971, and I collected good fruiting material in 1976.

4. The distribution of L. fulva is sharply delimited by the sandstone escarpment of the
Blackdown Tableland; it is most abundant on the summit and around the base of the
uppermost line of cliffs on the north and east sides of the plateau; it appears to reach
its lowest altitude in the gorge of Mimosa Creek below the very high waterfall; it is not
evident along the western escarpment, or on the highest parts of the plateau to the
south of Mimosa Creek. It is reasonable to infer that the factors limiting its spread are
rainfall and humidity, with temperature possibly also playing a part. Blackdown
Tableland is an island of high rainfall and cool temperatures in a hot, semi-arid region
with a long and severe dry season. The horizontally bedded sandstones have
doubtless also assisted its survival, the seepage at cliff-bases providing a permanent
source of soil moisture. The evident adaptation of this species to a mesic but relatively
cool environment is uncommon among the Australian Livistona species, though
L. australis appears to have similar requirements.

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9. L. muelleri Bailey

Bailey (1902:1683); Domin (1915: 501); Beccari (1921: 19; 1931: 94, t. 7ii); Burret (1941:
326); Moore (1963: 150; 1973: 60); Covacevich & Covacevich (1978: 92); Tucker (1980);
Johnson (1980:13); Jones (1984:138)

Type: Queensland: Cairns, £. Cowley Oct 1900; holo BRI n.v.; iso FI n.v., photo NSW.

L. humilis R. Br. 'var.' (nov.J F.M. Bailey, Queensland Agric. J. 2: 130 (1898) [same
specimen cited as type of L. muelleri],

L. humilis R. Br. var. sclerophylla Becc., Webbia 5(1): 76 (1921); Ann. Roy. Bot. Gard.
(Calcutta) 13: 99, t. 7iii (1931).

Type: Queensland: Bloomfield River, Miss E. Bauer no. 3; lectotype (here designated) FI
n.v., photo NSW; isolecto 'Miss Ellie Bauer, 12/[18]85' FI n.v., photo NSW. Of the two
sheets of this collection, the one bearing a leaf (as well as a fruiting branch and fruits
in a packet) is selected as lectotype.

?L. humilis R. Br. var. novoguineensis Becc., Webbia 5(1): 76 (1921).

Type citation: 'Nuova Guinea olandese meridionale sul fiume Maroke (Jaheri nell'Erb.
di Buitenzorg).'

Type: not seen.

L. brassii Burret, Notizbl. Bot. Gart. Berlin 12: 309 (1935).

Type citation: 'Brit. Neuguinea: Dagwa, Oriomo River, Western Division'

Type: Papua New Guinea: Western Division: Dagwa, Oriomo River, L.J. Brass 5950,
Feb-Mar 1934; iso BRI.

L. Crustacea Burret, J. Arnold Arbor. 20: 189 (1939).

Type citation: 'Papua: See Daviumbo, mittlerer Fly River'

Type: Papua New Guinea: Lake Daviumbu, Middle Fly River, L.J. Brass 7668, Sep 1936;
holo A, iso BRI.

Trunk solitary, to c. 10 m tall, 15-20 cm diameter at 1 m above ground, often
broadening gently but shortly at base into a cone up to 30 cm diameter on a pedestal
of roots of still larger diameter (almost hemispherical on some old plants). Surface
closely ringed with sheath scars; vertical fissures prominent, some long and deep,
others short and very shallow; basal 2 m or so clothed in persistent, semi-appressed
petiole-stubs often more than 10 cm long (if not burnt).

Crown ± globose, somewhat truncated at base, consisting of c. 25-35 stiffly radiating
leaves, most spreading at slightly above or slightly below horizontal. Ligules
moderately prominent, straw-coloured to pale brown, outer surface sparsely to
densely dotted with chestnut-brown (greying with age) ± circular, thick, corky scales
in pits, their surfaces flush with ligule surface.

Petiole 70-100 cm long, 14-19 mm wide; concave-convex to flattened-triangular in t.s.
Margins bearing minute dark red-brown linear calli; toward base bearing somewhat
sparse shiny blackish, antrorse, patent or retrorse, pungent prickles to c. 4 mm long;
much shorter or sometimes absent on apical half. Surfaces green, undulately striate,
the upper with numerous linear pits to 7 mm long containing conspicuous rows of
thick, corky, antrorsely overlapping, persistent red-brown scales (soon fading to grey),
their surfaces flush but papillose, appearing shaggy, their margins irregularly
erose-fimbriate; lower similar but scale-bearing pits shorter, more numerous, tending
to be aggregated into patches, the scales remaining red-brown longer. Hastula base
truncate to rounded or shallowly 3-lobed or rarely emarginate; rim at 10-60° to costa.

Rodd, Revision of Livistona (Arecaceae)

107

8-20 mm wide, sometimes rolled into a half-cylinder, entire to shortly cuspidate or
2-lobed or very irregular, the necrotic margin narrow to very broad and irregularly
disintegrating.

Lamina 70-80(-90) cm long, 0.20(-0.25) mm thick, weakly costapalmate, ± circular
with narrow sinus at base, chartaceous, brittle and readily splitting, to coriaceous and
moderately tough; flat or slightly contorted with 1 gentle adaxial fold either side of
costa and lower edges involute with only the lowest 1-2 segments either side
resupinate. Segments 28-30 either side of costa; largest segments 25-32 mm wide,
widest at base! free for 50-65% of their length, bifurcated for 5-14% of free length, the
lobes parallel, shortly acute or (occasionally) abruptly curved outward, or more
commonly tapering evenly to a fine bristle-like point. Intersegmental appendages
delicate, brittle, threadlike, to 7 mm long, mostly absent from fully expanded leaves.
Ribs: abaxial ribs in t.s. oblong, rounded to bluntly square-edged, 0.6-0.7 mm thick,
0.9-1.1 mm deep; adaxial ribs oblong, bluntly to quite sharply square-edged, 0.5-0.7 mm
thick, 0.8-1.0 mm deep. Venation: major longitudinal veins 8-9 either side of midrib,
prominent above, slightly less prominent beneath; transverse veins moderately
prominent to somewhat obscure above, more prominent beneath, mostly angled, often
quite steeply, frequently arcuate or sigmoid, frequently continuous across 2-3
longitudinal veins. Surfaces shiny olive green to somewhat grey-green above, below
dull bluish-green with densely crowded stomates and slightly to moderately glaucous
with thin, delicate waxy bloom, glabrous except for scurfy scales along angles of ribs
toward base.

Inflorescences mostly slightly shorter than leaves, sometimes much shorter, fairly stiff
and strongly ascending, somewhat pyramidal in form with stiffly spreading branches.
Partial inflorescences 5-10, reducing markedly in size toward inflorescence apex, the
lowest mostly exceeding 'A of total inflorescence length; each branched to 3, rarely 4
further orders; rachillae 1.5-13 cm long, 0.5-1.2 mm diameter (but only seen
post-anthesis, probably enlarged), these and other axes pinkish-brown to dark
red-brown, the surfaces wrinkled-striate and minutely granular-papillose giving a
slight 'frosted' appearance. Rachis bracts smoothly flattened-cylindrical, to c. 25 mm
diameter, tightly sheathing to somewhat undulately distorted or torn, dark red-brown,
greying with age where exposed to sun; surfaces closely striate with raised veins,
sparsely to densely flecked with pale straw-coloured to silvery appressed scales; bract
apices narrowly ovate-triangular, mostly ± cuspidate with necrotic tip, often with
dense fringe of whitish scales along part of margin. Flower-clusters 1-3 mm apart,
1-3-flowered; cluster axis present only as a slight steplike protrusion on rachilla, to 0.5 mm
long at anthesis, enlarging in fruit to up to c. 1.2 mm long and c. 1.4 mm diameter.
Cluster-bract strongly reflexed, triangular, acute, c. 0.3 mm long. Bracteoles hardly
detectable.

Flowers c. 1.6 mm long, campanulate. Anthopodium c. 0.2 mm long, somewhat
trigonous at base, deeply recessed at point of attachment to cluster axis. Sepals
moderately fleshy but membranous near apex, appressed to petals, slightly concave,
0.8-1.0 mm long, connate for c. 'A their length, broadly triangular, shortly cuspidate at
apex. Petals incurved, 1.3-1.6 mm long, connate for c. A their length, ovate-triangular,
subacute, slightly thickened at apex, auriculate at base with ± rounded auricles
overlapping those of adjacent petals, inner faces with 2 deep, narrow cavities close to
margins. Stamens slightly shorter than petals, connate for c. A their length, the
filaments broadly triangular, strongly shouldered just above base, narrowing rather
abruptly into a slender tapering apex; anthers 0.38 mm long. Carpels from c. % as long
as to almost equal to stamens; ovaries cylindric-ellipsoid, gibbous abaxially, narrowing
somewhat abruptly into slender curved style 0.3-0.5 mm long.

Fruit broadly ellipsoid, 10.5-12 mm long, 8.5-10 mm diameter; apex rounded, base
rounded or broadly conical often with small nipple at stalk position. Epicarp dark
red-brown to black, often pruinose, moderately thick but not tough. Mesocarp c. 1.0 mm

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Telopea 8(1): 1998

thick, oily and yellowish when fresh, red-brown and granular dried. Endocarp c. 0.15 mm
thick, smooth, cartilaginous but easily broken. Seed subspherical, 8-9 mm long, 7.5~8.5 mm
diameter, slightly flattened ventrally, dull red-brown, smooth. Intrusion narrow,
± cylindrical, penetrating from a lateral position for up to % seed diameter, in l.s.
slightly swollen or irregularly lobed at apex, occupying from less than 'A to % of seed
length, in t.s. usually with 2 erect to diverging apical lobes, occupying c. 'A or less of
seed width. Embryo lateral, at or somewhat below mid-point of seed (3 to 4 o'clock
relative to stalk). (Fig. Id, 2c, 4b, 13b, 15a)

Distribution: far northern Queensland, from Cape York south to about 17°45' (S of
Innisfail) on east coast, and south on the west coast of Cape York Peninsula at least to
mouth of Edward River, possibly to Mitchell River. South of Cooktown it is confined
strictly to coastal lowlands, but further north in the Peninsula it extends further
inland. Also in Papua New Guinea, in lowlands around Gulf of Papua, Fly River
region and probably also in adjacent parts of Irian Jaya.

Ecology: in North Queensland this species occurs in greatest numbers in poor
woodland communities dominated by Melaleuca viridiflora and Eucalyptus alba on flat,
seasonally swampy interfluves on sandy soils of low fertility frequently with a clay
hardpan below the surface. It may also occur on the outer margins of gallery rainforest
or, in northern Cape York Peninsula, in or adjacent to patches of vine forest. In Papua
New Guinea it is recorded as 'solitary or in small groves on rainforested shores of lake
and extending less commonly to the dry savannahs' and 'common palm of savannah
forest on open grass slopes'.

Conservation status: not considered at risk.

Specimens examined: Queensland: Cook: Jardine River, N bank, near crossing of Peninsula
Development Road, Hind 576, Oct 1974 (NSW); Musgrave Station, Hind 749, Sep 1979 (NSW); 14 km
WNW of Cooktown on Mclvor River Road, Hind 748, Sep 1975 (NSW); N side of Endeavour River
near Jones Lagoon, 11 km WNW of Cooktown, Rodd 3159 & Jacobs, 21 May 1976 (NSW, BRJ, K, BH);
Bloomfield River road c. 3-5 km N of Aylon, Rodd 3170, 22 May 1976 (NSW, BRI, K, BH); Craiglea,
Bruce Highway near Port Douglas turn-off. Hind 226, Aug 1972 (NSW); Trinity Beach area, north
of Cairns, Maconpchie 1717,22 May 1972 (DNA).

Papua New Guinea: See type citations of synonyms L. brassii and L. Crustacea. In field notes on his
New Guinea collections L.J. Brass describes it as 'Solitary or in small groves on rainforested shores
of lake [L. Daviumbu, middle Fly R.], and extending less commonly to the dry savannahs' and
'Common palm of savannah forest on open grass slopes, alt. 45 m.' [Dagwa, Oriomo R.]

Notes

1. A very distinctive species, not likely to be confused with any other. In the field it is
unmistakable among Australian species for its very stiff, short-petioled, almost flat
leaves, radiating horizontally. Herbarium specimens are most remarkable for their
petiole indumentum, consisting of rows of very corky brownish scales sunken in long
pits, their surfaces flush with the petiole surface.

2. Chapman (1991), in his listing of L. muelleri Bailey, comments ‘nom. illeg. non
Wendl.', referring to a prior name supposedly published by Wendland in Kerchove
(1878). But on examining this publication we find that it is merely a name mentioned
in passing, one item in a brief summary of all the palms known from Australia, thus:
'A la premiere region [i.e. "la region tropicale"] apartiennent les Calamus caryotoides et
radicalis, ... Livistona humilis, Ramsayi, Leichardti et Miilleri, Licuala Miilleri et Cocos
nucifera.’ It is clear that L. muelleri H. Wendl. is a nomen nudum and of no standing.

3. Burret (1941), in discussing the likelihood of his L. brassii proving conspecific with
L. muelleri, drew attention to the petiole scales. Oddly, though, he drew no comparison

Rodd, Revision of Livistona (Arecaceae)

109

between either of these species and his later-described L. Crustacea, the description of
which follows closely that of L. brassii.

4. L. muelleri appears to have some affinities with L. humilis and L. eastonii, sharing with
them shallowly bifurcated leaf-segments and distinctive rows of red-brown basal-cell
masses of fallen scales on the petioles. The corky, thick scales of L. muelleri, though, set
it apart sharply from these species. In another direction, it may have an affinity with
L. benthamii, sharing with it a number of inflorescence, flower and fruit characters.

5. One collection ( Rodd 3170) is remarkable for the inflorescence having empty tubular
bracts additional to the prophylls sheathing the bases of its lower partial
inflorescences, a departure from the virtually standard inflorescence pattern in
Australian Livistona. However, this does not appear in other collections and further
field observation is required to ascertain whether it is characteristic of a population.

6. Field observations indicate that in a population of this species a large proportion of
plants do not produce fruit, evidenced by their old inflorescences lacking thickened
fruiting cluster axes. Good collections of flowering specimens may reveal whether or
not there is any sexual dimorphism evident in the flowers. It is possible even that the
bract character described above may be correlated with sexual state.

7. The type material of Beccari's L. humilis var. sclerophylla in FI consists of two sheets,
the first consisting of a whole leaf, partial inflorescence and a packet of seeds; the
second of only partial inflorescence and a packet of seeds. Their labels are essentially
the same, except that the second bears the date '12/85' while the first has no date. As
indicated above, the first is here chosen as lectotype. Beccari presumably failed to
equate this collection with Bailey's L. muelleri (of which he had adequate material, also
with mature fruit) because he was unaware that the leaf was juvenile and hence much
smaller than the L. muelleri leaves he had available. Tire Bauer specimen s fruits are
also at one extreme of the length : diameter ratio found in L. muelleri, whereas the type
of the species has fruits tending towards the other extreme.

8. Hind 748 bears the note 'fruits blackish, flesh yellow smelling strongly of banana and
very oily'. The mature fruits of this collection were the chief source for the fruit and
seed description here.

10. L. humilis R. Br.

Brown (1810: 268); Martius (1838: 239); Wendland & Drude (1875: 231); Beccari (1921:
19; 1931: 95); White (1988: 23, 25).

Type: Gulf of Carpentaria, Island 's' [= Morgans Island, Blue Mud Bay], R. Brown s.n.,
20.1.1803 (Bennett No. 5796); holo BM, photo NSW, iso K, FI n.v., photo NSW.

Saribus humilis (R. Br.) O. Kuntze, Rev. Gen. PI. 2: 736 (1891).

L. leichhardtii F. Muell., Fragm. 8: 221 (1874).

Type citation: 'In Arnhemia et regionibus vicinis, ubi jam Leichhardtio notata.

Lectotype (here chosen): Arnhem's Land, McAdam's Ranges, F. Mueller s.n., 1855;
holo MEL.

Trunk solitary, to 7 m tall, 5-8 cm diameter, broadening slightly right at ground.
Surface patterned with petiole scars, sheath scars not prominent; vertical fissures
sparse, irregular, narrow; most of trunk bearing appressed petiole-stubs to c. 3 cm long.
Crown globose, very open, consisting of c. 8-15 stiffly ascending to spreading and
slightly arching leaves. Ligules not very prominent, disintegrating at an early stage
into a network of fibres with only a narrow entire rim, the outer surface pale brown
with raised veining, clothed with appressed fimbriate white scales.

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Telopea 8(1): 1998

Petiole (30-)40-60(-70) cm long, (7-)8-14 mm wide; flattened-triangular to
transversely obtrullate in t.s. with very rounded keel. Margins armed with prickles
interspersed with irregular small blunt teeth and minute linear brown calli; prickles
up to 3 mm long, narrow, very pungent, mostly dark-tipped, all antrorsely falcate
except for a few retrorsely sigmoid ones near base, sometimes few or absent on upper
third of petiole but mostly present on whole length. Surfaces green to brownish or
somewhat reddish, obscurely striate, densely and conspicuously speckled with small
dark red-brown linear markings representing sunken basal-masses of early-deciduous
appressed white fimbriate scales, fairly uniformly distributed over both surfaces.
Hastuln base truncate, shortly and bluntly apiculate; rim at 10-40° to costa, 3-8 mm
broad, convex, semicircular to irregularly lobed or split, the necrotic margin very
variable, broad or narrow, continuous or interrupted.

Lamina (28-)35-55 cm long, 0.2-0.3 mm thick, moderately costapalmate, ± circular,
truncate at base or with broad to fairly narrow sinus; chartaceous-coriaceous,
somewhat brittle and springy, only slightly contorted with one adaxial undulation
either side of the often strongly deflexed costa, base on either side involute but
segments not resupinate. Segments 15-22 either side of costa; largest segments 12-32 mm
wide, free for 60-87% of the lamina length, bifurcated for 49-89% of free length, the
lobes parallel to slightly divergent, tapering at first convexly and then concavely to a
fine but rarely threadlike point. Intersegmental appendages very fine, threadlike, to 3 cm
long on younger leaves, soon broken off. Ribs: abaxial ribs in t.s. oblong-elliptic,
bluntly to sharply square-edged, narrowest at junction with lamina, up to 0.8 mm
thick, 1.0 mm deep; adaxial ribs similar but slightly smaller. Venation: major
longitudinal veins 4—8 either side of midrib, very prominent above, slightly less
prominent beneath; transverse veins somewhat obscure above, prominent beneath,
mostly near-orthogonal and discontinuous, on some specimens tending to more
steeply-angled and continuous across several longitudinal veins. Surfaces glossy
green or slightly brownish green, glabrous except on ribs, or lamina as well as ribs
bearing numerous red-brown fine spots, as on petiole but not so elongated, pustular
rather than sunken, mainly on veins but a few between veins; denser scurfy white
hairs present on rib bases close to hastula, but disappearing at an early stage.
Inflorescences very markedly dimorphic; those on fruit-bearing plants twice as long
as leaves (to c. 2 m), straight and strongly ascending, completely unbranched except
for the apical 25-35 cm which is strongly deflexed and represents a single (terminal)
partial inflorescence; those on non-fruiting plants up to about 50% longer than leaves,
ascending but usually arching in upper part, with 4-7 subequal partial inflorescences,
these branched to 3 further orders; rachillae 2.5-12 cm long, 0.4-0.7 mm thick, these
and other axes obscurely striate and sparsely to densely bristly-pubescent with white
hairs c. 0.1 mm long or shorter. Rachis bract of fruit-bearing plants complemented
basally by a succession of 5-8 empty peduncular bracts, these (and rachis bract)
flattened-bicarinate, 10—15 mm wide, tightly sheathing, straight and regular, rigidly
coriaceous, green (or reddish brown toward apex), finely striate with alternating
coarser and finer wrinkles, densely to very sparsely but evenly scurfy-pubescent with
small, appressed, longitudinally oriented, transparent laciniate scales; rachis bracts of
non-fruiting plants similar but slightly broader and looser; bract apices narrowly
triangular-ovate, long-acuminate, necrotic, somewhat warped, looser on non-fruiting
plants. Flower-clusters 1-3 mm apart, 2-4-flowered; cluster axis (on fruit-bearing
plants) a mere raised bump at rachilla node, or (on non-fruiting plants) knob-like, to
c. 0.7 mm long. Cluster-bract ovate-triangular to 0.5 mm long, membranous except at
base, often discernible only as a narrow rim by anthesis. Bracteoles usually 1 per
flower, shorter and blunter than cluster-bract, or (on non-fruiting plants) mostly
not discernible.

Flowers c. 1.8 mm long, ± globose to cylindric-globose. Anthopodium extremely short,
concave at base with a downward-projecting bump below each sepal. Sepals strongly

Rodd, Revision of Livistona (Arecaceae)

111

concave and appressed to petals, membranous near apex, thickening toward base,
c. 1.3 mm long, connate for c. A their length, orbicular-ovate, apex very shortly and
± bluntly cuspidate. Petals moderately concave, thick and fleshy, c. 1.8 mm long,
connate for c. A their length, very broadly triangular-ovate, bluntly acute and
shallowly cymbiform at apex, broad-based and auriculate above base, with minute
triangular auricles; inner faces with well marked but not deep cavities. Stamens at
least % as along as petals, connate for c. 'A their length; filaments very thick, broadening
apically with well marked high shoulders, abruptly and rather shortly cuspidate;
filaments in non-fruiting plants shorter and broader with more rounded shoulders;
anthers on fruit-bearing plants 0.35 mm long, the loculi apparently lacking pollen; on
non-fruiting plants c. 0.4 mm long, the loculi containing pollen. Carpels slightly
shorter than stamens; ovaries broadly ovoid, abaxially strongly gibbous, tapering
gradually into slender styles c. 0.4 mm long; ovaries in non-fruiting plants globose,
very plump, with shorter styles and containing apparently well-formed ovules.

Fruit ellipsoid-pyriform to obovoid, sometimes flattened or slightly concave adaxially
and gibbous abaxially, 11—19 mm long, 8-10 mm diameter. Epicarp smooth, dull
pinkish-purple ripening to shiny black. Mesocarp c. 0.6 mm thick, soft and pulpy.
Endocarp to 0.2 mm thick, quite hard, the outer surface with a number of longitudinal
ridges. Seed ellipsoid to subglobose, flattened adaxially, 6-7 mm wide. Intrusion small,
penetrating from a lateral position for only 'A width of endosperm, in I.s. with short
lobes pointing toward base and apex and occupying only c. 'A of seed length, in t.s.
usually narrow with 2 slightly diverging lobes. Embryo lateral but distinctly below
mid-point of seed. (Fig. 4c, 13c, 14)

Distribution: Northern Territory, the 'Top End', virtually ubiquitous in suitable
habitats north of about the Fitzmaurice River in the west and the Roper River in the
east, though apparently absent from the Katherine-Mataranka area.

Ecology: appears to be restricted to flat or gently undulating interfluvial areas on deep
sandy soils, usually with pisolithic ironstone gravel, or on bauxite, in open eucalypt
forest nearly always dominated by Eucalyptus tetrodonta, usually with E. miniata
also present.

Conservation status: not considered at risk.

Selected specimens examined: Northern Territory: Oenpelli, Speclit 1221, 19 Oct 1948 (AD); c. 80 km
E ol : Oenpelli, Maconochie 1598,28 June 1972 (DNA, AD, PERTH); Darwin & Gulf: Arnhem's Land,
McAdam's Ranges, Mueller s.n., 1855 (MEL); Macmillan Road, 8 miles [13 km] SE Darwin,
Chippendale NT4391, 24 May 1958 (DNA, NSW); Cannon Hill Airstrip (near Darwin), Marlensz &
Schodde AE617, AE618, 30 Jan 1973 (DNA, CANB); Stuart Highway c. 35 km N of Batchelor, Eichler
17887,17888, 2 July 1964 (AD); Stuart Highway, c. 160 km SSE of Darwin, Jackson 1050, 4 June 1967
(AD); Melville Island, Spencer s.n., July-Aug 1911 (NSW, MEL); Strauss Airstrip, 28 miles [45 km]
S Darwin, Must 957, 8 Mar 1972 (DNA, CANB); Daly Range, 14 km S of Manton R on Stuart
Highway, Rodd 2910, 2911, 30 Oct 1974 (NSW, DNA, K, BH); 16 miles [26 km] NE of Tipperary
Homestead, Lazarides 6687, 27 July 1961 (CANB, NSW); c. 4 miles [6.4 km] W of Pine Creek
Township, Lazarides & Adams 227 , 15 Mar 1965 (CANB); Macdonnell Airstrip, 67 miles [108 km] N
of Katherine, Maconochie 1300, 1971 (DNA, CANB, K); About 2 miles [3.2 km] NW Caiman Ck,
Byrnes 1014 & Maconochie, 27 Sep 1968 (DNA, NSW); Mun marl ary Shi., Latz 957, 10 May 1973
(DNA, CANB); 10 miles [16 km] W Jim-Jim, Byrnes 1725, 21 Sep 1969 (DNA, CANB); Obiri Rock,
Maloney 2, 18 July 1975 (NSW); 36.3 miles [58 km] N Oenpelli, Chippendale NT8108, 15 July 1961
(DNA, NSW); c. 20 km SE of Maningrida, Rodd 2923,2924, 31 Oct 1974 (NSW, DNA, K, BH); 36 miles
[58 km] N Wilton R.-Bulman Crossing, Maconochie 1457, 15 June 1972 (DNA, CANB); Elcho
Island, Maconochie 2179, 12 July 1975 (DNA, NSW, CANB, K); Wessell Islands, Latz 3470, 10 Oct
1972 (DNA, CANB); c. 1.5 km W B.H.P. Camp [near Caledon Bay], Maconochie 1560, 22 June 1972
(DNA, CANB, NSW, K); 4 miles E of Lake Evella turnoff, Maconochie 1513, 18 June 1972 (DNA,
PERTH); Gove Peninsula, halfway between Gove Airport and Nhulunbuy Township, Rodd 2926,
2927, 2928, 2 Nov 1974 (NSW, DNA, CANB, K).

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Telopea 8(1): 1998

Notes

1. A distinctive species, easily recognised by its small stature, the smallest of any
Australian species. Herbarium material might possibly be confused with L. inermis,
from which it differs in its non-pruinose leaves, profuse brown linear markings on
both sides of petiole and frequently on leaf-blade, inflorescences greatly exceeding
leaves, and bristly-pubescent rachillae. Its unbranched 'female' inflorescences are a
striking feature, not found in any other species.

2. Brown's brief diagnosis of this species is barely adequate to allow its recognition,
but as well as the description there is Bauer's beautiful plate, correctly captioned with
the correct name (presumably with guidance from Brown) and published eventually
in Martius's (1838) Historic! Natumlis Palmarum. The photograph of Brown's type
specimen published at folio size in Beccari's 1931 monograph also allows no doubt as
to the application of the name. Beccari interprets L. humilis correctly but seems to find
some confusion in the Bauer illustrations (of L. humilis and L. inermis) in Martius. To
me these illustrations both appear correctly named and quite unambiguous.

3. Mueller makes no mention in his protologue of L. leichhardtii of the specimen that he
himself collected on the McAdam Range (near the Fitzmaurice River), here accepted
as type of that name. It is a very adequate specimen of L. humilis, which can be seen in
MEL, labelled L. leichhardtii in his own hand. Mueller later (1878) remarked that his
L. leichhardtii probably included Brown's L. inermis and L. humilis, implying that
Brown's descriptions were too vague for definite application of the names.

4. Although in many Australian Livistona species there is a strong suspicion of
functional dioecism or androdioecism, accompanied by slight dimorphism in
inflorescence structure, it is only in L. humilis that this is expressed strongly and
unequivocally. The briefest inspection of any stand reveals two extraordinarily
different inflorescence types, segregated on different plants, and (to the best of my
knowledge) fruit set occurs on only one of these. This is the inflorescence type with no
lateral partial inflorescences, only a solitary terminal one on a very long peduncle
sheathed in a succession of peduncular bracts. The non-fruiting plants have
inflorescences which, though small and slender, conform to the 'standard' Livistona
type with a succession of short, subequal partial inflorescences. The flowers in these
two types look rather different but a systematic survey of their structure reveals only
slight differences, principally empty anther locules on the fruiting plants; the carpels
and ovules in both types appear normal, though ovule structure has not been
examined at high magnification. Clearly there is scope for a fuller investigation of the
reproductive behaviour of this species, indeed of all Livistona species.

5. White (1988) illustrates an instance of inferred hybridism between L. humilis and
L. inermis, though makes no mention of this in his text. The two are sympatric over
most of the range of L. inermis, though generally ecologically segregated. I have found
a few vegetative specimens difficult to assign to one species or the other, and so believe
such hybrids may well exist. Again, further investigation is needed.

6. See also discussion under L. benthamii, L. inermis and L. eastonii.

11. L. eastonii C.A. Gardner

Gardner (1923: 36); Beard (1976: 183); Hnatiuk (1977); Jones (1984: 133); Wilson (1992:
1249. fig. 353A).

Lectotype (here designated, as no specimen was actually cited by Gardner): Western
Australia: Lower King Edward River, C.A. Gardner 1544, 22 Aug 1921 (PERTH
[specimen with a whole inflorescence and a juvenile leaf, bearing a 'holotype' sticker]);
isolecto PERTH [specimen with a whole leaf blade (but no petiole) and small piece of

Rodd, Revision of Livistona (Arecaceae)

113

inflorescence, with determination slip in Gardner's (?) hand and a red 'isotype'
sticker], CANB, K, BH, B.

Trunk solitary, to 11 (?20) m tall, c. 12 cm diameter, broadening abruptly close to
ground into a broad conical or bulbous base up to 30 cm diameter. Surface toward base
without noticeable sheath scars; vertical fissures deep, prominent, especially toward
base; petioles or sometimes whole dead leaves persistent up to a height of 2.5-3 m, but
mostly seen burnt off leaving regular spiral rows of appressed stubs 2-10 cm long;
leaves shed higher up leaving prominent, closely crowded scars.

Crown somewhat elongated, open, consisting of c. 10-15(-30?) rather stiff leaves, all
but the youngest few drooping due to bending of petioles. Ligules not very prominent,
soon tearing into a net of narrow criss-crossing ribbons, pale red-brown ageing to
silvery grey, with dense, appressed, transparent, very short scales.

Petiole (55-) 65-80(-90) cm long, 14-20 mm wide; flattened-triangular to broadly
obtrullate or concave-convex in t.s. Margins armed with mixed antrorse, patent and
retrorse, narrow, mostly blunt prickles to c. 4 mm long; shorter, blunter and sparser
near petiole apex; minute elongated brown calli also present. Surfaces pale yellowish
green, irregularly but sometimes markedly striate, with ± profuse, evenly distributed
linear red-brown to purple-black rows of basal-masses to 2 mm long in shallow pits,
bearing early-deciduous, often tightly twisted white or pale brown scales. Hnstuln
variable; base narrowly to broadly triangular or shortly and asymmetrically 3-lobed;
rim at 20-50° to costa, fairly smooth and flat, to c. 10 mm wide with broad but often
irregular necrotic margin, often disintegrating in patches.

Lamina 60-90 cm long, 0.25-0.30 mm thick, strongly costapalmate, truncate at base
with all segments forward-pointing, coriaceous, tough and moderately flexible; only
moderately contorted with 1 adaxial undulation either side of the slightly to strongly
deflexed costa, sometimes shortly involute at base with lowest 1-2 segments either
side resupinate. Segments 20-27 either side of costa; largest segments 22-31 mm wide,
free for 50-90% of their length, bifurcated for 49-63% of free length, the lobes parallel
or slightly converging or diverging, not or barely drooping, evenly tapering to fine,
acute, somewhat rigid apices mostly shortened by early necrosis. Intersegmental
appendages fine, brittle, threadlike, early-deciduous, occasionally up to 4.5 cm long,
mostly absent on mature leaves Ribs: abaxial ribs in t.s. oblong to trapeziform
(narrowing to lamina), square-edged, to 0.9 mm thick, 1.3 mm deep; adaxial ribs
rectangular to somewhat tapering or oval, not generally square-edged, to 0.8 mm
thick, 1.2 mm deep. Venation: major longitudinal veins 5-9 either side of abaxial rib,
evident but not always prominent above, ± obscure beneath; transverse veins obscure
to evident above, evident to moderately prominent beneath, mostly angled, many
continuous across several longitudinal veins and frequently arcuate. Surfaces pale
greyish-green, somewhat yellowish above, moderately pruinose on both sides but
with a more prominent and softer waxy bloom on underside; glabrous except for ribs
which bear rows of orange-brown calli mainly along angles.

Inflorescences approximately equalling leaves, steeply ascending but with pendulous
branches. Partial inflorescences 5-6, each branched to 4 further orders, the rachillae
0.2-8 cm long, 0.3-0.5 mm thick; all axes glabrous, wrinkled-striate, pinkish-brown in
bud stage soon changing to pallid whitish. Rachis bracts cylindric to moderately
flattened, bicarinate, to c. 20 mm diameter, smooth and tightly sheathing to somewhat
looser and warped, stiff and parchment-like, initially pinkish straw-coloured but soon
ageing to deep red-brown, closely striate, bearing closely appressed, fimbriate,
translucent white or pale brown scales giving a densely flecked appearance; bract
apices long-triangular, acute to slightly acuminate, frequently with scaly-tomentose
margins. Florver-clusters 0.5-3.5 mm apart, l-3(-4)-flowered; cluster axis + knob-like or
forked, 0.1-0.7 mm long. Cluster-bract broadly triangular, c. 0.4 mm long,
membranous, its apex initially delicately acuminate-fimbriate. Bracteoles 1-3,
resembling cluster-bract but often shorter.

114

Telopea 8(1): 1998

Flowers 1.6-1.9 mm long, campanulate to rotate. Anthopodium 0.2 mm long, not
distinct from base of calyx, bulging downward into 3 fleshy knobs below sepals, base
recessed at point of attachment to cluster axis. Sepals loosely appressed to petals,
moderately concave, somewhat fleshy right to apex, 0.6-1.0 mm long, connate for A
their length, broadly ovate-triangular, acute. Petals rather thin and tough, 1.6-1.9 mm
long, connate for Z-'A their length, narrowly oblong-ovate to triangular-ovate, rounded
toward the broadly acute apex, minutely auriculate at base with auricles usually
rounded, retrorse, overlapping; inner faces lacking obvious cavities though with
shallow longitudinal ridges. Stamens %-%> as long as petals, connate for A-Z their
length; filaments moderately thin, broadly triangular-ovate, tapering smoothly to a
slender, elongated point, not obviously shouldered; anthers 0.40-0.42 mm long,
containing pollen except in 1 specimen (Rodd 2887) from fruit-bearing plant, in which
cells are consistently narrow and empty. Carpels from % as long to virtually as long as
stamens; ovary broadly ellipsoid to cylindric-ellipsoid, strongly gibbous abaxially and
sometimes apically also, rather abruptly narrowed into slender, ± straight style
0.25-0.5 mm long.

Fruit obovoid-pyriform to ellipsoid, 13.5-16 mm long, 8-9 mm diameter, straight to
slightly curved but not flattened adaxially, narrowly conical to almost rounded at base,
rounded to bluntly apiculate at apex. Epicarp purple-black, smooth and glossy when
fresh, rather coarsely wrinkled when dry, sometimes pruinose, moderately thick and
tough. Mesocarp 1.0-1.3 mm thick, soft and pulpy. Endocarp 0.15 mm thick, quite
smooth, adhering closely to seed. Seed obovoid to ellipsoid, 10-13 mm long, 7 mm
diameter, barely flattened adaxially. Intrusion penetrating from lateral position for 'A-%
seed width, in l.s. irregularly lobed with short lobes directed toward seed apex and
base, occupying A-A seed length, in t.s. narrow and with 2 erect lobes. Embryo lateral,
almost exactly midway between seed base and apex. (Fig. lc, 4d, 16,17a-b)

Distribution: Western Australia: northern Kimberley Region, apparently confined to
the Mitchell Plateau south of Admiralty Gulf and the region to the east and south-east
as far as the lower King Edward River and the middle-upper Drysdale River.

Ecology: restricted to more or less level plateau areas on deeply weathered soils with
abundant lateritic gravel; on such sites the most robust plants occur in gully heads or
moist depressions, but the species appears to be absent from immediately adjacent
sites on sandstone such as gorges and cliff-lines where no laterite is developed.
Gardner describes it as occurring also on 'basalt plains and hills'. On its preferred sites
L. eastonii forms huge populations, with all age-classes well represented, and is most
frequently co-dominant with Eucalyptus miniata and £. tetrodonta, or sometimes with
the latter only, usually forming a community with few other conspicuous trees or shrubs.

Conservation status: not considered at risk.

Specimens examined: Western Australia: Gardner: 15 miles [24 km] S of Admiralty Gulf,
Kimberley, Gardner 972, 8 July 1921 (PERTH); 75 miles [ 121 km[ NNW of Gibb River Station, Speck
4949, 7 Sep 1954 (CANB); c. 15 miles [24 km] N of Amax Bauxite Camp, Mitchell Plateau,
Maconochie 1281, 1 May 1971 (DNA, CANB, K, PERTH); 15 miles [24 km] S of mouth of Mitchell
River, Schulze s.n., 20 Jan 1973 (PERTH); Mitchell Plateau, N Kimberley, Beard 6992, 7 June 1974
(PERTH, NSW); Amax Bauxite Camp, Mitchell Plateau, 20 km SSW of head of Port Warrender, Rodd
2888, 26 Oct 1974 (NSW, PERTH, K, BH); Mitchell Plateau, 25 km SSW of head of Port Warrender,
Rodd 2887, 26 Oct 1974 (NSW, PERTH, K, BH); woodland S of Airfield Swamp, Mitchell Plateau,
West Kimberley, Hnatiuk MP36, 13 June 1976 (PERTH); N end of mnway, Mitchell Plateau Airfield,
Hnatiuk MP28, 13 June 1976 (PERTH); 2.9 km SE of Mitchell Plateau Mining Camp, George 14503,
23 Apr 1977 (PERTH, CANB, LE); 100 miles [c. 161 km] S of Kalumburu, Hutchinson 84, 19 July 1970
(PERTH); 25 miles [40 km] N of New Drysdale Station Homestead, Maconochie 1228, 28 May 1971
(DNA, PERTH, CANB, MEL, NSW); c. 12 km N of Drysdale River crossing on Kalumburu road,
Symon 10254, 30 May 1975 (NSW); Doongan Station, Crossland Creek at Gibb River - Kalumburu
road crossing, Telford 6096 & Butler, 24 July 1977 (CBG, PERTH, NSW); 11 km N of Doongan H.S.

Rodd, Revision of Livistona (Arecaceae)

115

turn-off from Gibb R-Kalumburu road, George 15220 ,25 June 1978 (PERTH, CANB, NSW, AD, K);
8 km N of Doongan H.S. turn-off from Gibb R-Kalumburu road, George 15221, 25 June 1978
(PERTH, CANB, NSW, K).

Notes

1. A very distinctive species, instantly recognisable in the field though not always so
clearly from herbarium specimens. Geographically it is very nicely delimited, showing
overlap only with L. lorophylla, but they are not ecologically sympatric.

2. Gardner, whose botanical travels in the northern Kimberley in 1921 may still be
unrivalled in their coverage of some regions, described L. eastonii in 1923 without Latin
description and without designating a type but these were not mandatory under the
ICBN at that time. Gardner named the species in honour of William R. Easton,
surveyor, leader of the Kimberley Expedition of 1921.

3. Of the two collections in PERTH, the lectotype ( Gardner 1544, Lower King Edward
River) is more complete than the other (Gardner 972, 15 miles south of Admiralty Gulf)
which consists only of pieces of petiole and of inflorescence in bud. Both localities fall
within the geographical range he specifies, though the Admiralty Gulf one is
admittedly more explicitly stated.

4. Taken literally, 'Lower King Edward River' would refer to the area around the
present settlement of Kalumburu, but L. eastonii is not present there; more likely it
means somewhere closer to the middle reaches of this river, paralleling the Lawley
River, which ties in with present-day knowledge of this species' distribution.

5. Observations by L.A.S. Johnson (pers. comm.) were that travelling north in the
Kimberleys, L. eastonii is first recorded close to 'Doongan' and thence sporadically
toward the Mitchell River turn-off but cuts out before the turn-off; it is thereafter
absent on the Kalumburu road and east of Kalumburu toward Forrest River
Aboriginal Reserve; on the Mitchell River fork it appeared again close to the King
Edward River crossing and was present most of the way from there to 'Mitchell River'
Station. He noted its presence in a number of eucalypt associations additional to its
usual occurrence in the £. tetrodonta-E. miniata association.

6. L. eastonii appears to be most closely related to L. humilis and L. muelleri.
Morphological features shared by this trio include relatively shallow lobing of the
leaves with stiff, non-attenuate apices to the secondary lobes; lamina with very few
undulations; petiole with early-deciduous scales leaving profuse and conspicuous
elongated, dark-coloured basal-cell masses giving a distinctive surface-texture; rachis
bracts relatively narrow, smooth and tight with very regular, shortly cuspidate apices,
thin and tough in texture with silvery flecking of closely appressed translucent scales;
flowers and flower-clusters virtually sessile, the clusters not elongated; fruits
elongated. Within this group L. eastonii is readily distinguished by its strongly
pruinose leaves, their size intermediate between those of the other two, as is the trunk
diameter. Geographically and ecologically these three species show a striking pattern:
in the three major projecting land-masses of far northern Australia they extend south
to roughly similar latitudes (c. 15° in northern W.A., 14° in N.T., 17° in Qld), and occur
on similar topography and soils, i.e. flattish interfluvial areas on deep sandy or lateritic
soils, in this respect contrasting sharply with geographically sympatric species in all
three regions. They show a strong association with Eucalyptus tetrodonta, though in
Cape York Peninsula L. muelleri extends further southeast than this eucalypt. Such
habitats are arguably subject to a greater fire frequency than any other Livistona
habitats and the degree of fire-adaptation of these three species appears to reflect this:
all seem to have a very prolonged geophytic juvenile phase, frequently seen sprouting
new leaves after fires; but all Australian Livistonas appear fire-adapted to a
considerable degree.

116

Telopea 8(1): 1998

12. L. inermis R. Br.

Brown (1810: 268); Martius (1838: 239, t. 145, 146 iv-vi); Beccari (1921: 20; 1931: 102,
t. 7iv); Blake (1954:128); White (1988); Birch & Dowe (1989:11)

Type: Gulf of Carpentaria, Island h [= North Island, Sir Edward Pellew Group], Brown
[Bennett No. 5795], 16(?) Dec 1802; holo BM (mounted on 5 sheets), photos NSW; iso
FI, photo NSW.

Saribus inermis (R. Br.) O. Kuntze, Rev. Gen. PI. 2: 736 (1891).

Trunk solitary, to 10 m tall, 6-10 cm diameter, broadening near base into a short cone
up to c. 15 cm diameter; or sometimes branching from base to produce clump of up to
c. 6 aerial stems of mixed height together with many unelongated sucker shoots with
juvenile leaves, springing from root mass up to c. 70 cm wide at ground. Surface of
basal 1-2 m of trunk bearing closely appressed petiole stubs to 5 cm long; leaves shed
cleanly higher up leaving regular pattern of petiole scars but only rather obscure rings.
Crown globose, very sparse, consisting of c. 30 arching to drooping leaves with
somewhat drooping segments. Ligules straw-coloured, clothed with very closely
appressed transparent scales.

Petiole 65-95 cm long, 6.5-10.5 mm wide, concavo-convex to transversely obtrullate
in t.s., with slightly rounded keel. Margins obscurely crenate with minute pale brown
calli or variably armed with mixed antrorse and retrorse, narrow, falcate or sigmoid,
pungent, mostly dark purplish prickles to 5 mm long, if present on apical half then
much shorter, patent, blunt, not dark-coloured. Surfaces dull pinkish-purple especially
toward base, variably pruinose (sometimes thickly), somewhat striate, upper
glabrous, lower sparsely to densely clothed with closely appressed to loose, scurfy,
laciniate white to pale orange-brown scales. Hasiula irregular, variable; base shortly
3-lobed to truncate-apiculate to shallowly V-shaped to semicircular; rim at 20-90° to
costa, 3-7 mm broad, sometimes contorted, necrotic margin narrow and entire to
broad and interrupted.

Lamina 33-65 cm long, 0.25-0.40 mm thick, weakly to moderately costapalmate,
approximately semicircular to almost circular, truncate to broadly notched at base,
coriaceous, flexible; slightly to strongly contorted with 1 adaxial undulation either side
of the barely to strongly deflexed costa and base on either side involute with
lowermost 2-6 segments resupinate. Segments 12-24 either side of costa; largest
segments 11-21 mm wide near point of bifurcation, free for (80-)91-97% of their
length, bifurcated for (70)73-84% of free length, the lobes parallel or slightly diverging,
usually broadening above point of bifurcation, 6-13 mm wide at widest point, their
apices tapering evenly to a fine point. Intersegmental appendages to c. 5 cm long,
delicate and hairlike, soon broken off. Ribs: abaxial ribs in t.s. ± square but not very
sharp-edged, to 0.7 mm thick, 0.9 mm deep; adaxial ribs similar, to 0.6 mm thick, 0.8 mm
deep, or in some specimens hardly protruding above lamina. Venation: major
longitudinal veins 8-10 either side of abaxial rib, prominent on both surfaces;
transverse veins obscure above, prominent below, from orthogonal to steeply angled,
frequently continuous across several longitudinal veins. Surfaces concolorous, green
to strongly greyish-pruinose, glabrous except for tufts of dense, scurfy, whitish to
straw-coloured scales on bases of ribs.

Inflorescences shorter than to barely equalling petioles, but broader than long, the
peduncle to c. 30 cm long and the rachis nodes only 4-10 cm long. Partial
inflorescences usually 3, the lowermost ± equalling the remainder of the inflorescence,
branching to a further 3 orders; rachillae 1-7 cm long, 0.3 mm thick, these and other
axes wrinkled-striate, bright creamy yellow at anthesis, later darkening to
reddish-brown grading to whitish on rachillae. Rachis bracts commonly reduced to
only one tubular one (sometimes none) subtending lowermost partial inflorescence,
somewhat flattened-bicarinate, c. 12-15 mm diameter, rather loose but generally

Rodd, Revision of Livistona (Arecaceae)

117

smooth and entire, toughJy papery, finely and evenly striate, pale straw-coloured
outside, red-brown inside, with inconspicuous mats of strongly contorted transparent
hairs, densest toward apex; higher rachis bracts much smaller, linear-lanceolate; bract
apices somewhat elongated, triangular-ovate, not cuspidate, entire. ¥lower-clusters 1-4 mm
apart, 1-3-flowered; cluster axis slender, 0.2-2 nun long, elongating in fruit to up to 3 mm
long. Cluster-bract c. 0.5 mm long, narrowly triangular, acute to acuminate or
sometimes laciniate at apex, whitish-hyaline, very delicate, apical A-A usually
shrivelled away by anthesis. Bracteoles 2-3, resembling cluster-bract or shorter
and rounded.

Flowers broadly funnel shaped to rotate, 1.8-2.3 mm long. Anthopodium slender,
0.2-0.6 mm long. Sepals concave, very loosely cupping petals, whitish-scarious except
at base, 0.7-1.4 mm long, connate for 'A-A their length, narrowly triangular to
somewhat ovate, acute to slightly acuminate. Petals rotately spreading or forming a
funnel, leathery in texture, thin to moderately thick, 1.5-1.9 mm long, broadly
ovate-oblong to triangular-ovate with apiculate, thickened, slightly hooded apex,
auriculate at base with prominent, rounded, downward-pointing auricles (or auricles
occasionally lacking); inner faces with 3-4 parallel raised nerves close to centre.
Stamens %-A as long as petals, connate for up to c. 'A their length; filaments broadly
triangular-ovate, rather abruptly tapering into elongated acuminate apex, obscurely
shouldered low on each margin; anthers 0.3-0.4 mm long. Carpels c. 'A as long as
petals; ovaries very broad, cylindric-globose, their summits very gibbous abaxially,
rather gradually tapering into tapering styles under 0.2 mm long.

Fruit obovoid-pyriform or ellipsoid, usually slightly flattened or even concave
adaxially, 10-12 mm long, 6-7 mm diameter, base tapering or rounded or with abrupt
short conical projection, apex rounded sometimes with stigma persisting as short
prickle. Epicarp dull purplish, slightly pruinose, ripening to glossy black (finely
wrinkled with reticulum of sharp ridges when dry). Mesocarp c. 1 mm thick, soft and
pulpy, purple, with numerous deep red tannin bodies when dry. Endocarp adhering
tightly to both mesocarp and seed. Seed 8-9 mm long, 5-5.5 mm wide. Intrusion small,
penetrating from a lateral position for little more than 'A width of endosperm,
irregularly lobed mainly toward seed base and apex. Embryo lateral, midway between
seed base and apex or slightly above. (Fig. 4i, 20c-d, 21).

Distribution: Northern Territory, escarpments of far north and north-east, extending
south-west to around Katherine and south-east to just across the border into far
northwest Queensland; also islands of the western half of Gulf of Carpentaria.

Conservation status: not considered at risk.

Selected specimens examined: Northern Territory: Darwin & Gulf: Arnhem I lighway, 2 km W of
West Alligator River, Benson 952 ,11 June 1974 (NSW); 21 miles [34 km] NW of Katherine, Lazarides
6627, Sep 1961 (CANB); 91 km from Pine Creek towards UDP Falls, Gittins 2590, July 1973 (NSW,
DNA); Katherine Gorge, 15 miles [24 km] NE of Katherine Lazarides 6990 ,15 Sep 1964 (CANB,
NSW); Katherine Gorge, c. 'A km SE of National Park headquarters, Rodd 2931, 2932, 3 Nov 1974
(NSW, DNA, K, BH); 8 km NW of El Sharana, Benson 994 ,18 Dec 1974 (NSW); Vicinity of El Sharana
Mining Camp, Martensz & Scliodde AE384 ,17 Jan 1973 (CANB); Pine Creek-Oenpelli Road, vicinity
of Mudginberry Station, Syttlon 10347 (NSW); Munmalary Station, (Kay L.S.), Latz 3728, 10 May
1973 (DNA, CANB); Kakadu National Park, 6 km SW of Mt Brockman, Telford 8064 & Wrigley, 23
Apr 1980 (CBG, NSW); Kakadu National Park, 2.5 km NW of Koongarra Saddle, Telford 8114 &
Wrigley, 24 Apr 1980 (CBG, NSW); c. 90 km S of Maningrida, Maconochic 1581 ,25 June 1972 (DNA,
CANB, K); Gulf of Carpentaria, Maria Island, Dunlop) 2836, 14 July 1972 (DNA, DWL, BR1, AD,
CANB, NSW); South West Island, McKey 101, 24 May 1970 (DNA, K, NSW); Centre Island, Sir
Edward Pellew Group, Craven 3841 ,11 Feb 1976 (CANB); Rice 2037 ,10 Sep 1975 (NSW, BRI). Barkly
Tableland: Echo Gorge, Wollogorang Station, Thomson 802, 27 Nov 1984 (DNA, NSW).

118

Telopea 8(1): 1998

Notes

1. Most previous attempts to apply this name, which together with L. humilis forms
part of Brown's protologue of Livistona, seem to have been surrounded by uncertainty
and confusion. Martius (1838), who apparently saw no material, suffered no such
confusion, for he had available (and published magnificent reproductions of) Bauer's
very accurate and detailed drawings which can quite satisfactorily be tied to Brown's
type collection.

2. Mueller (1865, 1874, 1878) virtually ignored Brown's L. humilis and L. inermis,
apparently including them in his vaguely defined L. leichhardtii, together with the
entity now distinguished as L. benthamii.

3. Wendland and Drude (1875) appear to have based their fairly lengthy description of
L. inermis largely on Queensland east-coast specimens, citing '"Rockingham"! (Folia et
semina matura lect. initio Septembris 1865)' and "'Moore's Creek Range near
Rockhampton"! (leg. Thozet)'. These (seen in MEL) are respectively L. drudei and
L. decipiens according to the present treatment. In deference to Brown's concept of the
species Wendland and Drude added 'Regiones circa sinum Carpentaria ex
narrationibus plurum investigatorum (Burke, Wills, Stuart etc.)'.

4. Bentham (1878) confessed to knowledge of this species only from the accounts of
Brown and Martius, and concluded: 'It may prove to be a variety only of L. humilis.'

5. Beccari (1931) gave an accurate, amplified description of the type material, at least
as to flowers and fruits. However, the leaf he described seems more like that of
L. humilis than of L. inermis as described here or as figured by Bauer (in Martius).
Beccari, in discussion, infers from this discrepancy that Bauer's habit and leaf
illustrations represent a different species from the flowers and fruits, possibly his
(Beccari's) L. rigida (type also from Gulf of Carpentaria). Recent collections and field
observations fully support the accuracy of Bauer's illustrations, so Beccari's
hypothesis has turned out to be false.

6. Blake (1954), in notes on northern Australian plants, was the first to apply Brown's
name to any recently found palm in the Northern Territory, though only tentatively.
He noted, under L. 'inerme 'A small-leaved slender palm with smooth petioles
probably belonging to this species was observed in 1946 about SE of the junction of the
McKinlay and Mary Rivers, but as the plants were sterile no specimens were collected.'

7. Specht (1958) in an account of the Arnhem Land flora made no mention of L. inermis,
though the Oenpelli district, where it is known to occur, was one of his main centres
of collecting. He presumably did not distinguish it from L. humilis.

8. As now interpreted, this species exhibits considerable variation, and further
observation and collecting are needed. Basally branching trunks seem to occur only in
some populations, judging from specimen notes and accompanying photographs, but
do not appear to correlate with characters of leaf, inflorescence or flower. Leaf size is
very variable as is degree of dissection: most commonly encountered are leaves
dissected to within 1-3 cm of the hastula, but the other extreme is represented by
Benson 952, with leaves dissected only to within 7-8 cm of the hastula, with very broad
segments; this collection also shows a variant flower type, with shorter, thicker, more
triangular-acute petals than in others examined, though features such as calyx and
inflorescence agree with L. inermis. Alternatively it may be a hybrid with L. humilis (see
note under that species).

9. It is possible that some form of sexual dimorphism is expressed in the floral
variation, though there does not appear to be dimorphism in overall inflorescence
structure.

Rodd, Revision of Livistona (Arecaceae)

119

10. Thomson 802, from dose to the extreme south-east of the known range of the
species, is also somewhat atypical in the large size of its leaves and breadth of
segments; it is possible, but unlikely, that further collections and observations in this
area will reveal it as belonging to a distinct entity.

11. Birch and Dowe (1989) report L. inermis from Queensland, on Westmoreland
Station about 20 km from the Northern Territory border and 65 km inland from the
gulf. They describe its habitat as chiefly bases of cliffs of sandstone conglomerate,
which is consistent with its occurrences in the Northern Territory.

13. L. lorophylla Becc.

Beccari (1921:18; 1931: 87, t.6, f.5 ); Wilson (1992:1249, fig. 353B); as 'loriphylla'.

Type citation: 'Nord Ovest Australia nel Cambridge Gulf, a circa 120°20 long. E. Gr.e
15° L. S. (Johnson )'

Type: Western Australia: Cambridge Gulf, Johnson s.n.; holo FI, photo NSW (2 sheets);
iso 'Johnston 1885' MEL, NSW (fragment only). Tire geographical coordinates cited in
the protologue are not on any specimen; they represent a spot a long way out to sea
north-west of Cambridge Gulf.

Trunk solitary, to 5 m tall, 8-10 cm diameter, broadening slightly at the very base into
a short cone. Surface closely ringed with leaf-scars; vertical fissures rather sparse but
quite long and crooked; basal 2-3 m of trunk bearing crowded, appressed petiole stubs
1—4 cm long, leaves and sheaths shed cleanly higher up. Crown globose, very open,
consisting of 30-40 mostly arching to drooping leaves. Ligules with ragged edges,
outer surface dull straw-brown with mosaic of darker brown corky scales.

Petiole 90-100 cm long, 11-12 mm wide, flattened-triangular in t.s. with rounded keel.
Margins with regularly-spaced linear brown calli, toward base armed with a few
irregularly scattered, slender, patent or retrorsely hooked dark brown prickles up to 3 mm
long. Surfaces red-brown, darker at base, the upper smooth to obscurely striate,
largely glabrous, the lower ± densely scurfy-tomentose, less so toward apex, with
appressed tomentum of flaky white to pale brown scales. Hastuln rather irregular; base
shortly and broadly 3-lobed, ± truncate except for central apiculum; rim at c. 30° to
costa, convex, 4-9 mm broad, necrotic margin broad and irregular.

Lamina 65-100 cm long, 0.30-0.35 mm thick, strongly costapalmate, ± semicircular in
outline, with broad basal sinus and costal region often strikingly pinnate with
conspicuous gaps between segment bases, coriaceous, flexible; slightly contorted with
1 adaxial undulation either side of the barely deflexed costa and lower edges involute
with lowest 4-6 segments either side resupinate. Segments c. 17-24 either side of costa;
largest segments 14—20 mm wide narrowing to 8-9 mm at base, free for 95-98% of their
length, bifurcated for 55-78% of free length, the lobes diverging at 20-30°, apices finely
attenuate, threadlike. Intersegmental appendages threadlike, to c. 2.5 mm long. Ribs:
abaxial ribs in t.s. bluntly square-edged, up to 0.9 mm thick, 0.9 mm deep; adaxial ribs
hardly projecting from lamina. Venation: major longitudinal veins 7-8 either side of
abaxial rib, prominent on both surfaces; transverse veins prominent, mostly
discontinuous and near-orthogonal with respect to longitudinal veins. Surfaces
olive-green, concolorous, somewhat glossy, glabrous except for dense white to
brownish scurfy tomentum on bases of ribs on both surfaces.

Inflorescences shorter than or just equalling petioles, ascending and arching. Partial
inflorescences 4-6, subequal, branching to 3 further orders; rachillae 0.2-5 cm long,
0.3-0.5 mm thick; all axes glabrous, brown to reddish turning white post-anthesis with
spongy amorphous surface layer of cells. Rachis bracts slightly flattened, bicarinate,
c. 14 mm diameter, toughly papery, finely but deeply rugulose-striate, pale
reddish-brown, densely but not conspicuously scurfy-tomentose with matted,
crinkled, transparent hairs, densest near apices; bract apices triangular to rounded.

120

Telopea 8(1): 1998

obtuse to acute, not cuspidate, often densely fringed with hair-like scales. Flozver-
clusters 1-2 mm apart, 1-4-flowered; cluster axis knoblike, 0.2-0.5 mm long.
Cluster-bract c. 0.5 mm long, initially broad and pale brown at base narrowing to a
transparent, serrate to laciniate apical portion, mostly shrivelled by anthesis.
Bracteoles 2-3, lunate to triangular-acuminate, irregularly laciniate, often resembling
cluster-bract.

Flowers 1.2-2.0 mm long, broadly funnel-shaped to cup-shaped. Anthopodium very
short, hardly distinguishable from calyx. Sepals membranous, loosely cupping petals,
c. 0.7 mm long, connate for c. % their length, triangular-ovate, acute to slightly
cuspidate. Petals thin-textured, c. 1.2 mm long, barely connate, broadly ovate with
slightly thickened acute or very shortly cuspidate apex, strongly auriculate at base,
with no obvious cavities on inner faces. Stamens c. % as long as petals, connate for
almost A their length; filaments broadly triangular-ovate, mostly slightly shouldered
low on each side, thence tapering ± smoothly to a finely cuspidate apex; anthers
0.2 mm long. Carpels little more than 'A as long as petals; ovaries obovoid-cylindric,
summit strongly gibbous abaxially, abruptly narrowed into slender straight styles
c. 0.2 mm long.

Fruit (only seen immature) obovoid-pyriform, dorsally gibbous, c. 9 mm long, 6 mm
diameter. Mesocarp c. 0.6 mm thick. Seed not seen mature. Intrusion lateral, elongated
along long axis of fruit, irregularly lobed especially toward fruit base and apex.
Embryo lateral, closer to apex than to stalk. (Fig. 4h, 19, 20a-b, 21)

Distribution: scattered sporadically through the northern Kimberley region of north
Western Australia, from a little north of Derby eastward to the west shore of
Cambridge Gulf, and with an outlier in the Northern Territory; nowhere more than
about 100 km from the coast.

Ecology: usually associated with outcrops of sandstone or siliceous metasediments,
including low 'breakaways' in the broad river valleys, cliff ledges of major gorges, and
rims of plateaus. Associated vegetation is commonly dominated by the prickly grasses
of genera Triodia and Plectraclme, usually with sparse stunted trees including Terminalia
spp.. Ficus spp., Owenia vernicosa, Persoonia falcata, and some of the smaller Eucalyptus
and Corymbia species.

Conservation status: not considered at risk.

Specimens examined: Western Australia:Fitzgerald: West Kimberley (Brockmans Expedition),
House s.n., 1901 (PERTH); Stewart River, c. 68 km NNE of Derby, 14 km from Kimbolton HS.
towards Gibb River road, Telford 6404 b Butler, 2 Aug 1977 (CBG, NSW, PERTH). Gardner: Uwins
Is, Brunswick Bay, Wilson 11450 ,8 July 1973 (PERTH); W5 Marigui Promontory, Prince Regent River
Reserve, Kenneally 2162, 27 Aug 1974 (PERTH, NSW); North Kimberley, Speck 4916, Sep 1954
(PERTH); Lower King Edward River, Gardner 1544, 22 Aug 1921 (PERTH); Sir Graham Moore Is,
Wilson 11222, 30 June 1973 (PERTH); c. 3 km ESE of Kalumburu Mission, Rodd 2871, 2872, 25 Oct
1974 (NSW, PERTH, K, BH); 3 miles [5 km] NNW of Kalumburu Mission, Speck 4916, 4 Sep 1954
(CANB); Kalumburu Mission, Brigden DNA688S, Aug 1973 (CANB); Kalumburu, Crawford 58 ,6 Jan
1974 (PERTH); Douglas b Mees s.n., 3 July 1960 (PERTH); Lullfitz s.n., 18 Jan 1975 (PERTH);
Drysdale River National Park, George 13758 ,12 Aug 1975 (PERTH, NSW); Site B3, Cracticus Falls,
Kenneally 4153, 9 Aug 1975 (PERTH, NSW); Site C5, Fern Gully, Drysdale River National Park,
Kenneally 4557, 20 Aug 1975 (PERTH); Site C3, Planigale Creek, Drysdale River National Park,
Kenneally 4447 ,19 Aug 1975 (PERTH).

Northern Territory: Victoria River: Victoria River area, 15°16’S, 129°35'E, Dunlop 8203 b Leach, 9 Mar
1989 (NSW).

Notes

1. The epithet was published by Beccari as ‘loriphylla' but under article 60.8 of the ICBN
is to be corrected. Loron, a thong or strap, was adopted into Greek from Latin lorum.
Liddell & Scott (1864) list three combinations in the form loro- with other Greek words

Rodd, Revision of Livistona (Arecaceae)

121

(see also such botanical names as Loropetalum). The form /on- would, of course, be
correct in combination with a Latin word.

2. This species characteristically occurs on sandstone cliff ledges and around
'breakaways' over a large area of the northern Kimberleys. On heavier basaltic and
bauxitic soils in the central part of its range it is replaced by the locally very abundant
L. eastonii, a quite unrelated species; in the increasingly arid range country to the south
it is replaced by L. kimberleyana, which appears to be closely related.

3. The Northern Territory palm sometimes determined as 'L. lorophylla' is here treated
as L. inerviis R.Br. It is possible that a broad concept of the latter species might take in
the north Kimberley plants, in which case Beccari's name would be reduced to
synonymy, but differences in growth-habit, stature, colouring and inflorescence
structure seem sufficiently consistent to warrant their separation.

4. George 13758 and Kenneally 4153 (from Drysdale River National Park) both exhibit a
remarkable androecium: the antesepalous stamens are nearly all replaced by
staminodes varying from quite petal-like to much narrower, incurved at tip and of
irregular shape, but usually 2 of the larger staminodes have adnate to their inner face
(as do the normal petals) an additional complete stamen of normal appearance. This
bizarre flower structure was not found in other collections from areas nearer the coast.
However, the sample available is very small; it remains to be seen whether these
represent a localised aberrant population, or alternatively represent the normal
condition for one of the sexual states in the species as a whole. However, dissection of
flowers of the isotype fragment in NSW, and buds of two other pre-anthesis collections
revealed a norma! arrangement of stamens. In other respects this species appears to
show relatively little variation, considering its apparently rather fragmented distribution.

14. L. kimberleyana Rodd, sp. nov.

Palma solitaria magnitudine mediocris, trunco ad 15 m alto, c. 15 cm diametro; coma
globosa foliis cinereo-viridibus, debilibus, versus apices cemuis; petioli 150-180 cm
longi, 14-17 mm lati proxime laminam. Laminae costapalmatae, c. 90 cm longae,
dissectae fere ad hastulam; segmentis c. 25 mm latis, profunde bifurcatis apicibus
subtiliter attenuatis. Inflorescentiae quam petiolis breviores vel aequilongae, c. 7 ramis
lateralibus, illis denuo 3-plo ramosis rhachillis 0.4-6 cm longis, 0.8 mm diametris.
Flores l-2(-3?)-fasciculatae. Fructus obovoidei, parvi.

Type: Western Australia: SW base of Mt King, Durack Range, 17°20’S, 127°23 E, A.N.
Rodd 2866, 24 Oct 1974; holo NSW; iso PERTH, CANB, K.

[Livistona sp. A, Wilson (1992:1250); Livistona sp. 'Kimberlies' (Jones 1984)]

Trunk solitary, to c. 15 m tall, 13-15 cm diameter at 1 m above ground, generally
broadening gradually at base into a narrow cone up to 25 cm diameter. Surface closely
ringed with prominent sheath-scars; vertical fissures rather sparse, short, fairly
straight; basal 1-1.5 m bearing closely appressed petiole-stubs to c. 5 cm long, leaves
and sheaths shed cleanly higher up.

Crown globose, open, consisting of about 40 strongly arching to pendulous leaves,
usually with a skirt of dead leaves hanging beneath. Ligules not observed.

Petiole c. 150-180 cm long, 14-17 mm wide; flattened-triangular in t.s. with rounded
keel. Margins on upper half of petiole with closely spaced minute dark calli, toward
base with retrorse or less commonly antrorse prickles, often dark-tipped, hooked and
pungent, up to 5 mm long. Surfaces pale dull pinkish, strongly pruinose, obscurely to
prominently striate, the lower with irregularly scattered bands of elongated shallow
pits containing minute blackish dot-like or linear basal-masses obscured by appressed
white scurfy scales. Hastula short, very irregular; rim appressed against costa, 5-9 mm

122

Telopea 8 ( 1 ): 1998

wide, roughly semicircular, triangular or ovate, strongly cuspidate, asymmetric, with
irregular patches of necrotic margin.

Lamina 70-100 cm long, 0.4 mm thick, strongly costapalmate, almost circular in
outline with broad basal sinus, coriaceous, flexible; moderately contorted with 1
adaxial undulation either side of the slightly deflexed costa and involute at base with
few segments on either side resupinate. Segments 20-22 either side of costa; largest
segments 23-25(-33) mm wide, free for 85-90% of their length, bifurcated for 59-76%
of free length, the lobes flexuose, lying almost parallel, their apices finely attenuate but
not thread-like except on lower segments. Intersegmental appendages inconspicuous,
threadlike, c. 3 cm long, soon breaking off. Ribs: abaxial ribs in t.s. oblong, bluntly
square-edged, to 1.0 mm thick, 1.5 mm deep; adaxial ribs much shallower. Venation:
major longitudinal veins 7-8 either side of abaxial rib, prominent on both surfaces;
transverse veins clearly visible on both surfaces, mostly discontinuous, most
near-orthogonal with respect to longitudinal veins. Surfaces slightly discolorous,
greyish-green, both surfaces quite glabrous, moderately pruinose but wax layer not
easily damaged.

Inflorescences (seen only in immature fruit stage) as long as or slightly shorter than
petioles, stiff and straight. Partial inflorescences c. 8, these branching to 3 further
orders; rachillae 0.4-6 cm long, c. 0.8 mm thick; all axes pallid straw-coloured with a
hard, waxy surface layer. Rachis bracts c. 2 cm diameter, slightly distorted,
straw-coloured tinged pinkish, finely striate, glabrous except for patches of appressed,
transparent, colourless contorted scales toward apex; bract apices ovate, obtuse to
shortly cuspidate. Flower-clusters 1-3 mm apart, l-2(-3?)-flowered; cluster axis 0.3-0.8 mm
long. Cluster-bract c. 0.5 mm long, broadly triangular-cuspidate. Bracteoles 2-3, to 0.5 mm
long, resembling cluster-bract.

Flowers: Anthopodium c. 0.3 mm long, c. 1.0 mm diameter, bulging downward at
base, deeply recessed at junction with pedicel. Sepals fleshy-textured with rather
narrow hyaline rim, c. 1.8 mm long, connate for c. 'A length, broadly triangular, bluntly
acute. Petals thick-textured, deep reddish-brown, to 1.5-3.0 mm long, triangular,
hooded at the apex with somewhat indexed small callous point, cavities on inner faces
not very distinct. Stamens c. % or more as long as petals, connate for up to half their
length; filaments very broadly triangular, narrowing rather abruptly into shortly
acuminate apices; anthers c. 0.45 mm long and almost as wide. Carpels (seen as
squashed remains) equalling stamens, abruptly narrowing into slender oblique styles
c. 0.3 mm long.

Fruit (only seen very immature) obovoid, c. 8 mm long, 6 mm diameter. (Fig. 4g, 17d)

Distribution: Western Australia — ranges of central Kimberley Region, mainly King
Leopold and Durack Ranges, through both of which it appears to be widely distributed.

Ecology: occurs usually in rocky ravines and along cliff bases and on cliff ledges,
mostly associated with siliceous sediments and metasediments. Associated trees
include Corymbin aspera and Gardenia spp., while the ground layer is dominated by
Triodin spp. and Plectrachne spp.

Conservation status: not considered at risk by Briggs and Leigh (1996), if not being
listed can be interpreted thus. However, there is hardly evidence of the existence of
large populations of this species, and there appear to be no national parks coinciding
with its range (maybe Tunnel Creek N.P.?), so a listing of 3R may be more appropriate;
further investigation is needed.

Specimens examined: Western Australia: Fitzgerald: c. 26 km NE of Inglis Gap, Symon 10154,
23 May 1975 (NSW); Mt Ord Gorge, King Leopold Ranges, 20 miles [32 km] SSW of Mt House Stn,
Lazarides 6453, 29 July 1959 (CANB); Teroni Gorge, 12 miles [19 km] NNW of Elgie Cliffs Station,
East Kimberleys, Lazarides 6395, 22 July 1959 (CANB); SW base of Mt King, Durack Range, Rodd
2867 ,24 Oct 1974 (NSW, PERTH, CANB, K); Chamberlain River Gorge, 7 km E of El Questro Station

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123

homestead, Kenneally 10978, 30 June 1989 (PERTH, NSW); Mt Leake, Fitzgerald 1206, July 1905
(PERTH); Mt Broome, Fitzgerald 815, May 1905 (PERTH)

Notes

1. This was formerly confused with L. alfredii (e.g. Fitzgerald 1918; Gardner 1923 and
see note under that species).

2. Fitzgerald (1918) reported 'L. alfredii’ as occurring on 'Mts Herbert, Broome, Leake
and Barnett; Isdell, Harris and Phillips Ranges; junction of Hann and Barnett Rivers
(W. V.F.)' — some of which fill in the crescent formed by the King Leopold and Durack
Ranges.

3. Gardner (1923) gives for ‘L. alfredii ' the distribution 'Synnot Creek near Charnley
River, King Leopold Ranges near Prince Regent River, and sandstone ranges near the
lower King Edward River and Napier Broom Bay. Among sandstone rocks in elevated
situations. King Leopold Ranges and Sir Frederick Hills.' Some of these localities
would extend its distribution a long way north, but it is more than likely that they
represent occurrences of L. lorophylla, which Gardner apparently did not distinguish
from his 'L. alfredii'. Neither Fitzgerald nor Gardner appear to have collected
specimens to support their observations, which is perhaps understandable
considering the conditions under which they travelled.

4. Its closest relationship appears to be with L. lorophylla Becc., from which it differs
principally in its overall larger dimensions, its pruinose foliage and its much thicker,
more rigid inflorescence rachillae. It is possible that toward the northern part of its
range it may show some intergradation with that species, in that one or two specimens
seen have not been unequivocally referable to one or the other. It is possible even that
further study may show it to represent an arid race of L. lorophylla, but for the
meantime it seems preferable to treat it as a distinct species.

5. The type locality in the Durack Range is also the type locality for L. mariae subsp.
occidentalis, and indeed these two very different taxa grow together here in a rocky
ravine, but showing different ecological preferences, with a grove of L. mariae confined
to permanent soaks in the bed of the ravine, and scattered L. kimberleyana among rocky
outcrops on the lower slopes — the populations nonetheless intermingling at the
boundaries. At the time of my visit I observed (and photographed) some individuals
that did not seem clearly referable to one or another of these taxa and suspected that
they might be hybrids, but time did not permit collection of a specimen.

15. L. victoriae Rodd, sp. nov.

Palma solitaria magnitudine mediocris, trunco ad 12 (-20) m alto, c. 20 cm diametro.
Coma foliis pruinosis rigide radiantis; petioli c. 1 m longi, 25 mm lati proxime
laminam. Lamina valde costapalmata, c. 1 m longa. Inflorescentia 4-8 ramis
lateralibus, illis denuo 4-plo ramosis, rhachillis 0.5-3.0 cm longis, gracillimis. Fructus
subglobosi-ellipsoidei, in siccitate ad 11 mm longi, 10 mm diametro, sutura ventrali
manifesta. Semina intrusione testa laterali, parvo, embryone fere basali.

Type: Northern Territory: c. 3 km W (by road) of Victoria River crossing, Katherine-
Kununurra road, 15°35’S, 131°06’E, alt. c. 80 m, A.N. Rodd 2934, 3 Nov 1974; holo NSW;
iso DNA, K, PERTH.

[Livistona sp. 'Victoria River' (Jones 1984); ‘Livistona sp. B' (Wilson 1992:1250)]

Trunk solitary, to 12 (-20?) m high, c. 20 cm diameter at 1 m above ground, hardly
reducing upward, broadening close to ground into a somewhat bulbous swelling to
c. 30 cm diameter. Surface rather obscurely ringed with sheath scars, becoming smooth
on lower trunk with age; vertical fissures not very conspicuous; petiole-stubs c. 4 cm

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Telopea 8 ( 1 ): 1998

long present on lowest 2-3 m of trunk, closely appressed with irregular, abraded or
somewhat ragged ends.

Crown rather open, with pale blue-grey cast, globose or slightly elongated, consisting
of 30-40 leaves with gently arching to drooping petioles, the leaf-blades continuing in
the same plane with straight segment-lobes but conspicuously decurving costa.
Ligules prominent, whitish or pale brown, contorted, subglabrous, strongly impressed
between fibres, tearing early from petioles but leaving large, roughly triangular
remnants attached to them.

Petiole 100-110 cm long, 22-28 mm wide, strongly flattened-triangular in t.s. with
rounded keel, slightly concave above but often with slight median keel. Margins
obscurely denticulate or toward base armed with irregularly scattered blunt, antrorse
or patent prickles up to c. 2 mm long. Surfaces pale dull pinkish, strongly pruinose,
obscurely and finely striate; lower surface, and to a lesser extent upper, bearing
unevenly but densely scattered groups of slightly elongated shallow pits containing
brown linear rows of minute basal-masses, appressed white fimbriate scales spreading
from margins of pits, glabrescent with age. Hastula flat; base narrowly V-shaped to
somewhat ovate-apiculate, often very asymmetric; rim at c. 20° to costa, 3-7 mm wide,
with mostly narrow necrotic margin.

Lamina 90-110 cm long, 0.35-0.40 mm thick, strongly costapalmate, with broad basal
sinus, fairly stiff and harsh, tough and not readily split; strongly contorted with 1
adaxial undulation either side of the strongly deflexed costa, at base on either side
shortly involute with lowermost 2-3 segments resupinate. Segments 20-28 either side
of costa; largest segments 29^10 mm wide, free for 55-65% of their length, tapering
slightly toward point of bifurcation, bifurcated for 55-70% of free length, the lobes
evenly tapering but attenuated at apices into slender, necrotic threads, breaking off
with age. Intersegmental appendages present, moderately persistent, straight, white,
mostly c. 5 cm long. Ribs: abaxial ribs in t.s. oblong, somewhat bluntly square-edged,
c. 2.5 mm deep, c. 1.0 mm wide; adaxial ribs very similar. Venation: major longitudinal
veins 9-11 either side of abaxial rib, moderately prominent above, less prominent
below; transverse veins moderately prominent on both surfaces, often steeply angled,
often continuous across most of the width of the half-segment. Surfaces concolorous,
strongly pruinose, glabrous except for ribs bearing scales and brown markings as on
petiole.

Inflorescences variable in length, from slightly shorter than petioles to about petiole
plus half lamina length. Partial inflorescences 5-9, subequal, the largest slightly over
half total inflorescence length, each branched to 4 further orders; rachillae 0.4-3 cm
long, 0.4-0.6 mm thick, cream when fresh, glabrous; larger axes striate, often very
sinuous; all axes minutely papillose (appearing pallid with age). Rachis bracts usually
loose and distorted, becoming split and raggedly fibrous with age, thick and
harsh-textured, straw-coloured aging silver-grey though more reddish-brown toward
apices, finely striate, unevenly but often densely woolly with closely appressed,
matted, somewhat curly, narrow white scales; bract apices elongated, acute or obtuse,
never cuspidate. Flower-clusters 0.5-3 mm apart, 1-2-flowered; cluster axis 0.2-0.7 mm
long, cylindrical, moderately slender; longer cluster axes often with lateral flower
closer to base than apex. Cluster-bract very short, rounded or minutely apiculate.
Bracteoles not detectable.

Flowers: Anthopodium c. 0.5 mm long, bulging downward at base. Sepals
membranous, closely appressed to petals, connate for over M their length with very
broad shallow sinuses, triangular, acute. Petals c. 1.2 mm long, rather narrowly
triangular-ovate, acute and cymbiform at apex; inner face with 3 fairly deep cavities.
Stamens at least % as long as petals, connate for c. 'A their length; filaments broadly
triangular-ovate, tapering smoothly to an elongated slender apex. Carpels not seen.
Fruit (only immature and long-fallen fruits seen) spherical-ellipsoid, strongly flattened
adaxially, c. 11 mm long, 10 mm diameter, abruptly narrowed at base into a very short

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125

conical extension, ventral suture evident. Epicarp reddish-brown (immature, dry) or
grey-brown (fallen), minutely verruculose. Mesocarp apparently very thin, deep
red-brown on inner surface. Endocarp c. 0.15 mm thick. Seed c. 9 mm long, flattened
on adaxial face. Intrusion penetrating from lateral position for / to 'A seed diameter, in
l.s. with very narrow base and small lobes directed away from embryo. Embryo
sub-basal. (Fig. lb, 4f, 17c)

Distribution: confined to the region of the border between the Northern Territory and
Western Australia, north of about 18° latitude. It is abundant in the vicinity of the
middle and lower parts of the Victoria River (downstream of Victoria River Downs
Station), and much of the Ord River basin, though apparently not downstream of
Kununurra. Two sites where it occurs in great abundance are the Bungle Bungles,
south of Lake Argyle, and Jasper Gorge, a western tributary of the Victoria River a
short distance north of Victoria River Downs. To the northeast, populations of what is
very likely this palm were seen from the air on ranges forming the watershed between
the Fitzmaurice and Katherine Rivers. For references to possible southern extensions
of its range see Note 4 below.

Ecology: apparently virtually confined to sandstone range country, where it grows in
rocky ravines, on slopes below cliffs, on cliff ledges, and less commonly along gravelly
or sandy watercourses, all habitats where there is permanent seepage water available
to its roots. Trees with which it is commonly associated include Corynibia aspera, Ficus
leucotricha, Termimlia canescens, Owenia vernicosa. Gardenia spp. and Acacia spp. The
dominant grass is nearly always Triodia.

Conservation status: although this species has received the coding 3KC- (Briggs &
Leigh 1996), it is now known to occur in very large populations, with many local
concentrations scattered over a considerable area of the Ord and Victoria River
catchments, mostly on rocky, inaccessible sites and it is very adequately conserved in
the Bungle Bungle-Purnululu National Park (WA) and the Gregory National Park
(NT). For these reasons is should be regarded as 'not at risk and should no longer
have a ROTAP listing.

Specimens examined: Northern Territory: Victoria River: Yambarran Range, Leach 4554 & Walsh,

16 May 1994 (DNA, NSW,BRI, MEL); N side of Jasper Gorge, upper slope below cliffs of sandstone,
McGillivray 3804 & George, 15 June 1978; Jasper Gorge, Parker 1065, 13 July 1977 (DNA, NSW);
Byrnes 1718, 17 Sep 1969 (DNA, CANB, PERTH); 5 km E of Victoria River crossing, Maconochie 2493,
8 Oct 1980 (DNA, NSW).

Western Australia: Gardner: Ord River Basin, Koford & Dortch 120, Sep 1974 (PERIH), Tributary of
Keep River, Lullfitz s.n., Nov 1978 (DNA, NSW); Thompson Spring, c. 40 km SE of Kununurra, Ryan
s .17., 6 June 1979 (PERTH, NSW); 7 miles [11.3 km] E of Denham [=Dunham?] River Station, Perry
2529, 19 J uly 1949 (CANB, NSW); Cabbage Tree Creek, Carr Boyd Ranges, E Kimberleys, Rodd 2800,

17 Oct 1974 (NSW, K, PERTH); Carr Boyd Range, Wyndham to Halls Creek road, Hand s.n., 1978
(NSW 108860b); near Lucky Hill, 23 km NNE of Dunham River HS., NE Kimberleys, Lazarides 8552,
13 Mar 1978 (CANB, K, PERTH); Piccaninnv Creek Gorge, 15 km SE of Bungle Bungle Outcamp,
Bungle Bungle Range, NE Kimberley, Blackivcll BB419, 6 Apr 1985 (PERTH); Bungle Bungle Range,
gorge 3 km SE of Bungle Bungle Outcamp, Kenneally 9260, 8 July 1984 (PERTH), Gallery Forest
16 km NE of Bungle Bungle Outcamp, E Kimberley, Kenneally 9212, 5 July 1984 (PERTH), Frog Hole
Canyon, N end of Bungle Bungle Range, Briggs 9307 1 Aug 1994 (NSW); Gorge off Piccaninny
Creek, Bungle Bungle Range, Briggs 9309,5 Aug 1994 (NSW); Teroni Gorge, 12 miles [19 km] NNW
of Elgie Cliffs Station, E Kimberleys, Lazarides 6395, 22 July 1959 (CANB); Bens Springs, 3 km E of
El Questro homestead, Kenneally 10965, 29 June 1989 (PERTH, NSW).

Notes

1. A well-marked species showing clear morphological and geographical demarcation
from others in the region. In leaf characters it approaches most closely to L. alfredii,
differing slightly but consistently in depth of lobing and bifurcation of segments;

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Telopea 8(1): 1998

the trunk is also more slender. The large inflorescences, ramified to 5 orders and with
most rachillae extremely short (4-10 mm), together with the small fruits, and seeds
with small endosperm intrusion, combine to make this a very distinctive species.

2. The epithet victoriae alludes to its occurrence in the vicinity of the Victoria River, but
the genitive form is used to complete a trio with L. alfredii and L. mariae, named
respectively for Queen Victoria's son and daughter-in-law and occurring more or less
equidistant from the present species in similarly semi-arid habitats.

3. It is puzzling that this has remained unrecognised as a species for so long, or indeed
not even represented by any herbarium collection until Perry's in 1949. It must have
been a familiar plant to pioneer pastoralists of the eastern Kimberley Region from the
1880s onward. Further, Mueller himself must have seen it in the course of his journey
up the Victoria River with Gregory in 1854.

4. In his autobiographical book. Boss Drover, Keith Willey (1971: 113) refers to the
former presence of native palms 'at Lewis Creek and in isolated places in the desert',
and to 'groves of them at Flora Valley and Sturt Creek but the pioneers cut them all
down to build sheds and yards.' These three localities are all in Western Australia not
far from the Northern Territory border between 18°10’ and 19°10’S, much further south
than any of the collections cited here. He also mentions a solitary, tall palm at 'Palm
Spring' on Montejinnie Station, which is ESE of Victoria River Downs. From his
descriptions of the trunks these palms could well be L. victoriae, though L. marine is
another possibility.

5. Wilson (1992) describes the fruit as 'black, globular, ca 15 mm across'. Fresh mature
fruits may be somewhat larger than dry ones, but I have not seen fruits of such a large
diameter nor any that are quite black in colour in the dry state. Briggs 9307 from the
Bungle Bungle Range has apparently mature fruits — though described by the
collector as 'green' (August), when dry they had turned chestnut-brown with closely
wrinkled skin; they conform to the previously recorded dimensions of 11 mm length
and 10 mm diameter. B. Briggs (pers. comm.) considered that fruit colour was a
consistent feature distinguishing the green-fruited Bungle Bungle palms from the
black fruited plants of L. eastonii on the Mitchell Plateau. Maconochie 2493 bears the
label note 'Fruit globular, green, drying black, thin pericarp.' But the dried fruits of his
specimen, apparently virtually mature judging by the hardness of the endosperm, are
a somewhat reddish-tan with closely wrinkled skin.

16. L. alfredii F. Muell.

Mueller (1892: 112; 1893: 28); Drude (1893: 12); Dammer (1905: 297); Fitzgerald (1918:
24 p.p.); Beccari (1921: 18; 1931: 82, t.6).

Type: Western Australia: Millstream, Hamersley Range, J. Forrest, 1883(7); holo MEL.

[L. marine F. Muell. (1878), as to specimens from Fortescue River, W.A.]

Trunk solitary, to 10 or possibly 12 m tall, 30-35 cm diameter at 1 m above ground,
broadening more or less abruptly at base, often with flaring rim of corky plates above
conical root mass up to 30 cm high. Surface closely ringed with sinuous scars,
somewhat bulging between these to give a transversely corrugated appearance;
vertical fissures close-set, deep, somewhat zig-zagging; basal 1-2 m of trunk bearing
persistent, irregularly broken-off petiole stubs 3—5 cm long, or sometimes whole
dead leaves.

Crown globose, moderately dense, consisting of c. 25-30 leaves, the petioles mostly
gently recurved, usually with a few dead leaves hanging vertically below. Ligules
fairly prominent, with smooth, almost unbroken surface, very pale buff-coloured,
whitish scurfy-tomentose.

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127

Petiole 90-130 cm long, 24-33 mm wide; flattened-triangular in t.s. with rounded keel.
Margins almost completely smooth or armed with irregularly scattered narrow,
patent, bluntish pricldes under 2 mm long. Surfaces pale pinkish-green ageing to
pinkish-brown, smooth, pruinose with hard waxy feel, on younger leaves densely
clothed in minute white waxy appressed scales with patchily distributed tufts of
spirally coiled white laciniae up to 3 mm long, arising from dark brown basal-masses
in regularly distributed linear pits about 2 mm long, more numerous on abaxial
surface, the waxy scales persisting but losing their laciniae. Hastula short and broad;
base obliquely V-shaped or somewhat cuspidate; rim at about 45 ° to costa, 5-10 mm
wide, ± flat, continuous or split at apex, early-necrotic.

Lamina 90-140 cm long, 0.40-0.45 mm thick, strongly costapalmate, truncate at base
(lowermost segments spreading at right-angles to petiole), rather harsh and rigid;
fairly strongly contorted with 2 adaxial undulations either side of the gently deflexed
costa. Segments about 28-33 either side of costa; largest segments 25-42 mm wide, free
for 60-70% of their length, bifurcated for 60-75% of free length, the lobes slightly
diverging, tapering evenly and gradually, attenuated into very fine threadlike apices.
Intersegmental appendages threadlike, up to c. 5 cm long. Ribs: abaxial ribs in t.s.
trapeziform (narrowing slightly to lamina) sharply square-edged, up to 1.0 mm thick,
2.0 mm deep; adaxial ribs similar or slightly shallower. Venation: major longitudinal
veins 8-10 either side of abaxial rib, not very prominent above, even less so beneath;
transverse veins barely visible. Surfaces concolorous, pallid grey-green with waxy
bloom (bluish in field), minute disc-like wax flakes persisting in the fine striations on
older leaves, virtually glabrous except for scattered scales near bases of abaxial ribs.
Inflorescences apparently dimorphic, on fruit-bearing trees ± equalling leaves, on
'male' trees hardly exceeding petioles. Partial inflorescences 5-7(?), subequal; each
branched to 2 ('female') or 3 ('male') further orders; rachillae to 13 cm long, 0.7-1.2 mm
thick, uniformly deep dull pink or cream suffused pink, with whitish pruinosity.
Rachis bracts smoothly cylindrical but mostly irregularly tom a little below apex
revealing net of brown fibres, 2.5-3 cm diameter, hard, leathery, reddish-brown often
becoming whitish-necrotic, those on 'female' inflorescences more tightly sheathing
and on 'male' somewhat loose and wrinkled, bearing a sparse to moderately dense
indumentum of short pale brown scales in a fine pattern of tufts, often scurfy and
whitish on upper half; bract apices long, narrowly triangular or slightly cuspidate,
glabrescent. Flower-clusters 1-5 mm apart, 1-2-flowered (nearly always 1-flowered on
'female' inflorescences); cluster axis (in 'female') present only as a pad with fleshy
lobes protruding between bracts or (in 'male') about 0.5 mm long, knob-like.
Cluster-bract minute, triangular, apiculate. Bracteoles 1-3 detectable as narrow,
apiculate rims.

Flowers 2-3 mm long, ± cylindrical to funnel-shaped. Anthopodium 0.3-1.0 mm long
(shorter on 'male' flowers), mostly thicker than long, bulging downward at base over
cluster axis. Sepals closely appressed to petals, with broad hyaline margins, 0.8-1.0 mm
long, triangular-ovate to broadly triangular. Petals 2.0-3.0 mm long, triangular-ovate,
slightly cymbiform but only at extreme apex, minutely auriculate at base, inner faces
with distinct but shallow cavities. Stamens c. % as long as petals; filaments broadly
triangular-ovate narrowing smoothly into acuminate apex, not shouldered; anthers
c. 0.4 mm long. Carpels c. M as long as stamens; ovaries closely pressed together along
inner faces, plump and gibbous on abaxial side, abruptly contracted into very short,
inward-pointing styles.

Fruit spherical, 25-35(-c. 40) mm diameter; base with shallow conical extension; apex
with stigmatic remains evident as smaller conical projection, adaxially displaced.
Epicarp red-brown to dark brown, smooth when fresh, finely wrinkled when dry, with
pale brown lenticels in the form of circular blisters 0.3-0.5 mm diameter with central
pit, evenly dispersed at 1-2 nun intervals. Mesocarp said to be thick and juicy when
fresh, when dry c. 1.5 mm thick. Endocarp c. 0.2 mm thick, very brittle, pale buff-coloured.

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Telopea 8(1): 1998

Seed near-spherical, 17-20 mm diameter (dry), testa 0.4-0.6 mm thick, its surface pale
buff-coloured and verruculose, with a deeper red-brown hilum. Intrusion penetrating
from a near-basal position, 6-8 mm diameter at base, expanding to only slightly wider
in centre to fill much of centre, leaving shell of endosperm 3-5 mm thick, irregularly but
shallowly lobed. Embryo in near-apical position. (Fig. 4e, 15b, 18)

Distribution: Western Australia, Hamersley Region, Fortescue River in vicinity of
Millstream, also head of Ashburton River and its tributary Duck Creek; recently
discovered is a relict stand of a few trees in the Cape Range, on the west side of
Exmouth Gulf.

Ecology: appears to be restricted to alluvium along rivers, smaller streams and beds of
ravines where moisture is permanently available at least in the form of steam-bed
soaks. In southern tributaries of the Millstream portion of the Fortescue River it forms
dense populations in shallow ravines in soft concreted sediments, their beds with
permanent small pools of highly mineralised and somewhat saline water edged with
white incrustations. Associated trees include Eucalyptus camaldulensis and Melaleuca
leucadendra.

Conservation status: 3RC- (Briggs & Leigh 1996). A substantial proportion of the
populations of the species lies within Millstream Chichester National Park.

Specimens examined: Western Australia: Fortescue: On the 'Mill-Stream' of the Fortescue River,
Forrest 301, June 1878 (MEL); Hamersley Range, Forrest 1879 (MEL) [label worded thus in Mueller's
hand, but also present is a piece of original wrapping paper marked: 'Botanical specimen, flower of
the Marie Livistona Palm, Survey Office, Perth 19 Feb 79/ This collection consists of pieces of
inflorescence and a packet of flowers]; Millstream, Hamersley Range, Forrest (MEL) [a half-leaf and
a packet of dissected seeds, with various slips of paper including notes on nomenclature, possibly
in Ewart's hand]; Millstream, Hamersley Range, anon (Beresford?), 1892 [1 fruit in packet] (MEL);
Millstream, Fortescue River, George 3523, 3 Mar 1962 (PERTFI); Beaugleholes.n., Aug 1974 (PERTH);
Brooker 2064, 24 Sep 1969 (PERTH); 'Millstream' Station, Rodd 2847, 20 Oct 1974, [in homestead
garden, presumed part of natural vegetation] (NSW, PERTH, K, BFI); loc. cit., Rodd 2848, 2849, 20
Oct 1974 (NSW, PERTH); Longreach, Millstream, Fortescue River, Gardner s.n., 21 Aug 1932
(PERTH); Duck Creek c. 32 km W of 'Mount Brockman' Station, Wark spec. A, spec. B, 3 May 1975,
(NSW). Carnarvon: Cape Range, North West Cape, Waldocks.n., 29 Sep 1988 (PERTH, photo NSW).

Notes

1. This is the most isolated and one of the most geographically restricted of all the
Australian species. Its major occurrence at Millstream is spread along about 15 km of
the Fortescue River and the lowest few kilometres of its tributary creeks on the
southern side, coinciding with the surfacing of a strongly discharging artesian aquifer
yielding water of appreciable salinity and alkalinity.

2. Tire other major occurrences known are on Duck Creek, a tributary of the upper
Ashburton River, where Mr R.J. Wark reports there are possibly up to 1000 plants, and
at Silver Grass Pool on the upper Robe River, reported to Mr Wark by the owners of
Mt Brockman Station.

3. In 1989 Mr Kevin Kertneally of the Western Australian Herbarium reported to me
the discovery by a group of zoologists from the Western Australian Museum in
September 1988 of a few palms clinging precariously to life in a remote gully of the
Cape Range, which runs south from North West Cape; a specimen forwarded to me
has been identified as L. alfredii, and this site represents an extension of its range by
approximately 300 km to the west. From the coordinates supplied the locality appears
to be southwest of Learmonth and not within the Cape Range National Park.

4. Mueller's first (and only) use of the name 'L. Alfredi' in 1892 in the Victorian Naturalist
is taken as valid publication of this species, and is cited as the place of publication in
Index Kewensis and by Beccari (1921, 1931), Moore (1963a) and Chapman (1991).

Rodd, Revision of Livistona (Arecaceae)

129

It is, however, in the nature of an informal report rather than a botanical description.
The only parts in it that provide any sort of diagnosis are contained in the words: 'This
particular characteristic [i.e. juvenile leaves being 'a rich bronzy colour'] seems neither
to apply to the West-Australian species and ' ... I have always found transmitted
fruitlets considerably larger than those of the genuine L. Marine , and further some
minor differences exist also in the flowers of the two species, as recently ascertained.'
He continues: 'The West-Australian Fan-Palm has now been named
L. Alfrecii, in honour of H.R.H. the Duke of Edinburgh, at whose nuptial festival the
Central Australian Palm became dedicated to the Princess Marie of Russia.' It can be
argued, however, that the whole article contains an implied reference to the Millstream
element in Mueller's 1878 description of L. marine (based on a leaf specimen from the
'Glen of Palms' and inflorescence and fruit from 'The Millstream', and hence
constitutes valid publication. As noted under L. marine, the 1892 note certainly
signified Mueller's intention to restrict the application of his name to the Central
Australian species, and thus might at least be construed as a sort of lectotypification
of that species.

5. Index Kewensis cites, in addition to the Victorian Naturalist note, an additional
publication of the name by Mueller in Botanisches Zentralblatt in 1893. This is merely a
verbatim transcript (still in English) of the Victorian Naturalist note.

6. If the argument is not accepted that Mueller's 1892 publication was valid, then the
next significant use of the name was by Drude in 1893, in an article entitled 'Uber die
australischen Livistona-Arten', in Beiblatt zu den Botanischcn Jahrbiichern, Nr. 39. In
German with a Latin key, this separates L. alfredii into the monotypic Sect. Gregorya,
defined entirely on characters of fruit and seed (particularly the raphe), and the key
supplies a reasonable diagnosis based on these characters.

7. Beccari's posthumous 1921 (brief Latin) and 1931 (lengthy English) accounts of the
old-world coryphoid palms also provide adequate diagnoses of this species. But even
his description is based on only two collections, one by Forrest from Millstream (date
not cited), the other from 'Nickol Bay' by McRae , dated 1879. There has been no more
recent confirmation of its occurrence at the latter locality (near the towns of Roeboume
and Karratha), and it can probably be treated as a shorthand designation for a large
hinterland, including the Millstream area.

8. Fitzgerald (1918) reported L. alfredii from 'Millstream, Fortescue River (Alex.
Forrest), Mts Herbert, Broome, Leake and Barnett; Isdell, Harris and Phillips Ranges;
hills near the junction of Harm and Barnett Rivers (W.V.F.) ...' All except the first of
these are localities are in the Kimberley Region, and the palms occurring there are
most unlikely to fall within the concept of L. alfredii adopted herein. They are most
likely L. kimberleyana Rodd.

9. Gardner (1923) interpreted L. alfredii in a way which cannot possibly be sanctioned
by the ICBN, even as it existed then. Under this name he cites a number of localities,
all in the Kimberley Region. In the present treatment these would probably, like
Fitzgerald's citations, represent L. kimberleyana, which could conceivably be included
within a very broad concept of L. alfredii. But, under the heading 'L. marine F.v.M.' he
cites a collection from 'the Millstream Station'! In effect he was preserving Mueller's
early (1878) concept of L. mariae encompassing both the Macdonnell Range and the
Millstream palms, and at the same time displacing the application of L. alfredii from its
type locality of Millstream (Mueller 1892, Drude 1893) to occurrences in the
Kimberleys.

10. Colour of the fruit when fresh and fully ripe has not been observed at first hand,
but at the time of my visit to 'Millstream' Station I was informed by the owners that
the fruits ripened to black or purplish-black and were quite juicy. Tike a plum'.

130

Telopea 8(1): 1998

Acknowledgments

Peter Wilson, Barbara Briggs and Barbara Wiecek assisted and persevered in the final
stages of preparing the manuscript. Angela Benn and Sue McCahon typed the
manuscript. Robert Roden was responsible for the illustrations. Don McGillivray,
Barry Conn and Ken Hill obtained photographs of type specimens and other valuable
information while in Europe as Australian Botanical Liaison Officers. Especially
relevant were collections and information from Tony Irvine and John Dowe. Surrey
Jacobs, Clyde Dunlop, Mick Jackes, Peter Lavarack, the late John Maconochie and the
Benedictine Community at Kalumburu facilitated the field studies. My 1974 field trip
around northern Australia would not have been possible without the use of a light
plane piloted by my friend Kel Stillman. Karen Wilson drew my attention to the book
Boss Drover, with mention of palms in the southern Kimberley. The Directors of
herbaria in Australia and abroad provided loans of relevant specimens. Funding for
fieldwork and technical assistance was provided by a grant from the Australian
Biological Resources Study, the Royal Botanic Gardens and Domain Trust and Dames
and Moore Consulting. My thanks go to them all.

References

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Manuscript received 14 November 1997
Manuscript accepted 2 October 1998

Fig. 1. Trunks showing petiole stubs, a, L. benthamii, Adelaide R. crossing Arnhem Highway
Northern Territory, b, L. victoriae, Victoria R. crossing. Northern Territory, c, L. eastonii, Mitchell
Plateau, north Western Australia, d, L. muelleri, Silver Plains, Queensland.

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Fig 2. a, b, L. nitida, Delusion Creek Creek-Davvson R. junction, Queensland: a, fruits: whole, split
longitudinally, and endocarp stripped of mesocarp (1 cm grid); b, ligule of young leaf opened flat
(note overlapping petiole margin lower right), c, L. muelleri, Cooktown, Queensland, inflorescence
rachis and partial inflorescence, showing rachis bracts with indumentum of scales, d, L. decipiens,
Lowmead, Queensland, hastula and lamina base.

Rodd, Revision of Livistona (Arecaceae)

135

Fig. 3. Transverse sections from leaf lamina segments, a, L. marine subsp. marine Rodd 3215.
b, L. mariae subsp. rigida Rodd 2937. c, L. mariae subsp. occidentalis Rodd 2868. d, L. lanuginosa Irvine
1912. e, L. benthamii Rodd 2904. f, L. drudei Irvine 1833. g, L. australis Rodd 3030. h, L. nitida Rodd
3055. i, L. decipiens Rodd 3077. Tissues are indicated as in Tomlinson (1961), i.e. sclerenchyma black,
xylem hatched, phloem dotted, cells of hypodermis and bundle sheaths shown. Palisade lies above
the minor veins and spongy mesophyll below.

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Telopea 8(1): 1998

Fig. 4. Transverse sections of leaf lamina, a, L.fulva Rodd 3062. b, L. muelleri Hind 748. c, L. humilis
Rodd 2911. d, L. eastonii Rodd 2888. e, L. alfredii Rodd 2848. f, L. victoriae Parker 1065.
g, L. kimberleyana Rodd 2866. h, L. lorophylla Kenneally 2162. i, L. inermis Rodd 2931.

Rodd, Revision of Livistona (Arecaceae)

137

Fig. 5. a, L. marine subsp. marine, Palm Valley, Central Australia, b, c, L. marine subsp. rigida', b, Adels
Grove, Queensland; c, Mataranka, Northern Territory, d, L. marine subsp. occidentalis, Mt King,
Durack Range, north Western Australia (Some trees higher on slope are L. kimberleyana).

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Telopea 8(1): 1998

Fig. 6. L. marine subsp. rigida. a, part of 'female' inflorescence; b, c, 'female' flower bud. From Latz
7828. Scale bar: a = 4 mm; b, c = 1 mm.

Rodd, Revision of Livistona (Arecaceae)

139

Fig. 7. L. lanuginosa, a, part of 'female' inflorescence; b, c, 'female' flower; d, stamens.
From Irvine 1912. Scale bar: a = 4 mm; b-d = 1 mm.

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Telopea 8(1): 1998

Fig. 8. a, L. benthnmii, Howard R. near Darwin, Northern Territory, b, L. drudei, Rollingstone Creek
N. of Townsville, Queensland, c, L. austvnlis, Crediton Loop Road, S of Eungella, Queensland, d,
L. nitida, Delusion Creek-Dawson R. junction, Queensland.

Rodd, Revision of Livistona (Arecaceae)

141

Fig. 9. a, L. lanuginosa, 'Nosnillor' S of Charters Towers, Queensland, b, c, L. nitida, Delusion
Creek-Dawson R. junction, Queensland, d, L. decipiens, Palm Creek S of Miriam Vale, Queensland.

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Fig. 10. L. australis, a, part of 'male' inflorescence; b, c, 'male' flower; d, stamens; e, part of 'female'
inflorescence; f, 'female' flower; g, partly dissected female flower; h, stamens, a-d from Rodd 2733;
e-h from Rodd 2734. Scale bar: a, e = 4 mm; b-d, f-h = 1 mm.

Rodd, Revision of Livistona (Arecaceae)

143

Fig. 11. L. decipiens. a, part of 'male' inflorescence; b-c 'male' flower; d, stamens; e, part of 'female'
inflorescence and flower cluster; f, g, 'female' flower; h, stamens, a-d from Hind 662; e-h from
Briggs 2071. Scale bar: a, e = 4 mm; b-d, f-h = 1 mm.

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Telopea 8(1): 1998

Fig. 12. L.fiilva. a, part of inflorescence; b-d, flowers. From Henderson 1180. Scale bar: a = 4 mm;
b-d = 1 mm.

Rodd, Revision of Livistona (Arecaceae)

145

Fig. 13. a, L.fulva, Blackdown Tableland, Queensland, b, L. muelleri, WNW of Cooktown,
Queensland, c, L. humilis, near Gove, Northern Territory. Plant on left is 'female' with long
inflorescence branched only at apex; plant on right is 'male' with shorter, much branched
inflorescence (scale bar 0.5 m long).

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Telopea 8(1): 1998

Fig. 14. L. humilis. a, part of 'male' inflorescence; b, c, 'male' flower; d, stamens; e, part of 'female'
inflorescence; f, g, 'female' flower; h, stamens, a-d from Rodd 2910; e-h from Rodd 2911. Scale bar:
a, e = 4 mm; b-d, f-h = 1 mm.

Rodd, Revision of Livistona (Arecaceae)

147

Fig. 15. a, L. muelleri, Silver Plains, north Queensland, b, L. dfredii, near Millstream, northern
Western Australia.

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Telopea 8(1): 1998

Fig. 16. L. eastonii. a, part of 'male' inflorescence; b, c, 'male' flower; d, stamens; e, part of 'female'
inflorescence; f, g, 'female' flower; h, stamens, a-d from Rodd 2888; e-h from Rodd 2887. Scale bar:
a, e = 4 mm; b-d, f-h = 1 mm.

Rodd, Revision of Livistona (Arecaceae)

149

Fig. 17. a, b, L. eastonii, Mitchell Plateau, north Western Australia, c, L. victoriae, Victoria River,
Northern Territory, d, L. kimberleyann, Mt King, Durack Range, north Western Australia.

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Te/opea 8(1): 1998

Fig. 18. L. alfredii. a, part of 'male' inflorescence; b, 'male' flower; c, partly dissected 'male' flower;
d, stamens; e, part of fruiting inflorescence, a-d from Rodd 2847; e from Rodd 2848. Scale bar;
a, e = 4 mm; b-d = 1 mm.

Fig. 19. L. lorophylla. a, part of 'male' inflorescence; b, c, 'male' flower; d, stamens; e, part of 'female'
inflorescence; f, g, 'female' flower bud; h, bud opened out; i, stamens, a-d from Johnson s.n, isotype;
e-i from Kenneally 4153. Scale bar: a, e = 4 mm; b-d, f-i = 1 mm.

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Telopea 8(1): 1998

Fig. 20. a, b, L. lorophylla, Kalumburu, north Western Australia: a, habit (scale bar 0.3 m long); b,
leaf, c, d, L. inermis, Katherine Gorge, Northern Territory: c, habit, note basal branching; d, fallen
crown with 3 inflorescences at anthesis.

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153

Fig. 21. L. inermis. a, part of 'male' inflorescence; b, c 'male' flower; d, stamens; e, part of 'female'
inflorescence; f, g, 'female' flowers; h, stamens, a-d from Rice 2037; e-h from Rodd 2931 . Scale bar:
a, e = 4 mm; b-d = 1 mm; f-h = 1.2 mm.

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Amphipogon (Gramineae) does have microhairs

L. Watson, N.P. Barker and S.W.L. Jacobs

Amphipogon is a genus of eight species endemic to Australia. The genus has usually
been treated as a member of the subfamily Arundinoideae (Watson & Dallwitz 1992,
1994) despite the similarities of the microhairs of Amphipogon to those of Enneapogon in
the Chloridoideae (Amarasinghe & Watson 1988). Barker (1997) followed Renvoize
(1981) and 'a recent survey of the genus by Linder (pers. comm.)' in stating that the
leaf blades of Amphipogon species lack microhairs, despite also citing Watson and
Dallwitz (1992) who described them in detail. At best this statement is applicable only
to abaxial leaf blade epidermes. Microhairs on the adaxial surface of Amphipogon
leaves are recorded and illustrated by Johnston and Watson (1976), Jacobs (1986) and
Amarasinghe and Watson (1988). More recent information is summarised in Watson
and Dallwitz (1992, 1994) based on light microscopy of seven species of Amphipogon
plus transmission electron microscopy of longitudinally sectioned adaxial leaf blade
microhairs of A. cnricinnus by Amarasinghe and Watson (1988). From Johnston and
Watson (1976), Jacobs (1986), Amarasinghe and Watson (1988) and Watson and
Dallwitz (1992,1994) it is clear that microhairs are regularly found on the adaxial leaf
surface as well as on the lemmas. The generic description and some illustrations are
available on the WWW (currently at http://www.biodiversity.uno.edu/delta/).

The rbcL data support the placement of Amphipogon in the Arundinoideae by Watson
and Dallwitz (1992,1994), but the presence of bicellular microhairs similar to those in
Enneapogon (Chloridoideae) justify their caution.

References

Amarasinghe, V. and Watson, L. (1988) Comparative ultrastructure of microhairs in grasses.

Botanical Journal of the Linnean Society 98: 303-319.

Barker, N.P. (1997) The relationships of Amphipogon, Elytrophorus and Cyperochloa (Poaceae) as
suggested by rbcL data. Telopea 7: 205-213.

Jacobs, S.W.L. (1986) Systematics of the Chloridoid grasses. Pp. 277-286 in Soderstrom, T.R., Hilu,
K.W., Campbell, C.S. and Barkworth, M.E. (eds). Grass Systematics and Evolution. (Smithsonian
Institution Press: Washington).

Johnston, C.R. and Watson, L. (1976) Microhairs: a universal characteristic of non-festucoid grass
genera? Phytomorphology 26: 297-301.

Renvoize, S.A. (1981) The subfamily Arundinoideae and its position in relation to a general
classification of the Gramineae. Kew Bulletin 36: 85-102.

Watson, L. and Dallwitz, M.J. (1992) The Grass Genera of the World, ed. 1. (CAB International:
Wallingford).

Watson, L. and Dallwitz, M.J. (1994) The Grass Genera of the World, ed. 2 (CAB International:
Wallingford).

Les Watson
78 Vancouver St
Albany WA 6330
Australia

156

Telopea 8(1): 1998

Nigel P. Barker
Department of Botany
Rhodes University
Grahamstown, 6140
South Africa

^Surrey W.L. Jacobs
National Herbarium
Royal Botanic Gardens
Sydney NSW 2000
Australia

* author for correspondence

Manuscript received 19 May 1998
Manuscript accepted 30 July 1998

157

SHORT COMMUNICATION

Eucalyptus canobolensis (Myrtaceae), a new combination for a
former subspecies of Eucalyptus rubida
John T. Hunter

Johnson and Hill (1991) originally published this taxon as a subspecies of E. rubida. Its
circumscription was based on the larger size of the leaves, both juvenile and adult, the
square coppice stems, the larger buds and fruits and the smaller stature of the tree.
During a recent investigation into the distribution and abundance of Eucalyptus rubida
subsp. canobolensis involving literature, field and herbarium research it became
apparent that there are sufficient distinct morphological characters and published
information to justify specific rank. Recent research (Chappill 1988; Chappill &
Ladiges 1996) indicates that this taxon, although related, is not as closely allied to
E. riibida as Johnson and Hill (1991) suggest.

R.H. Cambage made the first herbarium collection of this taxon in 1899 (NSW 325551).
Maiden (1917) commented on the collections of E. rubida made by Cambage, Boorman
and Millthorpe from Mt Canobolas and Orange. He noted that Cambage's collection
from the top of Mt Canobolas was '... an intensely glaucous form, the operculum
larger in proportion to the calyx, the valves well exerted. It has the sharp [E.] Gunnii
[sic] rims to the buds'. Maiden suggested that the Orange-Canobolas collections
showed E. rubida' s affinities to E. gunnii. Cambage's collection is illustrated in this work.

Chappill (1988) in her phylogenetic study of the section Maidenaria included samples
of this taxon (as E. sp. aff. rubida). In this study, Chappill showed that E. sp. aff. rubida
is as distinctive as other species in the series Viminales and that it was significantly
separated from E. rubida sens, strict, in cladistic analyses. Chappill and Ladiges (1996)
published cladistic analyses showing that E. rubida sens, strict, and E. rubida subsp.
canobolensis do not form a monophyletic group. In this analysis E. rubida subsp.
canobolensis is placed in a clade that includes E. gunnii, apparently justifying Maiden s
(1917) comments on the similarity of the Mt Canobolas specimens to E. gunnii.

A number of morphological characters separate E. rubida subsp. canobolensis from
E. rubida sens, strict, namely: larger adult leaves; larger juvenile leaves; larger buds;
larger fruits; smaller stature; quadrangular juvenile stems and coppice shoots; and
glaucous buds and fruits (Table 1). Furthermore, the phylogenetic affinities of this
taxon are masked if it retains subspecific rank within E. rubida. Accordingly, I make the
new combination:

Eucalyptus canobolensis (L.A.S. Johnson & K.D. Hill) J.T. Hunter, comb, et stat. nov.

Basionym: Eucalyptus rubida subsp. canobolensis L.A.S. Johnson & K.D. Hill., Telopea 4:
239 (1991).

Holotype: New South Wales: Central Tablelands: Mt Canobolas, Orange, J.L. Boorman,
Jan 1908 (holo: NSW [NSW 325550]).

Illustrations: Maiden (1917: pi. 109 5a & b) as E. rubida; Hill (1991: 113 fig. 104b) as
E. rubida subsp. canobolensis.

Distribution: this species is restricted to the higher altitudes (c. > 1000 m) of Mt
Canobolas, south west of Orange on the Central Tablelands of New South Wales.

Habitat: Eucalyptus canobolensis occurs predominantly above 1100 m, but can be found
less abundantly down to 1000 m. The species is most common between 1200 and 1300 m

158

Telopea 8 ( 1 ): 1998

altitude. Above 1300 m E. pnuciflora becomes more prominent. Basalt occurs over most
of Mt Canobolas with many acid volcanic trachytic outcrops. The taxon occurs in sub-
alpine woodland and is associated with E. pnuciflora, E. dalrympleana, E. viniinalis and
E. dives as an overstorey and Poa sieberiana, Cassinia arcuata, Acaena ovina, Hydrocotyle
algida and Scutellaria humilis in the understorey.

Conservation status: Johnson and Hill (1991) allocated a ROTAP code (Briggs & Leigh
1996) of 2VC- for this taxon. This species was subsequently listed on the Threatened
Species Conservation Act (NSW Government TSC Act 1995) as vulnerable (Schedule 2).
Hunter (1998) has established that although the species is restricted to Mt Canobolas,
there are probably over 60 000 individuals. Briggs and Leigh (1996) suggest that 1000
individuals in a reserve is adequate for conservation purposes. This species dominates
most areas within the Mt Canobolas State Recreation Area and the evidence indicates
that this species is not vulnerable. A more appropriate ROTAP code is 2RCa and
removal from the TSC Act is warranted.

Table 1. Comparison of selected characters for Eucalyptus rubida sens, strict, and E. canobolensis.

Plant height
Adult leaf length
Adult leaf width
Juvenile leaf length
Juvenile leaf width
Juvenile stems
Bud length
Bud width
Fruit length
Fruit width

Buds and fruits glaucous

Eucalyptus rubida

to 40 m

3- 14 cm

0.8-2.4 cm

4- 5 cm
2.5-5 (-6) cm
Round

4-7 mm
2-4 mm

4- 6 mm

5- 7 mm
Sometimes

Eucalyptus canobolensis

8-12 m tall (rarely to 18 m)

(6.5-) 10-18 cm

1.5-3.5 cm

(4-) 5-10 cm

Quadangular

5-9 mm

3-5 mm

5- 9 mm

6- 8 mm
Always

References

Bn ggS' J-D. & Leigh, J.H. (1996) Rare or Threatened Australian Plants, revised edition. (CSIRO and
the Australian Nature Conservation Agency: Canberra).

Chappill, J.A. (1988) A Systematic Study o/Eucalyptus L'Her it. informal Subgenus Symphyomyrtus
Section Maidenaria (Myrtaceae). Unpublished PhD thesis. University of Melbourne.

Chappill, J.A. & Ladiges, P.Y. (1996) Phylogenetic analysis of Eucalyptus informal subgenus
Symphyomyrtus section Maidenaria. Austral. Syst. Bot. 9: 71-93.

Hill, K.D. (1991) Eucalyptus. Pp. 76-142 in Harden, G.J. (ed.) Flora of New South Wales, vol. 2. (New
South Wales University Press: Sydney).

Hunter, J.T. (1998) Threatened Endemic Eucalypts of the Central Tablelands. Unpublished report for the
Western Zone New South Wales National Parks and Wildlife Service (Dubbo).

Johnson, L.A.S. & Hill, K.D. (1991) Systematic studies in the eucalypts-3 New taxa and
combinations in Eucalyptus (Myrtaceae). Telopea 4: 223-267.

Maiden, J .H. (1917) A Critical Revision of the Genus Eucalyptus, vol. III. (Government Printer: Sydney).

John T. Hunter

149 Kentucky St

Arrnidale, NSW 2350

Australia

Manuscript received 30 June 1998
Manuscript accepted 2 October 1998

NOTICE TO AUTHORS

Telopea is published twice-yearly, in March and September. Preference will be given to papers relating to
the flora of New South Wales. Brief papers may be published as Short Communications.

Deadlines for the submission of papers are 1 June (for the March issue the following year) and 1 November
(for the following September issue). Authors are expected to have had their papers peer-reviewed before
submission. All papers will be refereed. Two copies of the manuscript should be submitted along with
originals of photographs and clear photocopies of all other figures. The full postal address (plus telephone
and fax numbers) of the author who will check the proofs and receive correspondence should be included.
Once a paper has been accepted for publication the author should provide the paper on a computer disk
along with final artwork. The disk should be in IBM compatible (MS-DOS) or Macintosh format and
clearly labelled with the word processing program used and the file name(s).

General formatting requirements • Text should be justified. This applies also for abstracts, headings, keys
and reference lists. Headings should be in upper and lower case, and not underlined. • Use only a single
space after all punctuation marks including fullstops. • Insert a space between a numeral and unit of
measurement, e.g. 3 mm, but no space between initials, e.g. L.A.R. Haegi, or between en dashes and
associated numerals, e.g. 5.2-6 mm or between extreme measurements and ranges e.g. (10—)25—35(—90).
Use double hyphens (—) to indicate en dashes (-). • Do not use the spacebar to indent or tabulate. Use
italics where necessary, and use single quote marks before double.

Organisation of the paper The title should be explicit and descriptive of the content. Include the family name
and broad geographic region where appropriate. Abstracts (except for Short Communications) should
be included. Check most recent issue for format. Bracketed keys are preferable especially for long keys,
but indented keys are acceptable. Long indented keys should be divided into groups. When giving
authors of botanical names follow the forms in the Kew Draft Index of Author Abbreviations. But note
unabbreviated use in references below.

Types Cite details in full, giving details from protologue and from specimen label separately if there
are important differences. Type citations should be in a consistent format, e.g: Type: New South Wales:
North Western Plains: 10 km W of Moree (29°08' S 129°48' E), B. Wiecek 1250, 2 Jan 1989; lecto NSW
(Weston 1990: 21); isolecto K, MO.

Selected specimens Cite no more than 20 (except for very widely distributed species) and arrange by
Botanical Divisions. Use accepted format: locality, collector & number, date (herbarium code plus
institutional number if there is no collector's number) and omit the initials of collectors, unless confusion
is likely. Only latitudes and longitudes on the original labels should be included. Give dates in the
following format: 12 Jan 1987, 2 June, 30 July, 10 Dec etc.

References In formal taxonomic citations use the fully abbreviated (Harvard) form: author (year: page) e.g.
Bentham (1878: 234). The traditional - abbreviated form, e.g. Bentham, FI. Austral. 7: 234 (1878), may be
used in shorter papers. Authors' names in these citations should be given unabbreviated. References to
books published before 1900 need not include the publisher and place of publication, but be consistent.

Index to taxa This is useful if the paper is large and deals with many species and synonyms. The author
should prepare the basic alphabetic listing including all names in recent use.

Illustrations/maps/photos • Supply bromides, photographs, transparencies or good quality artwork with
the final manuscript. Check that maps show their context clearly (by lat/longs or an inset map) and that
relevant place names in the text (but not for cited specimens) are shown. • Photos should be unmounted,
good-quality prints. Do not label photos. • Labelling that is part of an illustration, e.g. place names on a
map, should be added in Helvetica font by the person preparing the illustration. • Bar scales on the figure
are preferable to numerical scales in the caption. Any magnification levels in the caption should refer to
the size of the submitted original figure (not anticipated final size). • The maximum final size of the
illustration is 205 mm high x 125 mm wide.

Captions Use lower case letters for the parts of a figure e.g. Fig. 1. Jacksonia michaeliana. a, stem tissue (x 10);
b, calyx lobes (x 0.5). (a from holotype; b from Barson 234.)

Tables should preferably be portrait rather than landscape shape.

More detailed instructions are available on request from the editors.

TELOPEA 8(1): 1-158 • December 1998

ISSN 0312-9764

Contents

Dilhvynia glaucula (Fabaceae: Mirbelieae), a new species from the Southern
Tablelands, New South Wales Peter C. Jobson and Peter H. Weston 1

Aponogetonaceae of Australia, with descriptions of six new taxa

C. B. Hellquist and S.W. L. Jacobs 7

New combinations arising from a new classification of non-African
Restionaceae Barbara G. Briggs and L.A.S. Johnson 21

Two new rare species of Homoranthus tMyrtaceae: Chamelaucieae) from

the Northern Tablelands of New South Wales John T. Hunter 33

Nassella tenuissima (Gramineae) recorded from Australia, a potential new
weed related to Serrated Tussock

S.W.L. Jacobs, J. Everett and Maria Amelia Torres 39

Notes on Australian Meliaceae D.J. Mabberley 45

Revision of Livistona (Arecaceae) in Australia A.N. Rodd 49

SHORT COMMUNICATIONS

Amphipogon (Gramineae) does have microhairs

L. Watson, N.P. Barker and S.W.L. Jacobs 155

Eucalyptus canobolensis (Myrtaceae), a new combination for a former
subspecies of Eucalyptus rtibida John T. Hunter 157

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