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VOLUME 3
NUMBER 1 1989

AUSTROBHILEYA

A JOURNAL OF PLANT SYSTEMATICS

Or QUEENSLAND HERBARIUM
A DEPARTMENT OF PRIMARY INDUSTRIES
QUEENSLAND GOVERNMENT

Editorial Committee

E.M. Ross (editor)
R.J.F. Henderson (technical advisor)

Word Processing Cover design & Graphics
C.G. Watkins
K.M. Ryan W.A. Smith
J.M. Cronk
Austrobaileya
Vol. 1, No. | was published on 1! December 1977
Vol. 2, No. 14 July 1984

]

2 29 August 1985

3 24 July 1986

4 24 September [987
5 5 October 1988

Austrobaileya is published once per year.
Exchange: This journal will be distributed on the basis of exchange.

Subscriptions: Orders for single issues and subscriptions may be placed. The price is
(1989) A$20 per issue for individuals, A$35 for institutions, including postage.

All correspondence relating to exchange, subscriptions or contributions to this journal
should be addressed to The Editor, Austrobaileya, Queensland Herbarium, Department
of Primary Industries, Meiers Road, Indooroopilly Q. 4068. |

ISSN 0155-4131

© Queensland Government 1989
Queensland Department of Primary Industries
GPO Box 46
Brisbane 4001

Austrobaileya 3(1): i-172 (1989)

CONTENTS
A new species of Coix L. (Poaceae) from Australia
B.K. Simon ee) ee ae ee

Ceropegia cumingiana Decne Sra. dataa teal
P.V. Bruyns & P.j. Forster .

The genus Zornia J. Gmelin Fon soe ta in Australia
S.T. Reynolds & A.E. Holland .

Two new species of Eucalyptus inate from central Queensland
A.R. Bean & M.I.H. Brooker Ftetote lf eae wi nary ake Hanae Pd

A new species of Marsdenia R. Br. hiatal apie og: from eastern Australia
J.B. Williams . Figece Jah SOL er a eee

Kailarsenia Tirvengadum emend. Puttock (Rubiaceae: Gardenieae) in
Australia
C.F. Puttock

The genus Ancana F. Muell. (Annonaceae) in Australia
L.W. Jessup sate ee ey NRE OBL une Bie ong,

Secamone R. Br. (Asclepiadaceae: Secamonoideae) in Australia
P.I. Forster & K. Harold... .......... 0.0...

Studies in Australian grasses: 4. Taxonomic and nomenclatural studies
in Australian Andropogoneae
B.K. Simon anloees

Revision of the Australian Vitaceae, 4. Clematocissus Planchon
B.R. Jackes rye eee fe 76 FE eats
Bryophytes in a ee ORES mangrove CONTE
J WAR OIE 40s be Bed eee nk )

Notes on Asclepiadaceae, |
P.I. Forster

Notes on Trachymene ss ie feibiats in Bie ea l
A.E. Holland . kas re me

New species of Bulbophyllum section Oxysepalum (Orchidaceae) in

Australia
B. Gray & D.L. Jones .

Revision of the Australian Vitaceae, 5. 7 a toeaiie a aa Planchon
B.R. Jackes 21553) 6. 28 otek : re Oe aw

NOTES
Eriocaulon longifolium Nees ex Kunth (Eriocaulaceae), a new record for

Australia
G.J. Leach......

PLANT PROFILE

Austrobaileya scandens C. White
E.M. Ross .

Book Reviews ..

Corrigendum

13

39

45

51

63

69

79

LO]

1Q3

109

135

14]

149

159

163
167
172

Austrobaileya 3(1): 1-5 (1989) [

A NEW SPECIES OF COIX L. (POACEAE)
FROM AUSTRALIA

B.K. Simon

Queensland Herbarium, Meiers Road, Indooroopilly, Qld 4068

Summary

Coix gasteenii sp. nov., from northern Queensland, is diagnosed and compared with other known species of Coix.

Coix gasteenii B. Simon species nova afhinis C. /acryma-jobo L. sed utriculis
inflorescentiarum feminearum laminis et culmis rhizomatosis differt et affinis C.
ouwehandio Koord. sed laminis utriculorum inflorescentiarum feminearum
longioribus differt. Typus: Queensland. Cook District, Lakefield National Park,
1S°18’S, 144°36’E, Cabbage Tree Creek - 19 km E of Old Laura on road to
Cooktown, small branch of main channel ca 100 m upstream from road crossing,
13 May 1987, J.R. Clarkson 7215 & B.K. Simon (holo: BRI(AQ422941 - 3 sheets
BRI 422940, BRI 422941 and BRI 422942) iso: DNA,K,L,MBA,MEL,MO,
NSW,PERTH,PRE,QRS,SP,US). Figs 1-3.

Additional specimen: Queensland. Cook District: Lakefield — east of Cabbage Tree Creek, May 1980, Gasteen
50 (BRI).

This new grass was first brought to my attention in 1980 when an apical portion
of the plant was sent to the Queensland Herbarium for identification as part of a plant
collection made by Mr W.G.(Jim) Gasteen, during field work prior to the gazetting of
the area as the Lakefield National Park. At the time the fragmentary nature of the
material at hand revealed that it was different from C. /acryma-jobi, the only species
known to occur in Australia at the time, but the material was inadequate to use as a
basis for drawing up a description. Correspondence with Mr Gasteen established the
exact locality of the collection and John Clarkson and I were able to collect complete
material in May 1987. The new species 1s named after its first collector, a keen amateur
naturalist, who has conducted a number of vegetation surveys for the Queensland
National Parks and Wildlife Service, within the Department of Environment and
Conservation.

Gasteen’s original collection gave the impression that the male and female
inflorescences arose independently in the upper leaf axils. However examination of the
type material showed that their morphological arrangement is the same as that in C.
lacryma-jobi, which itself can be quite variable (Jacques-Felix 1961). It was noticed that
the male inflorescence may be produced at the apex of only one female involucre (utricle)
or a short chain of them. With this knowledge the Gasteen fragment was further examined
and it was discovered that the male racemes did arise from the apex of very immature
involucres which were affected by fungal damage.

A unique feature of C. gasteenii, when compared with C. lacryma-jobi, is the
extension of the utricle into small leaf blades. These are much smaller than the culm
leaf blades but have the same general shape and tuberculate margins. Morphologically
the utricle has been thought to be a metamorphosed leaf sheath and confirmation of
this is readily shown in C. gasteenii. This feature 1s developed to a much lesser extent
in some other species e.g. C. ouwehandii Koord. (Koorders 1919) or else is of abnormal
occurrence in others e.g. C. aquatica Roxb. (Jain & Banerjee 1974). Another striking
morphological character of C. gasteenii when comparing it with C. lacryma-jobi is the
rhizomatous base of the culm. Whether this occurs in other species is not certain as
either the character is not mentioned in species descriptions or adequate material of
species other than C. gasteenti and C. lacryma-jobi was not available for examination.

The morphological differences between C. gasteenti and C. lacryma-jobi may be
seen from the following descriptions of the two species generated by the DELTA system
(Dallwitz & Paine 1986).

Austrobaileya 3(1): 1989

Simon, Coix ;

C. gasteenil

Habit perennial; rhizomatous; 100-180 cm tall. Leaf blades 9-150 cm _ long,
{0-23 mm wide; with distinct forwardly pointing tubercles. Ligule 0.3 mm long. Utricles
(female involucres) 2 or 3 per inflorescence; continued apically into small leaf blade; 5-
7 mm long, 4-5 mm wide. Lower glume 7-9 mm long. Upper glume 8-10 mm long.
Lower lemma 8-9 mm long. Palea present (6-7 mm long). Upper lemma 7-8 mm long.
Palea 7-8 mm long. Caryopsis 4-5 mm long, 3-4 mm wide. Male racemes 1.4-1.8 cm
long; 3—5-jointed. Lower glume 6-7 mm long; 9-nerved. Upper glume 6-7 mm long; 9-
11-nerved. Lower lemma 6-7 mm long. Palea 5—~6 mm long. Upper lemma 5-6 mm
long. Palea 5-6 mm long.

C. jacryma-jobi

Habit annual; not rhizomatous; 90-250 cm tall. Leaf blades to 50 cm _ long,
2-4 mm wide; minutely tuberculate. Ligule to 0.5 mm long. Utricles (female involucres)
{[~3 per inflorescence; not continued apically into small leaf blade; 9-11 mm long; 7~9
mm wide. Lower glume to 9 mm long. Upper glume to 8.8 mm long. Lower lemma to
8 mm long. Palea absent. Upper lemma to 7.6 mm long. Palea to 5.4 mm long. Caryopsis
5-6 mm long, 4-5 mm wide. Male racemes 1—1.5 cm long; 3-5-jointed. Lower glume
to 9 mm long; |1l-nerved. Upper glume to 8 mm long; 11-nerved. Lower lemma to 38.4
mm long. Palea to 8.2 mm long. Upper lemma to 7.2 mm long. Palea to 6.8 mm long.

C. gasteenii differs from C. ouwehandii by its rhizomatous habit and by having
longer leaf blades from the utricles. However no material of the latter species was
examined and morphological details for comparison were those of Koorders (1919).

Coix is a tropical Asian grass genus with a natural distribution from India to
Japan to Papua New Guinea, with the Malesian Archipelago being suggested as a centre
of origin (Vallaeys 1948). It consists of five or six previously known species (Clayton &
Renvoize 1986; Nirodi 1955) and C. gasteenii. The best known and most widely spread
species is C. facryma-jobi, which has been introduced throughout the tropical and warm-
temperate regions of the world and has been classified into a number of varieties (Bor
1960; Mimeur 1951). The hard globose to fusoid and lustrous utricles (female involucres)
of this species are coloured white to bluish and have been used for a variety of decorative
purposes. Soft-shelled edible varieties have been grown in various areas of tropical Asia,
Africa and America. Although comparing favourably as a food source with wheat, rice
and corn, it has usually been replaced by one of these crops in regions where it has
been introduced.

The other species of Coix have a comparatively narrow distribution. C. gigantea
Koenig ex Roxb. and the closely related aquatic form C. aquatica Roxb. occur throughout
the range of the genus, although the latter has not been reported from Sri Lanka
(Lazarides 1980). A couple of Queensland Herbarium specimens cultivated or naturalised
in Queensland under the name C. /acryma-jobi have the male raceme dimensions of C.
gigantea and for this reason they are more appropriately placed there, e.g. J.W. Junes
AQ?2?81408 (BRI) from near Woombye and M. Cameron 118 (BRI) from Weipa. C.
puellarum Balansa occurs in Indo-China, Thailand and Burma (Lazarides 1980). The
two remaining species are narrow endemics, C. ouwehandii Koord. in Sumatra and C.
poilanei Mimeur in Laos (Nirodi 1955); for some reason they are not mentioned by
Lazarides (1980) although they are found within the geographical boundary defined by
this work.

Fig. 1. Coix gasteenti: A. habit X 0.25. B-J. female inflorescence. B. involucre (utricle) and associated leaf blade
and rachis X 2. C. L.S. of utricle showing female spikelets in situ X 3. D. portion of rachis within utricle x 3.
E. sessile sptkelet and two associated rudimentary pedicelled spikelets <x 3. F—J. sessile spikelet all * 3. F. lower
elume. G. upper glume. H. lower floret showing lemma and palea. I. upper floret showing lemma, palea and
young caryopsis. J. mature caryopsis. K. male raceme X 2. L-M. opposing views of spikelet triad of one pedicelied
and two sessile spikelets x 3. N—U. sessile spikelet. N~O. lower and upper glumes X 3. P-R. lower floret. P.
lemma X 3. Q. palea X 3. R. lodicules and anthers < 6. S—U. upper floret. S. lemma X 3. T. palea X 3. U. anther
x 6. All from hoio and isotype specimens, BRI.

Austrobaileya 3(1): 1989

Fig. 3. Colx gasteenii: A. photograph of several utricles (female inflorescences) and one male inflorescence. B.
| photograph of plant in natural habitat being examined by J.R. Clarkson. Both of type gathering.

Simon, Corx 5

Acknowledgements

I am grateful to Jim Gasteen for detailed notes on his collecting locality, to John
Clarkson, Botany Branch, Mareeba and Peter Stanton, National Parks and Wildlife
Service, Cairns for assistance in finding this locality, and for permission of the Queensland
National Parks and Wildlife Service to collect in a National Park.

References
BOR, N.L. (1960). Grasses of Burma, Ceylon, India and Pakistan. Oxford: Pergamon Press.
CLAYTON, W.D. & RENVOIZE, S.A. (1986). Genera graminum. Kew Bulletin Additional Series 13.

DALLWITZ, M.J. & PAINE, T.A. (1986). User’s guide to the DELTA system. A general system for processing
taxonomic descriptions, 3rd edition. CSIRO, Australian Division of Entomology Report No. 13.

JACQUES-FELIX, H. (1961). Observations sur la variabilite morphologique de Coix lacryma-jobi. Journal
‘Agriculture Tropicale et de Botanique Appliquee 8: 44-56.

JAIN, S.K. & BANERJEE, D.K. (1974). Preliminary observations on the ethnobotany of the genus Coix. Economic
Botany 28: 38-42.

KOORDERS, S.H. (1919). Beschreibung einer von Dr. Ouwehand im Toba-See, in Sumatra entdeckten neuen
Art von Coix. Bulletin du Jardin Botanique de Buitenzorg. Ser. 3, 1: 190-191, pl. 20.

LAZARIDES, M. (1980). The tropical grasses of southeast Asia. Vaduz: J. Cramer.

MIMEUR, G. (1951). Systematique specific du genere Coix et systematique varietale de Coitx flacryma-jobi.
Morphologique de cette petite cereal et etude de sa plantule. Revue International de Botanique Appliquee et
d’Agriculture Tropicale 31: 197-211.

NIRODI, N. (1955). Studies on Asiatic relatives of maize. Annals of the Missouri Botanical Garden 42: 103-130.

VALLAEYS, G. (1948). Le “Coix Lacryma-Jobi.” Bulletin Agricole du Congo Belge 39: 247-304.

Accepted for publication 20 December 1988

4

Austrobaileya 3({1): 7-11 (1989) 7

CEROPEGIA CUMINGIANA DECNE (ASCLEPIADACEAE)
P.V. Bruyns
Bolus Herbarium, University of Cape Town, Rondebosch, 7700 South Africa
and P.[. Forster

Botany Department, University of Queensland, St Lucia, Qld 4067

Summary

The recognition of subspecies in Ceropegia cumingiana Decne is unwarranted. An illustrated taxonomic account
of C. cumingiana and notes on habitat, distribution, ecology, affinities, variation, conservation status and ethnobotanical
use are provided. A lectotype is chosen for C. perforata N.E. Br.

Introduction

Species of Ceropegia L. are widely distributed in Africa, Arabia, Pakistan, India
and south-east Asia with the greatest concentration occurring in southern Africa (Dyer
1980, 1983) and peninsular India (Ansari 1984).

Ceropegia cumingiana Decne was first described from material collected in the
Philippine Islands by Cuming (Decaisne 1844). Huber (1957), in his monograph of the
genus, recognised one taxon (C. cumingiana subsp. horsfieldiana (Miq.) H. Huber) for
Java, Lesser Sunda Islands, Celebes, New Guinea and Australia, and one taxon (C.
cumingiana subsp. cumingiana) for the Philippine Islands. This paper provides a detailed
description of C. cumingiana. The nomenclature and infraspecific taxonomy of the
species and its synonymns are revised.

Taxonomic Treatment

Ceropegia cumingiana Decne in DC., Prod. 8: 643 (1844). Type: The Philippines. Luzon:

Calauang, Province of Laguna [fide Merrill 1915], H. Cuming 447 (holo: G(3
SEIN photo at BRI!); iso: BM,K(2 sheets, photo at BRI!),L(2 sheets),P(photo
BRI})).
Hook., Bot. Mag. 74: t. 4349 (1848); Miq., Fl. Ned. Ind. 2: 528 (1856); F. Muell.,
Fragm. 5: 159-160 (1866); Benth., Fl. austral. 4: 348 (1869); Fernandez-Villar in
Blanco, Fl. Filip., ed. 3, t. 313 (1880); F. Muell., Syst. census Austral. pl. 94
(1882); F. Muell., Second syst. census Austral. pl. 1: 159 (1889); K. Schum., FI.
Kais. Wilh. Land 110 (1889); Bailey, Queensl. fl. 3: 1015 (1900); K. Schum. &
Lauterb., Fl. Schutzgeb. Sidsee 508 (1901); Schitr., Nachtrage Fl. Schutzgeb.
Siidsee 354 (1905); Bailey, Compr. cat. Queensl. pl. 332 (1913); Merrill, Enum.
Philipp. fl. pl. 3; 343-344 (1923); Elmer, Leafl. Philipp. Bot. 10: 3548 (1938);
Tsiang, Sunyatsenia 4: 91 (1939); H. Huber, Mem. Soc. Brot. 12: 56-58 (1957);
Jones & Gray, Austral. Climbing pl. 88 (1976).

C. horsfieldiana Miq., Fl. Ned. Ind. 2: 528 (1856); C. cumingiana subsp. horsfieldiana
(Miq.) H. Huber, Mem. Soc. Brot. 12: 56 (1957). Type: Indonesia. Java. Soerakarta,
1802-1818, 7. Horsfield sn. (syn: K(photo BRI); Bali, Zollinger 1013 (syn:
P(photo at BRI!)).
panes Nachtrage Fl. Schutzgeb. Siidsee 354 (1905); Tsiang, Sunyatsenia 4: 90
(1939).

C. curviflora Hassk., Flora 40: 102 (1857). Type: Indonesia. Bali. Lobd s.n. (holo:
L!).

Tsiang, Sunyatsenia 4: 290 (1940); Bakhuizen van den Brink, Blumea 6: 373

(1950); Backer & Bakhuizen van den Brink, Fl. Java 2: 257-258 (1965).

C. obtusiloba Fawcett in Forbes, Nat. Wand. in East Archip. 511 (1885). Type:
Indonesia. Timor. [Kailakuk in the Kingdom of Bibicucu], [26-28 Apr 1883],
Forbes 3801 (holo: BM(photo BRI')).

8 Austrobatleya 3(1): 1989

C. perforata N.E. Br., Bull. Misc. Inform. 1901: 141 (1901). Type: Introduced from
New Guinea, Hort. F. Sander & Co., 13 August 1898 (lecto (designated here):
K(photo BRIJ)).

C. papuana Schitr., Bot. Jahrb. Syst. 50: 163 (1914). Type: New Guinea. “Kaiser
Wilhelmsland”, March 1887, M. Hollrung 659 (isosyn: MEL!); Im Strandgebiisch
bei Konstantinhafen, March 1902, R. Schlechter 14300 (syn: B +); Zwischen
Gestrauch hochsteigend in dem Sekundarwald am Djamu, December 1907, R.
Schlechter 16969 (syn: B 7).

Jones & Gray, Climbing pl. Austral. 177, 193 (1988).

C. borneensis Merr., Philipp. Jour. Sci. 13: 120 (1918). Type: Malaysia. Sabah (North
Borneo), Clement 9810 (n.¥.).

C. merrillti Schitr., Feddes, Repert. Spec. Nov. Regni. 13: 565 (1915); C. cumingiana
subsp. cumingiana forma merrillii (Schitr.) H. Huber, Mem. Soc. Brot. 12: 58
(1957). Type: The Philippines. Luzén. Bangued (“Benguet’’) distr., Bued River,
Oct-Nov 1905, E.D. Merrill 4863 (holo: K(photo BRI)).

Merrill, Enum. Philipp. fl. pl. 3: 344 (1923).

Perennial herb, sap clear. Rootstock a cluster of white, fleshy, fusiform roots; each root
{1-3 mm diameter, up to [5 cm long. Stems glabrous, twining, succulent, aerial parts
pale green, deciduous, to 5 mm diameter; internode length up to 15 cm. Leaves ovate-
elliptic, base rounded to cordate, tip acute; 3 or 4 main vein pairs, 2 or 3 minor vein
pairs; 4 extrafloral nectaries at lamina base; 20-130 mm long, 15-70 mm wide; petiole
25-40 mm long, 1-2 mm diameter; pale green, glabrous. Flowers borne near apex of
stem, arising at nodes, displaced to position between petioles of leaf pair; 1—20-flowered,
on glabrous, pale green peduncle to 80 mm long, 1 mm diameter. Pedicels 15-22 mm
long, 1.5 mm diameter, glabrous, pale green or slightly mottled with purple. Sepals
linear-lanceolate, acute-tipped, glabrous, 3-3.5 mm long, | mm wide; 2 or 3 glands at
base of confluence of each pair of lobes. Corolla tube 12-20 mm long, 3-6 mm diameter,
slightly inflated at base; exterior glabrous, tube with horizontal colour bands rising
vertically trom base, firstly of cream, then purple-black, then cream with deep purple-
red flecking or overall cream with pale purple blotches; interior with white, uniseriate
retrorse trichomes in inflated basal part, pale purple-red or cream. Corolla lobes connate
near tips, replicate where margins free, 6-15 mm wide; individual lobes 7-11 mm long,
2-6 mm wide, glabrous, with horizontal colour bands rising vertically from base, firstly
of cream, then yellow and tips red or alternatively firstly purple-red, then yellow, then
brown-red. Corona 3.5-4 mm long, 3.5-5 mm diameter, forming 3 cupular pockets
around staminal column. Outer corona 2.5 mm long, 3.5-5 mm _ diameter; each lobe
bifid into erect teeth, c. | mm long, | mm wide with white, uniseriate cilia 0.5 mm
long, either irregularly striped with purple and yellow or yellow at base with purple-red
tips. Inner corona 1.5-2.5 mm long, 1-1.5 mm wide, individual lobes cylindrical, 0.5
mm diameter; tips rugose, with few scattered uniseriate setae at tips; yellow to yellow-
cream; lobes fused to base of anthers for lower | mm. Staminal column 2~2.5 mm long,
1.5-2 mm diameter; lobes spathulate, Q.5 mm long X 0.5 mm wide; anther-wings 0.5-
Q./5 mm long. Style head 1-1.5 mm diameter. Ovaries c. 2 mm long, c. 1.5 mm wide.
Pollinia globose, inner top margin pellucid for 0.2 mm, yellow, 0.35-0.4 mm long,
0.29-0.32 mm wide. Corpusculum oblong, tan-brown, 0.33-0.35 mm long, 0.15 mm
wide. Caudicles translucent, 0.15 mm long, 0.12 mm wide. Follicles fusiform, green and
purple mottled, paired or single, to 21 cm long, 3-5 mm diameter. Seed oblong, tan-
brown, 14 mm long, 2 mm wide; coma white, up to 60 mm long. Fig. 1.

Selected specimens: The Philippines. Luzon. Bosoboso, Province of Rizal, Sep 1904, Ahern’s collector [Ramos],
Forestry Bureau No. 1854 [on mixed sheet with Cuming 447 & Vidal 1601] (K(photo BRI!)); Indonesia. Java.
Tambajangan, Kangean, Mar 1919, Backer 27541 (L); Tempoeran, SE of Semarang, Apr 1910, van Leeuwen (L):
Pekalongan, Bosch district, Apr [920, Beimee 5144 (L); Kedoe, Apr 1938, Brinkman 865a (L); Tanimbar Is,
Yamdena, Malentian, Apr 1956, v. Borsstum Waalkes 3322 (L). Sulawesi (Celebes). Palu Is, Apr 1957, Loeters
1302 (L); Flores Is, West of Nunang-Paku, Feb 1967, Schmutz 1304 (L). Timor. banks of Noil Fatoe, Apr 1929,
Walsh 208 (L). Papua New Guinea. Morose District: Bulolo Valley, Jun 1955, Floyd & Havel NGF7458
(BRI,LAE),; Wau-Salamaua Road, Kaisenik, Upper Bulolo Valley, Dec 1955, Womersley & Millar NGF7828
(LAE); Umi River Bridge, 6°15’S, 146°1S’E, van Roven NGFI606l (BRI,CANB,L,LAE); McAdam National Park,
side of Wau-Bulolo Road, Mar 1978, Verdcourt 5116 (LAE). CENTRAL District: between Maipa & Mankina
Villages, Kairuku subdistrict, Sep 1962, Darbyshire 957 (CANB). NORTHERN District: Kokoda, 8°53’S, 147°44’E,
Mar 1936, Carr 16260 (CANB). Australia. Cook District: Lockerbie, 10 miles [16 km] WSW of Somerset, {948,
Brass 18396 (A,BRI,CANB,L), 2.7 km past Lockerbie HS site on Cape York rd, 10°47’S, 142°29’E, Jun 1988.
Forster 4420 & Liddle (BRI); Weipa, Lake Patricia, [2°39’S, 141°50’E, Apr 1988, Forster 4075, Liddle & Bostock

(BOL,BRI*); Lamond Hill, Iron Range, {2°43’S, 143°18’E, Apr 1988, Forster 4208 & Liddle (BRI); MclIlwraith
Range, 13°52’S, 143°t7’E, Apr 1988, Liddle [AQ408494] (BRI*); Mt White near Coen, Jul £949, Flecker

yoTE SM MMM RUSE ROME SES Rhett SEE EEE EEA DAN EW NI DEAN hy i SME? ayy EA Pa BD Mh? a a See e ene ean ene Ham howto wih tty aaah Me vn Me Me a a » +e Pee ee)

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= 0.25 mm).

a cumingiana
ted corolla. E. corolla-lobe {scale 3 mm). F. face v

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. side view of staminal column (scale 1 mm). H. side v
(scale of B

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Gray 212 (very small flowered form).

Fig
I. poll

10 Austrobaileya 3(1): 1989

N.Q.N.C.13286 (BRI); Stannary Hills, 17°18’S, 145°13’E, Jan 1977, Gray 212 (QRS*). (* indicates material

preserved in spirit) Specimens from 35 localities examined.

Distribution, habitat and ecology: C. cumingiana is known from Bali in the south-west,
Borneo in the north-west, Papua New Guinea in the east, the Phillipine Islands in the
north to northern Queensland in Australia in the south. It occurs in deciduous microphyll
and notophyll vine forests in northern Queensland and Papua New Guinea.

At Lake Patricia (Forster 4075 et al.) and Lockerbie (Forster 4420 & Liddle) the
soil is a mixture of fine sand and humus while at Lamond Hill (Forster 4208 & Liddle)
it is of volcanic origin and red in colour. The species is probably pollinated by small
drosophilid flies of which several have been found inside the flowers of Gray 212. Larvae
of a Danaus sp. (probably either D. plexippus plexippus (L.) or D. chrysippus petilia
(Stoll)) (Nymphalidae) were observed feeding on leaves of C. cumingiana at Lake Patricia.

Notes: The locality for Cuming’s type collection of C. cumingiana is taken from the
listing of Merrill (1915).

There are two sheets at K labelled as “Type Specimen’ for C. perforata. One has
mainly juvenile leaves, no flowers, a red printed “Type Specimen’ label and an annotation
in N.E. Brown’s hand stating ‘Ceropegia perforata, N.E. Br.! Hort. F. Sander & Co. Dec
20. 1897 Imported from New Guinea’. The other consists of mature leaves with four
pressed cymes of flowers and is labelled in N.E. Brown’s hand “Ceropegia perforata, N.E.
Br.! in Kew Bull. 1901, p. 141 Type Specimen! Hort. F. Sander & Co. Aug. 13. 1898
Introduced from New Guinea N. E. Brown’. Brown (1901) did not specify a date for
his type and hence the latter specimen is designated here.

The live or pickled material examined (in particular Gray 212, Forster 4075 et
al., Liddle AQ408494 and Kitchen AQ408495) demonstrates the wide variation possible
both in the size of the flowers and degree of pubescence on floral parts in this taxon.
Specimens of Kitchen AQ408495 (cultivated at Tolga, but collected at Lockerbie Scrub)
were observed to produce flowers of different sizes in relation to the general moisture
content of its environment (G. Sankowsky, pers. comm. 1988). A plant of Gray 212 had
flowers that often lacked the small hairs on the corolla lobes.

Subspecies horsfieldiana was distinguished from subsp. cumingiana by the generally
longer corolla lobes, pubescent interior lower portion of corolla tube and the pubescent
interior of corona (Huber 1957). Both flower colour and the presence or absence of hairs
on various floral parts are very variable in genera like Ceropegia (Field & Collenette
1984). Hence in this account, no subspecies are recognised for C. cumingiana.

‘C. cumingiana was placed in a Section Hylopegia H. Huber together with C.
lucida Wallich from China, Thailand, Malaysia, Bangladesh, Burma and north-east India
(Huber 1957). Both species have entirely glabrous vegetative parts, a feature shared with
C. thwaitesii Hook., C. elegans Wall. and C. madagascariensis Decne of Sect. Janthina
Huber (from which section they are excluded by the less prominent basal inflation of
the corolla-tube). All these species have very slender, glabrous stems, large, glabrous
leaves and a slender, glabrous peduncle bearing many flowers. All of these taxa have
essentially the same coronal structure: the outer lobes are bifid into erect, pilose teeth
(long, slender lobules - C. madagascariensis) and the inner lobes, usually somewhat
clavate towards their tips, are connate in the centre above the stigmatic head. This
corona structure is shared with a wider group of species including the complex surrounding
C. longifolia Wall.

Huber (1957) separated C. cumingiana and C. lucida on the different shapes of
the corolla-lobes (strongly attenuated, often spathulate at apices in the latter; ovate,
oblong or obovate in the former). Unfortunately spirit or live material of C. lucida is
lacking for a more detailed comparison of the respective coronas. The outer corona of
C. cumingiana forms a rather deeper cup than is customary in any of the above-
mentioned taxa except for C. longifolia where both a shallow outer corona and a very
much deeper structure even than that in C. cumingiana sometimes occurs in subsp.
sinensis Huber.

Conservation status: C. cumingiana is widely distributed and the relatively few collections
present in herbaria are more a reflection of inadequate collecting than of rarity. The
species is not rare, endangered or vulnerable at present.

Bruyns & Forster, Ceropegia cumingiana {1

Ethnobotanical use: Bailey (1913) records C. cumingiana as being used as a yam by the
natives along the Batavia [Wenlock] River, Queensland. They called the plant “‘Anareata”’.

Acknowledgements

Field assistance was provided by D.J. & ILM. Liddle, P.D. Bostock, M.C. Tucker
and R.A. Harvey. B. Gray (QRS) and G. Sankowsky provided material and information
on localities. The Directors of B, BM, BRI, CANB, K, L, LAE, MEL and QRS either
provided information on holdings, or allowed access to collections. J. West (CANB)
while Australian Botanical Liasion Officer at Kew located types at BM, G, K and P. L.
Pedley (BRI) organised some loans. U. Eggli (ZSS) checked Z for type material. P.R.
Sharpe translated Schlechter’s 1914 paper dealing with C. papuana. The Australian
Biological Resources Study provided partial financial assistance during 1988. All are
thanked for their support.

References
ANSARI, M.-Y. (1984). Asclepiadaceae: Genus Ceropegia. Fascicles of Flora of India. 16: 1~34.
BAILEY, F.M. (1913). Comprehensive Catalogue of Queensland Plants. Brisbane: Government Printer.

BROWN, N.E. (1901). Decades Kewenses. Plantarum Novarum in Herbario Horti Regii Conservatarum. Bulletin
of Miscellaneous Information 1901: 138-145.

DECAISNE, J. (1844). Asclepiadeae. In A. De Candoile, Prodromus Systematis Naturalis Regni Vegetabilis 490-
664. Parisiis: Sumptibus Fortin, Masson & Sociorum.

DYER, R.A. (1980). Ceropegia. In Leistner, Flora of southern Africa 27(4). Pretorta: Goverment Printer.
DYER, R.A. (1983). Ceropegia, Brachystelma and Riocreuxia in southern Africa. Rotterdam: Balkema.

FIELD, D.V. & COLLENETTE, LS. (1984). Ceropegia superba (Asciepiadaceae), a new species from Arabia. Kew
Bulletin 39: 639-642.

HUBER, H. (1957). Revision der Gattung Ceropegia. Memorias da Sociedade Broteriana 12.

MERRILL, E.D. (1915). Genera and species erroneously credited to the Philippine Flora. The Philippine Journal
of Science, C. Botany. 10: 171-194,

Accepted for publication 10 March 1989

Austrobaileya 3(i): 13-38 (1989) 13

THE GENUS ZORNIA J. GMELIN (LEGUMINOSAE)
IN AUSTRALIA

S.T. Reynolds and A.E. Holland
Queensland Herbarium, Meiers Road, Indooroopilly, Qld 4068

Summary

The genus Zornia J. Gmelin is represented by 17 species including 4 subspecies and 4 varieties in Australia. The
identity of two additional taxa is uncertain. All species are described and notes and maps of their distribution
appended. A key to distinguish them is provided. Eight new species viz 2. acuta, Z. maritima, Z. disticha, Z.
floribunda, Z. oligantha, Z. pedunculata, Z. prostrata and Z. ramosa, two new subspecies Z. muelleriana subsp.
congesta and Z. muriculata subsp. angustata, and one new variety Z. prostrata var. macrantha are described.

The new combination Z. dyctiocarpa var. filifolia, based on Z. filifolia Domin is made. Lectotypes are chosen for
Z. chaetophora F. Muell., Z. muelleriana Mohl., Z. stirlingiti Domin and Z. filifolia var, adenophora Domin. Z.
nervata Mohl. is a synonym of Z. albiflora Mohl. The majority of plants previously identified as Z. nervata are
included in Z. prostrata. Z. gibbosa Span. apparently does not occur in Australia; Australian collections formerly
identified as such are included in Z. disticha.

Introduction

There has been much confusion in the genus Zornia in Australia, chiefly with
plants in Queensland, and Mohlenbrock’s monograph of the genus (Mohlenbrock 1961)
has been found to be generally unsatisfactory and in some cases adds to the confusion.
His descriptions of some species do not match the specimens he cited, or the type of a
species name was from a different taxon from that described and specimens cited for it.
In some cases, e.g. Z. gibbosa Span. he chose a lectotype when the type was still available
for study, and his description does not cover that type. In the case of Z. albiflora Mohl.
and Z. nervata Mohl., the types of the names were found to be from the same taxon,
and his description of the latter species and the specimens cited do not agree with the
nominated type.

There was also confusion in the Z. filifolia and Z. adenophora group. Mohlenbrock’s
concept of Z. filifolia is different from Domin’s Z. filifolia and represents a new species
(named Z. ramosa here). Confusion existed because Mohlenbrock did not cite types for
Z. filifolla Domin and Z. adenophora (Domin) Mohl. and synonyms under it. He
combined Domin’s Z. filifolia var. subeglandulosa (= Z. filifolia var. filifolia) and Z.
filifolia var. adenophora under Z. adenophora when they represent different species of
plants. He probably did not see Domin’s types, because under Z. muriculata he did not
cite all its syntypes or give locality details of the types for Z diphylla var. hirsuta Domin
or Z. diphylla var. xerophila Domin.

The species vary greatly in the size and shape of leaflets, stipules and bracts, in
occurrence of pellucid dots, in number of articles per fruit and in the presence or absence
of bristles on the fruit. Some of these characters vary in the one population or within
the one individual but the species nevertheless remains distinctive. Many species require
further collections to ascertain their full range of variation.

Note: The revision of Zornia is part of the Revision of the tribe Aeschynomeneae (Benth.)
Hutch. in Australia, funded by the Australian Biological Resources Study. A revision of
the remainder of the tribe will be published 1n a future issue of Austrobaileya.

Taxonomy
Zornia J. Gmelin, Syst. Nat. 2: 1076, 1096 (1792). Type: Z. bracteata J. Gmelin

Derivation of name: After Johann Zorn, a German botanist and senator who wrote on
rare plants and published illustrations of useful plants.

Erect or prostrate herbs usually with pellucid or pustular glands on stems, leaflets, stipules
and bracts. Leaves alternate, petiolate, 2—4-foliolate (bifoliolate in Australia); leaflets

14 Austrobaileya 3(1): 1989

digitate; stipules well developed, paired, peltate, usually with a prominent basal spur,
prominently nerved; stipels absent. Inflorescences spicate, terminal or axillary, rarely 1|-
flowered: bracts separated or overlapping, paired, stipule-like, peltate, enclosing flowers
when in bud, prominently nerved. Flowers sessile. Calyx hyaline, ciliate, persistent,
usually with a short tube and 5-lobed; tube + campanulate, prominently nerved, upper
pair of lobes joined; lateral lobes smaller than other three. Corolla mostly yellow often
streaked with orange, purple or red; petals all clawed; standard + rounded; wings
subobovate, short, transversely plicate-rugose; keel incurved. Stamens monadelphous;
upper part of tube circumciss at maturity and usually shed with petals; anthers unequal,
alternately small and large. Ovary subsessile, with (2~)4-9 ovules. Fruits sessile, 2-9-
jointed, separating into 1-seeded, indehiscent articles, or sometimes of only | article;
included within persistent bracts or exserted; articles subsimilar, mostly quadrate, usually
+ compressed, reticulately veined and often with bristles or glands; bristles usually with
short fine hairs; seeds mostly + oblongoid, pale or dark brown.

About 86 species widespread throughout the tropics and warm temperate areas;
19 species in Australia, eight new.

In some cases in the following key to species, 1t has been found necessary to use
a combination of characters to distinguish the species because of their variability and
because some characters, e.g. habit, are poorly known.

Note: Measurements of stipules and bracts include the basal spur, and the peduncles are
measured to the base of the spike.

1. Flowers solitary . BS ahs 2
Flowers 2-several in a "spike. ‘rarely solitary and then i in “combination
with spikes on the same plant ...... 2... ee ee ee 3

2. Leaflets linear, 0.5-1 mm wide. Fruits with insti bristly articles. Pedun-

cles 1.2- 2.5 cm long ...... Il. Z. ramosa
Leaflets ovate, 3-5 mm ‘wide Prnits siatirous, “articles with a few short

bristles or non bristly. Peduncles 3-4 cm long . Wed) 12. Z. pedunculata

3. Bristles on fruits 3~5 mm long. Articles 3-5 mm long...... 1. Z. chaetophora

Bristles less than 2 mm long. Articles less than 3.5 mm long .......... 4

4. Plants with conspicuous pellucid dots on leaflets; bracts, stems and

stipules usually dotted ........ er 5
Plants epunctate or rarely with few obscure pellucid dots - on ‘Jeaflets
Gowell SUTTACES WANG DPACtS is) os. sy ele ey ele ee ie oe ee, sree ch eet 14

5. Bracts (13-)16-24 X (4-)6.5-8 mm, densely pellucid-dotted, overlapping.

Leaflets (2.5-)4-6.5 cm long .. . . 2. Z. stirlingii
Bracts 4-14 X 0.5-6 mm, usually sparsely pellucid-d dotted, } overlapping
or separated. Leaflets 0. 8-3.8 cm long... ean 6
6. Leaflets usually pellucid-dotted on both surfaces. Stems, stipules and
bracts also with pellucid dots...... oes 7
Leaflets only pellucid-dotted on lower surfaces. Stems, stipules and bracts
occasionally with small obscure pellucid dots ...............0..0... 11
7. Stemserect ...... ees Aa Pet cee ae 08s een BPO A mh, 8
Stems. prostrate or eeniiprostidite: ey een ae se ey eee ae ee 9

8. Leaflets 0.5-1 mm wide, usually linear, concave or flat. Fruits with 2-
6, glabrous, conspicuously gland-dotted, non bristly articles 3. Z. adenophora
Leaflets 1~2.5 mm wide, narrowly elliptic, flat. Fruits with 2-4, nti
hairy, eglandular bristly articles .... . ? .... & sp. (1)

Reynolds & Holland, Zornia 15

9.

10.

Ll.

12.

1 4.

LS.

16.

Leaflets (18-)32-38 x 1.5-3 mm, 10-15 times as long as wide, drying
black. Bracts ovate, 9-13.5 X 4-7 mm, overlapping or more Or less

SePAPALCHS oa. Bie cee wre ik Se ne Belo eee Ae PEA Teo 4. Z. acuta
Leaflets 8-33 X (1-)2-13 mm, usually less than 10 times as long as

wide, not drying black. Bracts subobovate, elliptic or ovate, 4-11 xX

1.5-5 mm, mostly separated .. .. 2. 6. ee ee 10

Bracts 4-7 X 1.5-4 mm, subobovate to elliptic, usually very separated
(internodes to 3 times as long as bracts). Leaflets 1-5.5 mm wide,
narrowly elliptic to linear oblong. Articles very bristly, glabrous or
Anely Wanye. ceca eh ee te re a SE Be ea Se Aan ece oe 5. Z. areolata

Bracts 7-14 X 2-7 mm, ovate or elliptic, overlapping half their length
to separated. Leaflets 4-13 mm wide, ovate-oblong, elliptic to oblong.

Articles usually with few bristles (mostly peripheral), or non bristly,
glabrous or hairy .. —. 2. 56 ee ee ce ee ee ee ee ee oe 6. Z. muelleriana

Pellucid dots on leaflets and stems black. Inflorescences with distichous

overlapping, narrowly elliptic bracts. Stems ascending ...... 7. Z, disticha
Pellucid dots on leaflets and stems not black. Inflorescences with over-
lapping or separated, ovate or elliptic bracts. Stems usually prostrate .. .. [2

Bracts to 2 mm wide, narrowly elliptic with a small spur at base, pellucid-
dotted, prominently nerved. Fruits with 4 or 5, pale articles with
reddish purple reticulation and bristles .... ... a . ££. sp. (2)

Bracts 3-6 mm wide, ovate, usually with a conspicuous spur; pellucid
dots sparse or absent; nerves prominent or indistinct. Fruits with 2-
6, pale articles with pale reticulation .. .. ©) 6.) ee ee ee re {3

. Leaflets 6-14 mm wide, ciliate, usually shiny and prominently reticulate

helow. Petioles 0.8-5.7 cm long. Bracts overlapping half their length
to. separated.6 ee ee Se Ka ee a ee te i ee be 8. Z. maritima
Leaflets 4-6.5 mm wide, neither ciliate nor shiny, mostly drying
pale, with reticulate venation inconspicuous. Petioles 1.5-2 cm )

long. Bracts overlapping to about a quarter their length.. .. .. 13. Z. pallida
Stems erect ok sc ke wre eh ee ee fee hp OEM Hees Eee Pe ee me re Hah A 15
Stefiis prostrate. 20.05. c.n eG eee yore ie Os Gee ey er re a PE OP OR HE ot 17

Bracts 4-8 mm wide, broadly ovate, obtuse, prominently reticulate.
Fruits hairy with hairy long bristles. Plants hairy ........ .. 15. Z. albiflora

Bracts 1-5 mm wide, narrowly elliptic to ovate, acute or acuminate,
obscurely reticulate. Fruits glabrous or hairy; bristles long or short.

Plants glabrous or hairy .....-...-..- 16

Leaflets 26-40 X 1-2.5 mm (9-30 times as long as wide). Inflorescences
{4-22 cm long, on peduncles 2.7-8 cm long. Bracts overlapping or
nearly so, acute. Fruits 3-6-articulate, usually hairy with few usually
peripheral bristles.. .. 2... 6. 6. ee ee ee te ee 10. Z. floribunda
Leaflets (5-)7—-32 x 0.5-9 mm (2-10 times as long as wide). Inflorescences
2.5-9.5(-15) cm long, on peduncles 0.3-1.5(-2.8) cm long. Bracts
overlapping or widely separated, acuminate or acute. Fruits
|-4(-6)-articulate, usually glabrous, muriculate or covered with long
bristles eet Mee ree ee el ae, cles Etat tele gs Eases eee ns 9, Z. muriculata

- Leaflets and bracts less than 2.5 mm wide .. .. 2-2. 6) ee ee ee ee ee es 18

Leaflets and bracts more than 3 mm wide... .. .... 2 2. ee ee ee ee ee 19

Sha

16 Austrobaileya 3(1): 1989

18. Inflorescences with 1-4(rarely-10) very separated bracts. Fruits hairy
with few small bristles or without bristles. Stipules not elongate,
+ falcate.. .. 2... ee ee ee ee ee ee ee ee we ws... «614, Z. dyctiocarpa

Inflorescences with (4-)7-20 very overlapping or + _ separated
bracts. Fruits usually glabrous, with blunt or acute, small,
short or long bristles. Stipules mostly elongate, + _ falcate,
CONSPICUOUS ht 2 estes CON es Bee yee fit lo wo oe 9. Z. muriculata

19. Leaflets less than 6 mm wide, lower ones usualty smaller and broader
than upper ones... ... eng eae, 20
Leaflets 6.5-15 mm wide, gees ones okie eftiatiiee ani upper ones...... 22

20. Fruits (4—)6—-9-articulate, glabrous rarely hairy; bristles bias absent.

Bracts separated, ovate ...... ... 14. Z. dyctiocarpa
Fruits 1—5-articulate, glabrous or hairy: bristles present or absent. Bracts
overlapping, especially towards the apex, ovate or elliptic ............ 21

21. Leaflets pale; nerves not apparent. Bracts 5-6 mm wide, acute.
Fruits with hairy articles and many hairy bristles; articles 2-3 xX
3-—3.5 mm. Stipules 8.5-10 cm long, not elongate or + falcate 13. Z. pallida

Leaflets not pale, usually with prominent lateral nerves. Bracts 2—3(—4)
mm wide, acuminate. Fruits usually glabrous; bristles few or absent;
articles 2-3 X 1.5-2.5 mm. phimamiats aes mm sais Ean elongate
+ falcate...... ) 9. Z. muriculata

22. Fruits hairy; articles 2-3.5 mm long, eglandular, prominently reticulate
veined and with conspicuous, hairy bristles. Inflorescences (4—)8-21-
flowered; bracts prominently reticulate. Leaflets usually bec ovate,
elliptic or obovate, obtuse... .. .. 16. Z. prostrata

Fruits glabrous; articles 1.5-2 mm lite. glanchilar-dotied. obscurely
reticulate veined, with small bristles, mostly at periphery, or bristles
absent. Inflorescences 2-8(- ied flowered: bracts obscurely reticulate.
Leaflets -ovate, acute os 25 2.2 a5 4 tig k yee Ad Sele ae do 17. Z. oligantha

1. Zornia chaetophora F. Muell., Trans. & Proc. Philos. Inst. Vic. 3: 56 (1858). Type:
Western Australia. Sturt Creek, February 1856, F. Mueller (lecto (here designated):
K; 180: MEL(MEL 1544961)).

Z. diphylla var. conjugata subvar. perglandulosa Domin, Biblioth. Bot. 89: 209
(1926). Type: Northern Territory. Victoria River, F. Mueller (holo: n.v., 180:
MEL(MEL 1544961)).

[Z. diphylla var. gracilis auct. non (DC.) Martius: Bentham, Fl. austral. 2: 229
(1864), quoad specimine Sturt Creek, F. Mueller.}

Herbs to 75 cm high, with erect, much-branched, glabrous or subglabrous stems. Stems,
leaves, stipules and bracts with dense reddish or pale brown ellipsoid or globose glands.
Petioles 4-13 mm long; leaflets variable, ovate, elliptic, linear oblong or narrowly elliptic,
1.5-4 cm long, 1-6(-—10) mm wide, lower ones usually smaller and broader than upper
ones, apex acute or mucronate, base obtuse or acute, margins rarely ciliate; midrib
prominent; petiolules to | mm long. Stipules subrhombic, usually subfalcate, 6-13 x 1-
5 mm, attenuate or acuminate at apex, produced into a narrow spur at base, 5- 7-nerved;
reticulate venation obscure; spur acuminate, 2-6 mm long. Inflorescences 7-25 cm long,
3-9-flowered, with very widely separated bracts; peduncles to 10 cm long, punctate;
bracts narrowly ovate or elliptic, 6-10 * 1.5-—3 mm, acute, with broad or narrow spur
at base, glabrous or puberulent, ciliate, 5-nerved; reticulate venation obscure; spur 1-2.5
mm long, acute, truncate or obtuse. Calyx 3-4 mm long; petals yellow; standard 6-8
mm long. Fruits 1—5-articulate, exserted; articles broadly elliptic or + obovate, concave
above, 3-S(-7) * 2-2.5 mm, prominently reticulate veined, eglandular, glabrous. ciliate,
or puberulent, covered with long, filiform, patent-hairy bristles: bristles 3-5 mm long.

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Reynolds & Holland, Zornia 17

Selected specimens: Western Australia. Near Derby Airport, 8 km S of Derby, Apr 1985, Aplin et al. 32 (PERTH);
Derby Pindan Block, Aug 1973, Holm 27 (PERTH); Junction of Gupungi and Cable Beach roads, Broome, Aug
1985, Kenneally 9431 (CANB,PERTH); Great Sandy Desert, Bruces Field, May 1982, Harris 48 (AD); Sturt
Creek, Feb 1856, Mueller [MEL 1544961] (K,MEL). Northern Territory. Victoria River, Mueller [MEL 1544961]
(MEL); SSE of mouth of Daly River, Jul 1946, Blake 16663 (BRI,CANB,NT); Creek Crossing, 3 miles [4.8 km]
S of Litchfield homestead, May 1964, Robinson R362 (NT); 5 miles [8 km] W of Wangi homestead, Aug 1969,
Byrnes 1674 (CANB,NT); Fish River Stn, Aug 1983, Silversteen 911 (DNA); Port Keats, Sep 1972, Robinson
[NT37493] (DNA,NT); 30 miles [48 km] S of Borroloola, Mar 1972, Byrnes 2541 * (CANB,DNA); 5 miles [8
km] S of Elhott, Stuart Hwy, Feb 1969, Must 462 (BRI,CANB,NT); 45km N of Renner Springs, Jul 1974, Carr
46357 & Beauglehole 46379 (NT); Murchison Range area, Apr 1983, Latz 9699 (NT); 24 km S of Soudan
homestead on Annitowa Rd, May 1977, Henshall 1791 (AD,BRI,CANB,NT). [* Byrnes 2541 has shorter bristles
(1.5-2 mm long) on the loments than is usual for the species but is otherwise typical. ]

Distribution and habitat: This species occurs in north Western Australia (from the
Kimberleys to the Great Sandy Desert) and in Northern Territory (around Fish River
and Victoria River, and as far south as the Murchison Range) (Map 8); occurs mainly
in sandy or loamy soil on red sandstone piains, in shrubland, eucalypt woodlands or
disturbed areas.

This species differs from all the other species occurring 1n Australia in the large
articles (3-7 X 2-2.5 mm) and long (3-5 mm long) patent-hairy bristles. Leaflets are
also long (to 4 cm) and densely pellucid-dotted. 3

Typification: The Kew sheet is chosen as a lectotype because it 1s part of the original
material and is a better specimen than the ones at MEL. Moreover the MEL sheet (MEL
1544961) consist of three specimens, all of this species, and carries two labels with
aewtaie localities viz Sturt Creek, Feb 1856, F. Mueller, and Victoria River, Jan 1856,
F. Mueller.

2. Zornia stirlingii Domin, Biblioth. Bot. 89: 210, t. 25, f. 1-4 (1926). Type: Queensland.
Cook DISTRICT: in xerodrymio ad pedem colles, calcifer Bluff prope opp Chillagoe,
ceeon 1910, Domin (lecto (here designated): PR(PR 527338); isolecto: PR(PR
527339)).

Z. diphylla var. stirlingii Bailey, Queensland Agric. J. 15: 491 (1904). Type: Queens-
land. COOK DISTRICT: Herberton, June 1904, J. Stirling (holo: BRI).

Tall, robust or spindly, glabrous herbs to 1 m high with thick, erect, few-branched stems.
Leaflets, stipules and bracts with reddish brown or pale brown pellucid dots, dense on
leaflets, stipules and bracts, fewer on stems, petioles and peduncles. Petioles 0.5-—2.2 cm
long; leaflets narrowly elliptic-ovate or linear, (2.5—)4—6.5 cm long, 1.5-4(-5.5) mm wide,
acute or acuminate at apex, acute at base; petiolules 0.5—0.7 mm long. Stipules narrowly
elliptic or subrhombic, 10-20 X 1.5—3 mm, acuminate at apex, with a long narrow spur
at base, 5-nerved; reticulate venation obscure; spur 5—7 mm long, acuminate. Infloresc-
ences 11-21 cm long, densely flowered with bracts crowded and overlapping especially
towards apex; peduncles to 10 cm long; bracts broadly elliptic or ovate, (1.3-)1.6-
2.4 X 0.4-0.8 cm, acute at apex, with a short broad spur at base, ciliate, 5—7-nerved;
reticulate venation prominent; spur acute or obtuse, 1-5 mm long. Calyx 4-5 mm long;
petals yellow; standard 7-11 mm long. Fruit 2—7-articulate, exserted or included; articles
2.5-3 X 2-3 mm, with reddish brown reticulate venation, eglandular, puberulent, densely
Scalia with fine, retrorse hairy bristles especially at the periphery; bristles 0.5-2 mm
ong.

Selected specimens: Queensland. COOK DISTRICT: 7 km from Kennedy River crossing on the Fairview—Kimba
Rd. on track to King River Outstation, Apr 1980, Clarkson 3232 (BRI); Mt Molloy, Apr 1932, Brass 2470 (BRI);
Chillagoe, Feb 1910, Domin (PR); 7 km W of Rockwood Ck on the Burke Development Rd, Mar 1980, Clarkson
3002 (BRI); Stannary Hills, May 1909, Bancroft 514 (BRI); Parada, 20 wb angel te km] SW of Mareeba, Apr 1962,
McKee 9358 (BRI); Herberton, Jun 1904, Stirling (BRI); southern slopes of Mt Masterton, Jan 1960, Gocdall
[AQ239526] (BRI); Springmount Stn, ca 13 km from the Mareeba—Dimbulah Rd, on the road to Collins Weir,
Apr 1983, Clarkson 4630 (BRI); Montalbion on Petford—Herberton Rd, Apr 1962, McKee 9429 (BRI); Lappa,
Jan 1982, Pedley 4840 (BRI,CANB); Bullock Ck, Feb 1922, White 1350 (BRI); Etheridge River, without date,
Armit 913 (MEL). NORTH KENNEDY DistTRICT: Christmas Creek Stn, 120 miles [192 km] NW of Charters Towers,
elle Compton 89 (BRI); Mt Saunders about 1.5 miles [2.4 km] NW of Yabula, Aug 1942, Smith T.118

Distribution and habitat: So far known only from north eastern Queensland (Map 8);
occurs mostly on rock or grassy hillslopes in eucalypt woodlands usually near creeks
and drainage lines.

The large (1.3-2.4 cm long) overlapping bracts and the densely pellucid-dotted
leaflets, stipules and bracts distinguish this species from other Australian species.

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12 Austrobaileya 3(1): 1989

Typification: This plant was described as Z. diphylla var. stirlingii by F.M. Bailey in
1904. K. Domin specifically described it as a new species in 1926, included Bailey’s Z.
diphylla var. stirlingii in the synonymy and took up the epithet “stirlingii” for it. He
was not, however, making a new combination of Bailey’s name. Domin’s description
was based principally on his own collections, though he cited also the type of Bailey’s
name with his protologue. He indicated, however, that he had not seen it. His protologue
also included the illustration in Bailey’s Comprehensive Catalogue (Bailey 1913). PR
527338 and PR 527339 from Domin’s original collections are chosen as lectotype and
isolectotype as the rest of the specimens are inadequate.

3. Zornia adenophora (Domin) Mohl., Webbia 16: 50, f. 33 (1961); Z. Alifolia var.
adenophora Domin, Biblioth. Bot. 89: 211 (1926). Type: Queensland. Nortu
KENNEDY DISTRICT: in monte Mt Remarkable prope opp Pentland, February
1910, Dormin (lecto (here designated): PR(PR 527344)).

Z. diphylla var. filifolia Bailey, Bot. Bull. 3: 10 (1891). Type: Queensland. Cook
DistRIcT: Walsh River, March 1891, 7. Barclay Millar (holo: BRI).

Glabrous herbs to 50 cm high, with erect much-branched, sparsely pellucid-dotted stems.
Leaflets, stipules, bracts and fruits usually with dense, ellipsoid, reddish or brown glands.
Petioles 0.4-1.3 cm long; leaflets linear-oblong, 1-3.4 cm long, 0.5~1 mm wide, acute or
mucronate at apex, acute at base, flat or concave; midrib prominent: petiolules to 0.5
mm long. Stipules narrowly ovate, elliptic or subrhombic, 2~3.5 & 0.5-1 mm, acute or
acuminate at apex, usually with a short spur at base, 3-nerved: reticulate venation
obscure; spur 0.5-1.5 mm long, acute. Inflorescences 3.5-10.5 cm long, 5-13-flowered,
with usually very widely separated bracts (internodes 2~3 times as long as bracts):
peduncles glabrous, to 3 cm long; bracts narrowly elliptic, often falcate, 3-5 x 0.5-1.5
mm, acute at apex, shortly spurred at base, rarely ciliate, 3-S-nerved; nerves pale:
reticulate venation obscure; spur 0.5-1 mm long, acute or toothed. Calyx 2-3.5 mm
long; petals yellow-orange; standard 5-6 mm long. Fruit (2-)4-6-articulate, exserted:
articles 1.5-2 x 1-2 mm, glabrous, usually prominently reticulate veined, covered with
large glands; bristles absent.

Selected specimens: Northern Territory. Calvert River, Jun 1987, Thompson 1867 (NT). Queensland. BURKE
District: Almaden, Jun 1907, Blackman [AQ239371] (BRI); Westmoreland Stn, 145 km WNW of Burketown,
May 1974, Pullen 9193 (CANB); 23 km NW of Old Corinda Outstation along Doomadgee to Westmoreland
road, May 1974, Pullen 9115 (CANB); ca 9 km SW of Normanton, on road to Cloncurry, Apr 1973, Henderson
Hi807 (BRI); 100 km from Georgetown on Normanton Rd., Apr 1975, Craven 3316 (BRI,CANB,MEL): 64 km
from Georgetown on Croydon Rd, Mar 1988, Forster 3840 (BRI); Cook District: Tate River, undated, Armif
537 (MEL); 3 miles [4.8 km] E of Mareeba, May 1959, Thorne 21136 & Jones (BRI); Einasleigh River, undated,
Armut 916 (MEL). NorTH KENNEDY DiIsTRICT: near source of Poison Ck, about 90 miles [144 km] N of
Hughenden, Apr 1935, Blake 8556 (BRI,CANB); Warrigal, on Great Dividing Range, Feb 1931, Hubbard &
Winders 7101 (BRI); Mt Remarkable, Pentland, Feb 1910, Domin (PR); 36 km E of Torrens Ck, Charters Towers
Rd, Jun 1984, Hacker 693 (BRI). SOUTH KENNEDY DISTRICT: near junction of Campaspe and Cape Rivers, Apr
1945, Blake 15739 & Webb (BRI,CANB). :

Distribution and habitat: Gulf of Carpentaria and northern Queensland extending south
to Pentland (Map 7); occurs in sandy soils in open forests and grassland, on flood plains
and in disturbed areas.

The glandular articles distinguish this species from all other Australian species. It
is allied to Z. areolata and Z. dyctiocarpa var. filifolia, it differs from the former in its
erect habit and more narrow leaves and bracts, and from the latter in its densely glandular
fohage and glandular fruits.

Typification: Prague sheet PR 527334 is chosen as the lectotype as it is the only sheet
of the original material that agrees with the original description. See also discussion
under Z. ramosa below.

4. Z. acuta S. Reyn. & Holland species nova a speciebus Australianis ceteris foliolis
elongatis angustatis, bracteis latis ovatis, stipulis elongatis, acuminatis differt.
Typus: Northern Territory. Shoal Bay Road Darwin, 2 February 1961, H.S. McKee
8377 (holo: BRI; iso: NT).

Herbs with + prostrate, glabrous, obscurely and sparsely pellucid-dotted stems. Petioles
(0.8~)1.3-1.5 cm long; leaflets narrowly elliptic, (1.8-)3.2-3.8 cm long, 2-3 mm wide,
acute to acuminate and mucronate at apex, subacute at base, glabrous, pellucid-dotted:

Reynolds & Holland, Zornia 19

petiolules to 1 mm long. Stipules narrowly elliptic, (8-)12-17 x 1.5-3 mm, acuminate
at apex, prominently spurred at base, 3-5-nerved, glabrous, pellucid-dotted, reticulate
venation prominent; spur 2-4.5 mm long, acute, acuminate, obtuse or truncate. Inflo-
rescences 4-11 cm long, 3-8-flowered with overlapping or + separated bracts (internodes
to 1.5 times as long as bracts); peduncles 2-4 cm long; bracts broadly ovate, 9-13.5 Xx
4.7 mm, acute at apex, S-nerved, glabrous, conspicuously reticulate veined, sparsely
pellucid-dotted. Flowers not seen. Fruits (immature) 1-5-articulate, included; articles
elabrous or hairy, gland-dotted; bristles to 1 mm long, hairy or glabrous.

Selected Specimens: Western Australia. Gibb River, Jan 1951, Gardner 9942 (PERTH). Northern Territory. Shoal
Bay Rd, Darwin, Feb 1961, A¢fcKee 8377 (BRILNT).

Distribution and habitat: North-western Australia (Map 13): known only from the above
collections. |

Z. acuta differs from all the Australian species with elongate narrow leaflets, in
the broad ovate bracts, and elongate acuminate stipules. It 1s closely allied to Z. areolata
differing chiefly in its long narrow leailets.

Etymology: The specific epithet refers to the acute leaflets and bracts.

5. Zornia areolata Mohl., Webbia 16: 53, f. 35 & 36 (1961) Type: Queensland. BURKE
District: Creen Creek, Gulliver s.n. (holo: MEL, n.v.; iso: MELCMEL 1544960)).

Herbs with prostrate, spreading, much-branched, reddish, sparsely hairy to glabrous stems
to 1 m long. Leaflets, stipules and bracts with conspicuous reddish or dark glands.
Petioles 4-18 mm long, puberulent or glabrous; leaflets narrowly oblong or elliptic to
linear, 0.8-2.5 cm long, (1-)2~3.5(-5.5) mm wide, acute or obtuse and mucronate at
apex, obtuse at base, glabrous or lower surface subglabrous; petiolules to 0.5 mm long.
Stipules narrowly ovate or subrhombic, 2-7 xX 1-2 mm, acuminate at apex, shortly
spurred at base, 3—5-nerved; reticulate venation obscure; margins pale; spur to 1.5 mm
long, acute, obtuse or toothed. Inflorescences 2-5.5 cm long, (1-)2—6-flowered, with
separated bracts (internodes to 3 times as long as bracts); peduncles filiform, 1-4.5 cm
long: bracts broadly subobovate or elliptic, 4-7 x 1.5-4 mm, obtuse or subacute at apex,
suboblique, with narrow spur at base, ciliate, 5-nerved; nerves pale, with fine distinctive
reticulate venation; spur 0.5-1 mm long, acute, obtuse or toothed. Calyx 2-3 mm long;
petals orange-yellow; standard 6-8 mm long. Fruit 2-5-articulate, exerted, crenate on
lower margins; articles + obovate, concave above, or oblong, 2-3 X 1.5-2 mm, con-
spicuously reticulate veined, eglandular, puberulent or subglabrous, covered with fine
glabrous bristles; bristles to 1 mm long.

Selected Specimens: Northern Territory. Near Knucky Lagoon, Nickson Place, Apr 1956, Burbidge 5106 (BRI);
Batchelor, Mar 1963, Muspratt SSO100 (NT); Yam Ck, near Brocks Ck 130 km SSE Darwin, Jan 1883, Foelsche
41 (AD,K,MEL); Pine Ck, Apr 1904, Niemann (MEL); 3 miles [4.8 km] N of Hays Ck, Apr 1969, Byrnes 1535
(BRI); Near Douglas River, Apr 1967, Robinson CSR16 (NT); 1/2 mile [ca 1 km] NW of Edith River Crossing,
Jan 1965, Wilson 210 (NT); Katherine Gorge National Park, Apr 1977, Dunlop 4493 (NT); Queensland: Cook
District: Newcastle Bay, May 1948, Brass 18678 (BRI); 0.8 km S of Laura River crossing on the Peninsula
Development Rd, Mar 1987, Clarkson 6800 & McDonald (BRI); Einasleigh-Forsayth Rd, Jul 1983, Hacker 359
(BRI). BuRKE District: Between Norman and Gilbert Rivers, in 1874, Gulliver 57 (MEL); Westmoreland Stn,
May 1976, Pullen 9193 (BRI). SourH KeNNepy District: Yarromere, 100 miles [160 km] SSW of Charters
Towers, Apr 1969, Walker [AQ239374] (BRI). LEICHHARDT District: Marylands stn, head of Connor’s River,
Apr 1962, Hinton [AQ239431} (BRI).

Distribution and habitat: Northern Northern Territory (Darwin to Katherine Gorge) to
northern and central Queensland (Map 9); occurs in sandy or sravelly soils in open
forests, creek banks, red soil flats, slopes of granite outcrops.

Z. areolata is usually a slender-looking plant with prostrate slender stems, narrow
pellucid-dotted leaflets and usually small, very separated subobovate or elliptic bracts.
The peduncles of some plants also tend to be zig-zag. The species varies in hairiness
and inflorescences, e.g. plants from Queensland tend to be more hairy with broader
leaflets and bracts while those from Katherine (N.T.) are + glabrous with small bracts
and very narrow leaflets.

Typification: We were not able to locate the holotype at MEL. A collection (at MEL),
from between Norman and Gilbert Rivers, in 1874 by Gulliver, (Gulliver 57), was

0 Austrobaileya 3(1): 1989

probably collected at Creen Creek and is probably an isotype. It agrees with the protologue
and illustation of the holotype. |

Note: Mohienbrock (1.c.) cites Z. diphylla var. conjugata subvar. perglandulosa Domin,
Biblioth. Bot. 89: 209 (1900) as a synonym of Z. areolata, but this plant from Upper
Victoria River, N.T., F. Mueller (photos BRI), is Z. muelleriana subsp. congesta, which
occurs in the area.

6. Zornia muelleriana Mohl., Webbia 16: 131, f. 89 (1981). Type: Queensland. Cook
District: Walsh, March 1891, 7. Barclay Millar (lecto (designated here):
BRI(AQ022928, excluding pieces of Z. muriculata indicated as such)).

Herbs with prostrate or semi-prostrate, laxly branched, densely hairy to glabrous, pellucid-
dotted stems 12.5-44 cm long. Stems, leaflets, stipules, bracts and petioles pellucid-
dotted with dense conspicuous yellow to darkish glands, rarely sparsely pellucid-dotted.
Petioles 8-16 mm long; leaflets ovate, elliptic, ovate-oblong or subobovate, (0.8-)1.4-
3.3 cm long, 4-13 mm wide, lower ones usually smaller and broader than upper ones:
apex acute or obtuse, rarely + truncate or retuse, base obtuse and oblique, both surfaces
conspicously pellucid-dotted, hairy or glabrous; lateral nerves usually conspicuous; petiol-
ules to 1 mm long, pilose. Stipules narrowly elliptic or rhombic, (3-)5-7 X 1.5-2 mm,
acuminate or acute at apex, usually narrowly spurred at base, 3-8-nerved, glabrous,
reticulate venation obscure; spur 1-2.5 mm long, obtuse, acuminate or truncate. Inflo-
rescences 3-25 cm long, (2—-)5-27-flowered with bracts overlapping (towards apex) to
separated, especially towards base (internodes 0.5-3 times as long as bracts); peduncles
1.5-10.5 cm long; bracts elliptic, ovate or nearly suborbicular, 7-14 x 2-7 mm, 3-6-
nerved, ciliate, prominently reticulate veined and gland-dotted: spur 1-2.5 mm long,
acute, obtuse or truncate. Calyx 3.5-4 mm long; petals yellow, pinkish or red: standard
6-7 mm long. Fruits (1-)2—5-articulate; articles 1.5~2.5 * 1.5-3 mm, usually pale with
conspicuous reddish brown reticulate venation, glabrous or sometimes finely hairy, ciliate,
sparsely gland-dotted, bristly (mostly at periphery) or rarely non bristly; bristles up to 2
mm long, usually hairy especially near the tip.

Z. muelleriana Mohl. is characterised by the usually densely pellucid-dotted leaflets,
stems, stipules and bracts; the ovate, elliptic or ovate-oblong and pellucid-dotted (on
both surfaces) leaflets; the inflorescences with usually overlapping to separated bracts,
and by the fruits with mostly whitish or pale coloured articles which have reddish
reticulate venation, few small glands and weak bristles. Some forms of this species
resemble some forms of Z. prostrata with obscure scattered pellucid dots but the petioles
are short in Z. muelleriana and long (longer than leaflets) in that species.

Typification: When Mohlenbrock described Z. muelleriana he did not indicate either in
the publication or on the specimen which of the two collections of Barclay Mullar
(December 1890 or March 1981) was the holotype. The above collection is selected as
lectotype because it is a better specimen and agrees with the original description. The
sheet 1{ is mounted on also includes small pieces of Z. muriculata which have been
indicated as such.

__ Leaflets, bracts, hairiness and number of glands in Z. muelleriana are all very
variable. Two subspecies are recognisable as follows.

Bracts elliptic or ovate, 7-11.5 * 2-5(~7)mm, overlapping by one-third their
length near apex, separated near base. Peduncles 1.5~-10.5 cm
long. Leaflets acute or obtuse at apex. Stems sparsely hairy to
SET CUE odo ec Bea ay os Re en tere Pu ae i Ln BES ends oe, subsp. muelleriana
Bracts broadly ovate to suborbicular, 8-14 X 3~7 mm, overlapping by about
half their length throughout. Peduncles 2-2.5 cm long. Leaflets obtuse,
retuse or + truncate at apex. Stems hairy ee subsp. congesta

#. muelleriana Mohl. subsp. muelleriana
Distinguishing characters are as set out in the key above.

Stems + densely gland-dotted, usually sparsely antrorse hairy. Leaflets mostly obiong-

Reynolds & Holland, Zornia 2]

ovate to elliptic or oblong, obtuse or acute at apex, glabrous or hairy, densely pellucid-
dotted (rarely with only a few dots); lateral nerves usually conspicuous.

Selected specimens: Queensland. Cook District: Walsh, Mar 1891, Barclay Millar (BRD; Mt Molloy, Apr 1962,
McKee 9122 (BRI); Southedge Stn, Yalkula, Feb 1978, Anning 269 (BRI); Gilbert River, Bick (BRI); Parada
Research Station, 32? km NSW of Mareeba, Mar 1975, Staples 2018 (BRI). NORTH KENNEDY District: 2 km N
of Fletcher Ck, about 40 km NW of Charters Towers on Lynd Rd, Jul 1983, Hacker 338 (BRD; Range View,
S.E. of Ravenswood, May 1954, Everist 5528 (BRD). SOUTH KENNEDY District: Collinsville, Mar 1984, Thompson
[AQ 440712] (BRI); Lou Lou Park Stn, about 50 km N of Jericho, Feb 1984, Cheffin 71 (BRD. PoRT CurTIs
District: Fitzroy River near Rockhampton, Jan 1961, Howard [AQ239427]} (BRI). LEICHHARDT DISTRICT: Burton
Downs, May 1982, Johnson 2338 (BRI).

Distribution and habitat: Z. muelleriana subsp. muelleriana is known only from Queens-
land (from near the Gilbert River to near the Fitzroy River) (Map 1); occurs usually in
sandy or rocky soil on sandy ridges or creek levees, 1n open eucalypt forests.

Subspecies muelleriana is very variable with at least three distinctive forms as
indicated below. They are not formally recognised here because of the presence of
intermediate forms and also because their distributional ranges overlap. Plants from
around Gilbert River are typical. They have densely gland-dotted stems, leaves, stipules
and bracts, and stems glabrous or nearly glabrous. Plants from the Atherton Tableland,
near Ravenswood and Suttor River are mostly with hairy stems, leaflets and bracts. This
form resembles subspecies congesta but differs from it in its separated smaller narrower
bracts. Some plants of this form are intermediate between this species and Z. prostrata
e.g. Herbert River, Dallachy (MEL); Mareeba, Pedley 2254 (BRI), and Blake 13476
(BRI); 85 km from Lynd, Einasleigh—Forsayth Rd, Hacker 351 (BRI) (Map 12). They
have similar leaflets to those of Z. prostrata with prominent nerves and also prominently
reticulate veined bracts. They are tentatively retained under Z. muelleriana subsp.
muelleriana because of the pellucid-punctate leaflets. Further collections are necessary
to ascertain relationship of these plants. Besides the above forms, a group of plants from
around Townsville to Fitzroy River area has fairly large, + showy inflorescences with
separated small bracts and hairy fruits, and resembles Z. disticha. However, in that
species the inflorescence has congested distichous bracts. Further collections of this taxon
may show it warrants recognition as a distinct variety or subspecies.

Zornia muelleriana subsp. congesta S. Reyn. & Holland subsp. nov. a subsp. muellerianae
bracteis congestis plerumque latioribus et pedunculis inflorescentiarum brevioribus
(2—2.5 cm longis) differt. Typus: Western Australia. Great Northern Highway,
200 km north of Hail’s Creek along road to Kununurra (16°45’S, 128°17’E), 25
April 1985, Aplin et al. 393 (holo: PERTH).

Distinguishing characters are as set out in the key above.

Stems, leaves, stipules and bracts densely hairy and pellucid-dotted with conspicuous
yellowish glands. Leaflets elliptic, broadly ovate-oblong or subobovate, obtuse, + truncate
or retuse at apex: lateral nerves conspicuous.

Selected specimens: Western Australia. Carr Boyd Ranges bordering Lake Argyle, Mar 1978, Hartley 14558
(CANB,NT,PERTH); Smoke Ck, SW of Lake Argyle, Apr 1980, Weston 12085 (PERTH); Kimberley, Meda-
Oobagooma Rd, 80 km by road N of Gibb River Rd, 70 km NE of Derby, Jun 1976, Beauglehole 52761 (PERTH),
Derby-Broome Rd, 19.6 km S of Derby, Apr 1985, Aplin et al. 63 (PERTH), Beagle Bay Mission, Dampier
Peninsula, Mar 1985, Mfartin 14 (PERTH); Head of Breadon Valley, South Esk Tableland, Apr 1979, George
15488 (NT,PERTH); Great Northern Highway, 200 km N of Hails Ck along Rd to Kununurra, Apr 1985, Aplin
et al. 393 (PERTH). Northern Territory: Victoria River, Dec 1855, Adueller (MEL); 130 km W of Timber Ck,
Victoria Highway, Sep 1974, Parker 476 (CANB,DNA,NT).

Distribution and habitat: From the Kimberleys, Western Australia to Victoria River area,
Northern Territory (Map 1); occurs usually in sandy soil, on sand plains, low lying areas
and also near creeks, usually amongst grasses and herbs.

Subspecies congesta is distinguished from the typical subspecies by the overlapping
bracts, short peduncles and densely hairy, gland-dotted stems, leaves, bracts and stipules.
The bracts vary greatly; some plants have wide bracts similar to those of Z. albiflora
Mohl. e.g. Royce 6892 (PERTH) from Mt Anderson Station, but Z. a/biflora differs in
the erect or ascending stems and the epunctate or rarely sparsely dotted bracts.

Etymology: The subspecific epithet refers to the congested bracts.

22 | Austrobaileya 3(1); 1989

7. Zornia disticha S. Reyn. & Holland species neva ad Z. muellerianam Mohl. foliolis
et caulibus glanduloso-punctatis accendens, autem caulibus plerumque erectis,
bracteis distichis congestibus et leguminibus puberulis differt. Typus: Northern
Territory. Night Cliff, Darwin, (12°22’S, 130°53’E), 24 March 1948, R.L. Specht
43 (holo: BRI; iso: AD,MEL).

[Z. gibbosa auct. non Span.: Mohl., Webbia 16: 112-113 (1961) quoad specimina
Australiensia et decriptionem pro parte. ]

Herbs to 30 cm high with laxly branched, usually ascending, glabrous or sparsely antrorse
hairy, and sparsely pellucid-dotted (with black glands) stems. Petioles (0.8-)1.4-2 cm
long, glabrous or subglabrous; leaflets narrowly elliptic or oblong, (1—-)1.4-2.5 cm long,
2.5-6.5 mm wide, subacute or obtuse and mucronate at apex, suboblique at base,
glabrous, usually with black or dark brown pellucid dots on lower surfaces and margins:
lateral nerves obscure; petiolules 0.5-1 mm long, pilose. Stipules narrowly elliptic or
rhombic, 6-9 X 1~2 mm, acuminate at apex, with narrow spur at base, 3-nerved, pellucid-
dotted; spur to 2 mm long. Inflorescences 2.5-13 cm long, 6-25-flowered, with flowers
densely clustered and with distichous overlapping bracts (bracts overlap half to one-
third their length); peduncles 5-14 mm long; bracts narrowly elliptic, 7.5-12 * 3-4 mm,
with small ovate spur at base, 5-nerved, pellucid-dotted (with yellow dots), glabrous
except for ciliate margins; spur 1-2 mm long. Calyx 2-3 mm long; petals yellow; standard
4-5 mm long. Fruits (2- or)3- or 4-articulate, included; articles 2 = 1.5 mm, hairy,
usually with prominent dark reticulate venation and hairy bristles, sparsely gland-dotted;
bristles 0.5-1 mm long, retrorse-hairy. -

Specimens examined: Papua New Guinea, exact locality unknown, Brass 1186 (BRI). Northern Territory. Night
Cliff, Darwin, Mar 1948, Specht 43 (AD,BRI,MEL); East Point Reserve, Darwin, 12°24’S, 130°49’E, Mar 1979,
Rankin 1846 (CANB,DNA).

Distribution and habitat: Known only from the collections cited (Map 10); occurs usually
in sandy lateritic soil along the beach and in disturbed areas.

Z. disticha can be distinguished from other related species by the black dots on
stems and lower surface of leaflets, by the showy usually densely flowered inflorescences
with distichous congested bracts and by the hairy fruits with hairy bristles. It is related
to Z. muelleriana Mohl. in the pellucid-dotted leaves and stems, but differs chiefly in
the usually dark or black dots, congested bracts and ascending stems.

Note: Z. disticha includes plants from Australia previously included under Z. gibbosa
Span. by Mohlenbrock (l.c.). The type of Z. gibbosa Span. 1s from Koepang (Kupang)
Timor, Spanoghe, at L, yet when Mohlenbrock wrote up the species in his monograph,
he selected a neotype, namely Thorel 1426 (NSW, US) from Cai Cong, Cochinchina,
collected between 1862-66. His description of Z. gibbosa matches his neotype and agrees
quite well with the Australian plants and some of the Asian collections, e.g. Merri// 3787
(NSW), and Clemens 12476 (AD) from Philippines, and Mokin 805 (NSW) from Burma,
but does not agree with the type or the original description of Z. gibbosa, or that of Z.
graminea Span. from Timor, Spanoghe, at L which he synonymised under Z. gibbosa.
Z. disticha and the above Asian plants are distinguishable from Z. gibbosa as follows:

Leailets broadly elliptic or subovate; apex broad, + rounded. Inflorescences

to 8-flowered; bracts broad, + separated ..............2.... Z. gibbosa
Leaflets narrowly elliptic or oblong; apex subacute to obtuse and mucronate.
Inflorescences 6-—25-flowered; bracts narrow, congested ........ Z. disticha

(and Asian plants cited above)

The Asian collections, although superficially similar to Z. disticha, differ in having
larger leaflets, stipules and bracts (the latter becoming membranous with age as in Z.
prostrata and Z. albiflora).

_ Because of the differences of Z. disticha from the type of Z. gibbosa Span. and
Asian collections of that species, and also from the type of Z. graminea Span., we have
decided to keep it distinct from Z. gibbosa.

Reynolds & Holland, Zornia 23

§. Zornia maritima S. Reyn. & Holland species nova propter flores, folia et legumina Z.
prostratae S. Reyn. & Holland proxima autem praecipue floribus bracteis et
stipulis grandioribus (circa duplo grandioribus) et plerumque foliolis manitfeste
nervaturis reticulatis in pagina infera + nitentibus differt. Typus: Queensland.
Cook District: Lizard Island, 29 September 1988, G.N. Batianoff 10143 (holo:
BRI; 1so: BRI).

Herbs with prostrate, glabrous, epunctate stems 17-60 cm long. Petioles 0.8-5.7 cm long;
leaflets elliptic, ovate or ovate-oblong, 1.8-3.6 xX 0.6-1.4 cm, acute or subacute and
mucronate at apex, obliquely obtuse at base, glabrous or lower surfaces sparsely hairy;
lateral nerves oblique, conspicuously reticulate veined especially on lower surfaces;
margins densely ciliate; lower surfaces usually shiny and with few pellucid dots; petiolules
0.5-1 mm long, hairy. Stipules narrowly elliptic, 10-18 x 1.5-3 mm, acuminate at both
ends, 7-nerved, glabrous, sparsely pellucid-dotted, ciliate; spur 3-6.5 mm
long. Inflorescences 9.5-17.5 cm long, 5-17-flowered, usually with large, + distichous,
overlapping bracts towards apex, separated towards base (internodes 1.5-3 times as long
as bracts); peduncles 2.7-8 cm long; bracts ovate, (8—)11-13.5 X 3-6 mm, acute at apex,
usually with prominent truncate, bilobed or obtuse spur at base, 5-nerved with prominent
nerves and reticulate venation, ciliate, obscurely pellucid-dotted; spur 1.5-3.5 mm long.
Calyx 3-4 mm long; petals orange-yellow; standard 7-12 mm long. Fruits (2—)4-6-
articulate; articles 2.5-3.5 * 2—2.5 mm, hairy, pale with pale reticulate venation, covered
with bristles; bristles about 1 mm long, retrorse hairy.

Selected specimens: Queensland. Cook District: Weipa, track between Pappan Ck and Isenbert’s landing, Mar
1986, Foster 2001 (BRI), Lizard Island, Jun 1973, Fosberg 54927 (BRI); ditto, May 1975, Byrnes 3192 (BRI);
ditto, Sep 1988, Batianoff 10143 (BRD; Cape Flattery, 53 km NNE of Cooktown, Apr 1975, AfcDonald & Batianoff
{581 (BRI); Cape Bedford, Jun 1968, Pedley 2614 (BRI); Cooktown, Mar 1977, Scarth-Johnson 386A (BRI); Mt
Saunders, May 1964, Scarth-Johnson 1443A (BRI); Endeavour River, in 1882, Persieh 354 (MEL); Bloomfield
Beach, ca 1.6 km N of Bloomfield River mouth, Sep 1960, Smith 1020 (BRD.

Distribution and habitat: Known only from Queensland, from Cape York Peninsula
south to the Daintree River (Map 10); chiefly coastal, on sand dunes and headland
ridges.

Z. maritima is characterised by the usually shiny finely pellucid-dotted lower
surface of leaflets which are also usually conspicuously reticulate veined. Although the
leaflets are usually narrowly ovate-oblong, some plants with broader leaflets are difficult
to separate from the very variable Z. prostrata, but in the latter species the leaflets are
neither shiny nor dotted and not prominently reticulate on lower surfaces and the bracts
are also smaller. Further collections of Z. maritima may possibly show the two species
to be conspecific. Plants with conspicuous pellucid dots on leaflets also approach some
forms under Z. muelleriana Mohl. but the dots are too infrequent to allow inclusion
under the latter species.

Etymology: The specific epithet refers to the habitat in which this species is usually
found.

9, Zornia muriculata Mohl., Webbia 16: 75, f. 36 & f. 55 (1961). Type: Northern Territory.
Morgan’s Island, Blue Mud Bay, Gulf of Carpentaria, 20 January 1803, R. Brown
s.n. (holo: BRI; 1s0: KSMEL,NSW)

Z. diphylla var. hirsuta Domin, Biblioth. Bot. 89: 209 (1926). Type: Queensland.
Mareeba, February 1910, K. Domin (syn: PR); Chillagoe, February 1910, K. Domin
(syn: PR); Castle Hill, Townsville, February 1910, K. Domin (syn: PR), South
Percy Island, 5 March 1906, H. Tryon (syn: PR; isosyn: BRI).

Z. diphylla var. xerophila Domin, |.c. Type: Queensland. Cook District: Walsh
River, north of Chillagoe, February 1910, K. Domin (syn: PR); near Lappa
Junction, February 1910, K. Domin (syn: PR).

Herbs to 35 cm high with usually erect, tufted, hairy epunctate stems, rarely plants
semiprostrate or glabrous. Petioles 0.5-—2.5 cm long; leaflets very variable, obovate,
elliptic, oblong, obovate or linear, (0.5-)0.7-3.2 cm long, 0.5-9 mm wide, lower ones
smaller and broader than upper ones, obtuse or acute and mucronate at apex, base
obtuse or subacute, glabrous or subglabrous above, sparsely hairy below, epunctate or
rarely with few obscure pellucid dots; lateral nerves 1-3 pairs, basal pair erect ascending

24 Austrobaileya 3(1): 1989

along margin, conspicuous or obscure, usually darkish coloured; petiolules 0.5-1 mm
long. Stipules usually conspicuous, elongate, narrowly elliptic, ovate, + falcate, 5-14 x
i-2.5 mm, acuminate at apex, spurred at base, 3-45-nerved; reticulate venation obscure:
spur 0.5-4 mm long, acute, acuminate or truncate. Inflorescences 2.5-—9.5(—15) cm long,
3-15(very rarely 20-23)-flowered, with overlapping to separated bracts; peduncles 0.3-
1.5(-2.8) cm long; bracts narrowly ovate or + falcate, 5.5-15 * 1-5 mm, acuminate at
apex, usually narrowly spurred at base, ciliate, epunctate or rarely with few pellucid dots,
3~5-nerved; spur 0.5-5.5 mm long, acute or acuminate. Calyx 2-3 mm long; petals
yellow with red markings; standard 5-9 mm long. Fruits 1-4(—6)-articulate, exerted or
included; articles 2-3 X 1.5-2.5 mm, usually whitish with reddish reticulate venation,
epunctate or with a few glands, glabrous (sometimes finely hairy), with small, blunt or
acute, or well developed bristles; bristles mostly at periphery, to 1 mm long.

Z. muriculata 1s characterised by the usually erect, hairy (or rarely glabrous) stems,
congested bracts, usually glabrous fruits with long or small short bristles and usually
elongate conspicuous stipules. The majority of plants have prominently nerved and
reticulate veined leaflets and bracts.

Leaflets, bracts, stipules, fruits and hairiness are all very variable, sometimes even
in one individual plant but the species nevertheless remains distinctive. Usually plants
are erect but sometimes those near coastal areas are semiprostrate, and have laxly
branched, elongate, glabrous or sparsely hairy stems, conspicuous long stipules, and 1-
3-articulate fruits which are muriculate or with few long weak bristles. Plants from
further inland are usually densely hairy with much-branched, tufted, erect stems, small
stipules and 2-4-articulate fruits with prominent muriculations or with + rigid, long
bristles. This is the common form in the Northern Territory. Some plants from coastal
areas with long, + semiprostrate stems resemble some of those under Z. dyctiocarpa
sat filifolia but that taxon has very interrupted few-flowered inflorescences and hairy
ruits.

Two subspecies are recognised, although they are very variable and merge into
each other.

Leaflets (3-)4-9 mm wide, 2-6(—7) times as long as wide, sometimes pellucid-
dotted; nerves usually prominent. Stems erect (rarely semiprostrate),
laxly branched, glabrous or hairy. Stipules conspicuous, (7-)10-14 mm
long. Articles 1-4, with few short or long weak bristles .. .. | subsp. muriculata

Leaflets 0.5-3.5 mm wide, (5-)7-10 times as long as wide, usually not
pellucid-dotted; nerves obscure or absent. Stems erect, tufted, usually
hairy. Stipules 5-7 mm long. Articles (1-)2-5, covered with conspic-
uous, + rigid, long bristles, or bristles few, small, acute or
blunt . beweh 4a Aen ah A AUR Ss Grud etl th aes ah ee: subsp. angustata

Zornia muriculata subsp. muriculata
Z. diphylla var. hirsuta Domin, Lc.

Distinguishing characters are as set out in the key above.

Plants usually with prominently nerved leaflets, stipules and bracts. Leaflets 1.8-3.2 cm
long, (3-)4-9 mm wide, rarely less, nerves mostly dark and ascending. Stipules (7-)10-
14 mm long, sometimes longer than petiole, often + falcate. Inflorescences 3—10(—13)-
flowered; bracts (8-)10-15 mm long, overlapping towards apex, interrupted below:
peduncles 0.7—2.5 cm long. Articles 1—4, mostly pale or glaucous with reddish reticulation,
muriculate or with few weak fine bristles at periphery.

Selected specimens: Northern Territory. Morgan’s Island, Blue Mud Bay, Jan 1803, Brown (BRI,CANB,K,NSW).
Queensland. Cook District: Goode Island, in 1881, Powell (MEL); Mabiuag Island, May 1911, Bick 64A (BRI);
Andoom, Arunta, 13 km N of Lorim Point, Weipa, Apr 1982, Aforton 1681 (BRI); Near Atherton, Jan 1966,
Hyland 3880 (BRD. NoRTH KENNEDY DISTRICT: Rockingham Bay, Dallachy (BRI,MEL). Port Curtis District:
S Percy Island, Mar 1906, Tryon (BRI).

Distribution and habitat: Chiefly coastal Northern Territory and Queensland, from Gulf
of Carpentaria and Cape York Peninsula to Central Queensland, also on offshore islands

Reynolds & Holland, Zornia 25

(Map 5); occurs usually in sandy soil in open woodlands or grasslands, or in cleared
areas.

Subspecies muriculata is very variable with at least two taxa recognisable within
this subspecies but because of the presence of intermediate forms they are not formally
segregated here. The typical form has elongate, ascending (or semiprostrate), glabrous or
sparsely hairy stems and narrow leaflets and occurs along the coast of northern Queensland
to as far south as Shoalwater Bay, including islands of the Gulf of Carpentaria and
Torres Strait. Leaflets of these plants are elliptic-oblong or + falcate and 4-6 times as
long as wide. Their stipules are conspicuous, + falcate and sometimes longer than the
petioles, and their bracts usually have a narrow falcate spur. Fruits 1-4-articulate; articles
have short, sharp or blunt bristles, or weak, slightly longer fine bristles.

The broad leaved hirsute form occurs on the Atherton Tableland and extends to
Percy Island. Plants are hairy with usually short tufted stems and broad leaflets. Their
leaflets are elliptic, oblong or obovate, and 2-4 times as long as wide. Stipules and bracts
are smaller than in the typical form and the bracts mostly have a short spur. Fruits are
2-4-articulate: articles usually have short blunt bristles or with weak acute longer bristles.
Some of these plants were described as Z. diphylla var. hirsuta by Domin.

Collections from the Percy Islands are intermediate between these two forms and
also subsp. angustata. Future collections may show them to represent a distinct variety
or subspecies.

Zornia muriculata subsp. angustata S. Reyn. & Holland subsp. nov. a subsp. muriculatae
Mobhl. foliolis angustatis minoribus, stipulis bracteisque minoribus, caulibus ramo-
sissimis caespitosis plerumque villosis, et leguminibus setosis muriculatisve differt.
Typus: Northern Territory. Near Balbirini Creek (16°35’S, 135°20’E), alt. 189-
192 m, May 1947, S.T. Blake 17742 (holo: BRI; iso: CANB,NT).

Z. diphylla var. xerophila Domin, Lc.
Distinguishing characters are as set out in the key above.

Plants usually densely villous, with tufted much-branched, erect stems and very narrow
and usually small leaflets, small stipules and bracts. Leaflets narrowly elliptic or linear,
0.5-2.1(-2.5) cm long, 0.5~2(-3.5) mm wide, (5-)7-10 times as long as wide. Stipules
5-7 mm long. Inflorescences (4~-)9-15(rarely 20-23)-flowered; bracts 5.5-9.5 mm long,
overlapping to separated; peduncles 0.3-1.5(-2.8) cm long. Articles 1-5, covered with
prominent, + rigid bristles (mostly Northern Territory plants) or with few, small, blunt
or acute ones (chiefly Queensland plants).

Selected specimens: Western Australia. Packsaddle Ck, northern Carr Boyd Ranges, Mar 1978, Hartley 14397
(CANB): Mt Beli, Jun 1978, George 15148(PERTH). Northern Territory. Near Batten Ck, about 10 km along
Ryan Bend waterhole Rd, Feb 1976, Craven 3854 (CANB); 94 miles {162 km] E of Daly Waters, Mar 1972,
Byrnes 2509 (CANB,DNA,NT); Tanumbirini Station, Mar 1979, Henshall 2543 (NT), Balbirini Ck, May 1947,

Blake 17742 (BRI,CANB,NT). Queensland. Cook District: Walsh, in 1891, Barclay Miller (BRI). BURKE
District: 7 km W of Kajabbi, on Mt Cuthbert Rd, Feb 1978, Schmid 420 (BRI). NoRTH KENNEDY DISTRICT:
Ravenswood, Mar 1943, Blake 14862 (BRI). MircHeLt District: about 15 miles [24 km] S of Muttaburra, Jun
1949. Everist 3889 (BRI). DARLING Downs District: Brookvale Park area, 12 km W of Oakey, Apr 1980,
Ballingall 2139 (BRI). New South Wales. 7 miles [11.5 km] NW of South Grafton, Gwydir Highway, Feb 1973,
Tindale 2047 & Wilson (NSW), Oxiley’s Station, Mar 1843, Leichhardt [NSW 183612] (NSW).

Distribution and habitat: Widespread in northern Australia, from the Kimberleys, Western
Australia, to Darwin and Gulf District, Northern Territory and in north west and eastern
Queensland to as far south as Grafton in New South Wales (Map 6); occurs usually in
— ae soil, on stony ridges, slopes, rocky outcrops and sandstone hills, in open
woodlands.

Subspecies angustata is very variable; several forms are included but because they
intergrade into each other throughout their range, they are not formally segregated here.
Plants from northwestern Queensland and Northern Territory have small narrow leaflets
and 2—4-articulate fruits covered with + rigid fairly long bristles, and inflorescences with
congested overlapping small bracts and short peduncles. Most of these are densely hairy
(except a collection from near Katherine which is glabrous). The majority of plants from
Queensland and from New South Wales have longer, narrow leaflets, interrupted inflo-
rescences and 2—4(-5)-articulate fruits, articles are with few small blunt or acute bristles.

26 Austrobaileya 3(1): 1989

A group of plants from around the Palmer and Walsh Rivers, and from Ravenswood
and Townsville in Queensland, have linear, + concave, small leaflets, very interrupted
inflorescences with usually small bracts and few flowers, and 1-3(—4)-articulate mostly
included fruits; articles have a few weak bristles only at margins. These plants were
described as Z. diphylla var. xerophila by Domin. Determining whether they are distinct
from other forms of subsp. angustata or not must await study of further material.

Etymology: the subspecific epithet refers to the narrow leaflets of the subspecies.

10. Zornia floribunda S. Reyn. & Holland species nova; planta inter Z. muriculatae
Mohl. et Z. dyctiocarpae DC. quasi intermedia, quoad formam foliorum et
leguminum cum Z. dyctiocarpae sat congruens sed foliolis longioribus, bracteis
congestibus et caulibus erectis differt; a Z. muriculatae haud aegre foliolis inflo-
rescentis et pedunculis longioribus et leguminibus (1—-)3-6-articulatis puberulis
setulosis distinguitur. Typus: Queensland. Port Curtis District: Gladstone, 10
March 1937, S\7T. Blake 12814 (holo: BRI).

Herbs to 50 cm high, with erect, sparsely branched, glabrous or sparsely antrorse hairy,
epunctate stems. Petioles 12-25 mm long; leaflets narrowly ovate-oblong, elliptic, or +
linear, 2.6-4 cm long, 1-2.5 mm wide, acute at apex, glabrous, rarely with few obscure
marginal pellucid dots on lower surfaces; petiolules to 1 mm long, pilose. Stipules
narrowly elliptic or + triangular, 11-15 X 1.5-2 mm, with prominent basal spur and
acuminate at both ends or truncate at base, 5-nerved; spur 2—3.5 mm long. Inflorescences
14-22 cm long, (4—-)13-20-flowered with bracts barely touching to overlapping about half
their length towards apex or separated, especially towards base (internodes to 3 times
as long as bracts); peduncles 2.7-8 cm long; bracts narrowly ovate-elliptic (7-)
11-12 x 3-5 mm, with a small basal spur, 5-nerved, with few obscure dots, ciliate; spur
1-2.5 mm long, acute. Calyx 3-4 mm long; petals yellow, streaked red: standard 7-9
mm long. Fruits (1-)3~6-articulate; articles 2 X 2.5 mm, finely hairy, with a few peripheral
bristles or with small bristles scattered over the article, prominently reticulate veined
and with small and obscure glands, rarely glabrous and long bristly.

selected specimens: Queensland. NORTH KENNEDY District: Approx. 40 miles [64 km] SE of Mt Garnett, Nov
1967, Morain 231 (BRD. SourH KENNEDY District: 8 km SE of Eungella Dam, Apr 1978, Byrnes 3627 &
Clarkson (BRI). Port Curtis District: Gladstone, Mar 1937, Blake 12814 (BRI); 52 miles (83.2 km] from
Gladstone on Biloela Rd, Apr 1962, Johnson 2268 (BRI). LEICHHARDT District: 4 miles [6.4 km] NE of New
Twin Hills H.S., Aug 1964, Pedley 1736 (BRI). BURNETT District: 30 km NNE of Gayndah, Dec 1972, Pedley
4010 (BRI). Wipe Bay District: Elliott Railway Bridge near Bundaberg, Dec 1938, Goy & Smith 585 (BRI);
Toolara $.F. Cpt £02, South Dempster, Feb 1983, Podberscek 2 (BRI).

Distribution and habitat: From near Mt Garnett in northern Queensland to northern
New South Wales (Map 2); occurs usually in sandy soil, on sandy ridges and hillsides
in eucalypt forests.

Z. floribunda is distinguished by the usually densely flowered long inflorescences
with + congested bracts and long peduncles, exserted, hairy or glabrous, moderately or
sparsely bristly fruits and also by the long narrow leaflets and erect stems. It is closely
related to Z. muriculata Mohl. and Z. dyctiocarpa DC. and is almost intermediate
between these species and has been confused with both in the past. It agrees with Z.
dyctiocarpa in the nature of the leaves and to a certain extent the fruits but differs in
habit, in longer leaflets, congested bracts, bristly, hairy or glabrous fruits. From Z.
muriculata it can be distinguished by the long narrow leaflets, long inflorescences and
peduncles, and usually hairy, (1-)3-6-articulate fruits.

Note: Leaflets and bracts are very variable. Plants from around Gladstone and Rock-
hampton are typical of the species having elongate narrow leaflets and long many-
flowered inflorescences with + congested bracts. Plants from southern Queensland and
northern New South Wales (e.g. Bingara Lookout, Feb 1977, Wilson (NSW)) are, however,
atypical and approach Z. dyctiocarpa DC. in leaves and bracts (but in the latter species
the bracts are separated and the stems are prostrate). More collections are needed
(especially from the latter localities 1.e. southern Queensland and New South Wales) to
determine whether the stems are always erect and the bracts always + overlapping.

Reynolds & Holland, Zornia 27

11. Zornia ramosa S. Reyn. & Holland species nova, Z. dyctiocarpae var. filifoliae
(Domin) S. Reyn. & Holland primo aspectu maxime similis sed floribus solitariis
et caule plerumque erecto ramosissimo differt. Typus: Queensland. COOK DISTRICT:
Thursday Island, 14 May 1901, Jaheri s.n. (holo: BRI; iso: K).

(Z. filifolia auct. non Domin: Mohlenbrock, Webbia 16: 52, f. 32, 34 (1961).]

Herbs 20-60 cm high, with + erect or ascending, slender, usually much-branched,
glabrous, epunctate stems. Petioles filiform, 5-16 mm long, glabrous; leaflets narrowly
ovate to linear, 1-3 cm long, 0.51 mm wide, acuminate at apex, obtuse at base, glabrous,
epunctate, rarely with few small dots on lower surfaces; midrib prominent; petiolules to
1 mm long. Stipules narrowly rhombic, 2-6 X 0.5 mm, with narrow basal spur, acuminate
both ends, glabrous, rarely with few pellucid dots, 3-5-nerved; nerves inconspicuous,
margins pale; spur 0.5-1.5 mm long. Inflorescences of solitary flowers usually on long,
filiform, glabrous peduncles, 1.2-2.5 cm long; bracts narrowly ovate or falcate, 5-7 x
[-2 mm, acute or acuminate at both ends, with slender spur at base, 3-nerved, epunctate
or with few pellucid dots; spur 1-2 mm long. Calyx 2-3 mm long; petals yellow or
purplish; standard 5-7 mm long. Fruits (1-)2—4-articulate, crenate on lower margins,
exerted: articles 2-2.5 X 1.5-2 mm, puberulent, prominently reticulate veined, bristly;
bristles to 0.5 mm long, hairy.

Selected specimens: Queensland. Cook District: Thursday Island, May 1901, Jaheri s.n. (BRI,K); ditto, without
date, Cowley 14 (BRI); Lockerbie, 10 miles [16 km] WSW of Somerset, Apr 1948, Brass 18441 (BRICANB);
ditto, May 1948, Brass 18584 (BRI,CANB); 45 km N of Coen (13°36’S, 143°02’E), Jun 1981, Maconochte 2708
(BRI); Hann River Crossing, Lakefield National Park (15°1~’S, 143°-’E), Jul 1983, Hacker 420 (BRI).

Distribution and habitat: So far known only from Cape York Peninsula, Queensland
(Map 13); occurs in sandy soil on lateritic ridges, riverbanks and in disturbed areas,
among grasses.

Z. ramosa is distinguished from other Australian species (except Z. pedunculata
S. Reyn. & Holland) by its solitary flowers. (It is distinguished from Z. pedunculata by
the narrow leaflets and hairy bristly fruits). It resembles Z. dyctiocarpa var. filifolia S.
Reyn. & Holland, in leaflets, stipules, bracts and fruits but the flowers are not solitary,
and the stems are prostrate in the latter species.

Note: Z. ramosa is a name provided for plants previously called Z. filifolia by Mohlen-
brock (1961) and plants identified as such using his monograph.

Mohlenbrock’s treatment of Z. filifolia Domin and Z. adenophora (Domin) Mobhl.
has created much confusion with regards to the identity of the species concerned. When
he described Z. filifolia, he cited a single specimen, namely Cowley 14 from Thursday
Island, without date, (BRI), and his description and illustration matched this specimen.
It is not, however, conspecific with the type of Z. flifolia Domin, viz Domin’s collections
from Rocky Island, Yarrabah (PR). It differs chiefly in the type of inflorescence, indicating
two distinct species are involved. In addition to this, when Mohlenbrock raised Z.
filifolia var. adenophora Domin to species rank as Z. adenophora, he included Z. filifolia
var. subeglandulosa Domin in synonymy. That name was not validly published as the
taxon it referred to included the type of Z. filifolia and hence should have been published
as Z. filifolia var. filifolia. Mohlenbrock’s concept of Z. adenophora thus includes the
type of Z. filifolia so the name he used for that species should have been Z. filifolia
Domin. The plants Mohlenbrock identified as Z. filifolia thus require a different name,
and since no other names appear applicable, the name Z. ramosa has been provided for
the species concerned. Since the type of Z. filifolia var. filifolia and of Z. filifolia var.
adenophora are not conspecific, two other taxa are involved. They are treated here as
Z. dyctiocarpa var. filifolia and Z. adenophora respectively. Mohlenbrock’s confusion
with these plants is highlighted by his determination of the BRI sheet of Cowley 14 as
Z. ramosissima, a name that was not subsequently published, and of the BRI sheet of
Jaheri s.n. (type of Z. ramosa) as Z. cantonensis Mohl., both belonging to the one taxon.
The latter specimen was not cited in his account and did not match collections of Z.
cantonensis he did cite.

The three taxa that have been confused, namely Z. filifolia Domin (now Z.
dyctiocarpa var. filifolia), Z. adenophora and Z. ramosa can be distinguished as follows:

28 Austrobaileya 3(1): 1989

|. Leaflets linear, conspicuously densely pellucid-dotted. Fruits covered with
large glands; bristles absent ........................ Z, adenophora
Leaflets linear or narrowly ovate, not pellucid-dotted or rarely with few
obscure pellucid dots. Fruits eglandular, bristly or non bristly... .. .... .. 2

2. Inflorescences few-flowered with very separated bracts, rarely flowers
solitary (then in combination with spikes on same _ plant).
Stems prostrate, usually laxly branched. Fruits usually scarcely
bristled .............................. 4. dyctiocarpa var. filifolia
Fiowers solitary throughout. Stems + erect or ascending, usually much-
branched. Fruits bristly .. .. .. aast ey he. ete Ae zetia, es te Z.. ramosa

12. Zornia pedunculata S. Reyn. & Holland species nova quoad flores solitarios cum Z.
ramosa S. Reyn. & Holland congruens sed caulibus + prostratis, foliis ovatis et
leguminibus glabris differt. Typus: Queensland. PortT Curtis District: 60 km
SW of Gladstone, Kroombit Tableland, tributary of Kroombit Creek, 3.2 km
SSW of Amy’s Peak (24°22’S, 150°59’E), alt 800 m, 4 June 1977, ALD. Crisp
2766 (holo: BRI).

Herbs with slender, few-branched, glabrous, obscurely pellucid-dotted stems. Petioles
0.9-1.6 cm long; leaflets ovate, 0.9-19 cm long, 3-5 mm wide, acute and mucronate at
apex, obtuse at base, slightly oblique, glabrous, epunctate, lateral nerves obscure; petiolules
ca | mm long, subglabrous. Stipules narrowly elliptic or rhombic, 7-9.5 * 1.5 mm,
acuminate at apex, acute and prominently spurred at base, 5~7-nerved, sparsely punctate:
spur ca 2.5 mm long. Inflorescence of solitary flowers; peduncles 3-4 cm long, filiform:
bracts ovate, 10 X 2.5 mm, acuminate at apex, with prominent basal spur, 5-nerved,
glabrous, obscurely yellow pellucid-dotted; spur ca 2.5 mm long. Flowers not seen. Fruits
3- or 4-articulate; articles glabrous, prominently reticulate veined with red venation; with
short bristles or bristles absent, sparsely gland-dotted.

Distribution and habitat: Known only from type from central Queensland (Map 13):
rare on sandy clay loam on creek flats in open forests.

Z. pedunculata is easily recognisable by its solitary flowers, long peduncles, ovate
leaflets, and glabrous fruits with short bristles or lacking bristles. Further collections are
needed to determine whether the flowers are always solitary as in Z. ramosa which has
narrowly ovate to linear leaflets and hairy bristly fruits. Nevertheless the plants are
sufficiently distinct to be described as belonging to a new species. A number of new
species in other families have been found on Kroombit Tableland in recent years.

Etymology: The specific epithet refers to the long peduncles.

13. Zornia pallida Mohi., Webbia 16: 130, f. 36 & 87 (1961). Type: Queensland. MARANOA
DisTRICT: Bungeworgoral, May 1913, J. Soutter s.n. (holo: BRI(AQ022931)).

Prostrate herbs with glabrous epunctate stems. Petioles 15-20 mm long; leaflets narrowly
ovate-oblong, 1.1—3.2 cm long, 4.5-6 mm wide, acute at apex, obtuse at base, (lower
ones usually smaller and broader than upper ones), glabrous, sparsely pellucid-dotted
below; lateral nerves obscure; petiolules ca 1 mm long. Stipules narrowly elliptic, 8.5-
1Q x I.5-2 mm, acute or acuminate at apex, prominently spurred at base, 5-nerved.,
epunctate;, spur ca 2.5 mm long, acuminate or truncate. Inflorescences 12-16 cm long,
9-13-flowered with bracts overlapping about 1/4 their length or barely touching each
other, usually separated towards base; peduncles 4—-4.5 cm long; bracts elliptic-ovate, 10-
[2 X 5-6 mm, acute at apex, prominently spurred at base, 5—-7-nerved, obscurely reticulate
veined, epunctate or rarely with few pellucid dots; spur 1-2 mm long, acute or sometimes
truncate. Calyx 3.5-5 mm long; standard to 12 mm long. Fruits 3—5-articulate; articles
2-3 X 3-3.5 mm, hairy or glabrous with hairy or glabrous prominent bristles; bristles
I-5 mm long.

Selected specimens: Queensland. PORT CURTIS DisTRICT: Broadsound, collector unknown [MEL 1551174] (MEL):
Stanage Point, about [00 miles [160 km] N of Rockhampton, Aug 1964, Gittins [NSW 14261] (BRI,NSW).
LEICHHARDT District: Nebo Creek, Bowman [MEL 1554084] (MEL). MARANOA District: Bungeworgorai, May
1913, Soutfer (BRI). BURNETT District: Gayndah, Aug 1959, Schoneveld 134 (BRI).

Reynolds & Holland, Zornia 29

Distribution and habitat: Known only from Queensland (from near Rockhampton to
Gayndah) (Map 9); occurs usually in eucalypt woodlands or disturbed areas.

Z. pallida is characterised by the narrowly ovate-oblong leaflets with sparsely
pellucid-dotted lower surfaces and overlapping, comparatively large bracts with a prom-
inent basal spur, and 3—5-articulate fruits with large, usually hairy, bristly articles.

It is closely related to Z. dyctiocarpa DC. having similar leaflets and bracts but
differs in the overlapping bracts, the large hairy articles and prominent hairy bristles.
The fruits and flowers especially those in collections from Broadsound and Stanage Point,
are not unlike those of Z. prostrata, but in that species the leaflets are usually wide and
bracts separated.

Note: This species is known from only a few, mostly inadequate collections. It is almost
impossible to separate sterile specimens of it from the variable Z. dyctiocarpa. More
collections are needed to determine its full variability and ascertain its relationship with
the latter species.

14. Zornia dyctiocarpa DC., Prod. 2: 317 (1825). Type: Novae Hollandiae, Sieber 610
(Microfiche BRI).

Z. filifolia Domin, Biblioth. Bot. 89: 210 (1926). Type: Queensland. CoOK DISTRICT:
Rocky Island, Yarrabah, January 1910, K. Domin (PR(PR 527341-42)).

Z. filifolia var. subeglandulosa Domin. l.c., nom. inval.

[Z. diphylla auct. non (L.) Pers.: Bentham, Fl. austral. 2: 228 (1884); Bailey, Qd FI.
2: 409 (1900); Domin, Biblioth. Bot. 89: 208-209 (1926).|

Herbs with prostrate or semiprostrate, usually dichotomously branched stems 9-52 cm
long. Stems, petioles, leaflets, stipules and bracts glabrous or with few appressed hairs,
epunctate, rarely with few minute glands. Petioles 1.5-3 cm long; leaflets variable on
one plant, ovate, narrowly ovate-oblong or elliptic to linear, 1-3 cm long, 1-8 mm wide
(lower ones smaller and broader than upper ones), apex acute or acuminate and
mucronate, base obtuse or acute, midrib usually sunken above; lower surfaces sometimes
with few minute marginal dots; petiolules to 1 mm long. Stipules narrowly ovate or
subrhombic, 5-13 X 0.5-2 mm, acute or acuminate at apex with a narrow spur at base,
slabrous, rarely ciliate, 3~5-nerved, reticulate venation obscure; spur |-} mm long, acute
or acuminate. Inflorescences 4-28 cm long, (1-)2—12-flowered, with widely separated
bracts (internodes 2—5 times as long as bracts); peduncles 3-13 cm long; bracts ovate or
elliptic, sometimes narrow, oblique or falcate, S-15 x 1-5 mm, acute or acuminate at
apex, usually with a conspicuous long and narrow spur at base, glabrous, ciliate, 5-
nerved, reticulate venation obscure, rarely with few pellucid dots; spur 1-3 mm long,
acute or toothed. Calyx 2-4 mm long; petals yellow or orange, with red flecks; standard
5-9 mm long. Fruits (2-)4—-9-articulate, rarely: less; articles 1.5-3 < 1.7-3 mm, thickly
margined, glabrous or rarely puberulent, conspicuously reticulate veined with reddish
brown venation, eglandular or rarely with few glands; bristles absent or if present to 0.5
mm long, retrorse hairy, usually at periphery of article.

Z. dyctiocarpa is characterised by the long inflorescences with very interrupted
bracts, and long, exserted, 4—9-articulate fruits with prominent + dark, reddish brown
reticulate venation, by the prominent long spur at the base of bracts and by the narrowly
ovate-oblong, elliptic or linear leaflets, with smaller broader ones at base.

Note: Leaflets, bracts, stipules, number of flowers and articles vary considerably in the
same population as does the presence or absence of bristles on the articles. The majority
of plants from New South Wales have ovate, elliptic or narrowly ovate-oblong leaflets
and prominently spurred bracts, while most of the plants from Queensland have narrower
leaflets and smaller bracts with a small spur. The inflorescences of some of these latter
plants are few-flowered and the bracts narrower than is usual in the species, and the
fruits are 2—4-articulate and hairy. Z. filifolia Domin is based on some of these north
Queensland plants which we consider varietally distinct but the extreme forms of each
eroup merge into each other. The two varieties may be distinguished as follows:

30 Austrobaileya 3(1): 1989

Leaflets (2.5-)3-7.5 mm wide, less than 10 times as long as wide. Bracts
2.5-5 mm wide, spur 1.5-4 mm long. Fruits (4-)6-9-articulate. Inflo-
rescences (2—)6—-13-flowered eee vege ree ee var. dyctiocarpa
Leaflets 1-2.5 mm wide, 10-22 times as long as wide. Bracts 1-2 mm wide:
spur 0.5-1.5 mm long. Fruits(1—)2-4(—5)-articulate. Inflorescences
(1~)2-S(very rarely—7)-flowered ........................ var. filifolia

Zornia dyctiocarpa var. dyctiocarpa
Distinguishing characters are as set out in the key above.

Stems prostrate; leaflets ovate, elliptic, ovate-oblong or narrowly ovate, 1-3.5 cm long,
3-7.5 mm wide, (2-10 times as long as wide), lower ones usually smaller and broader
than upper ones. Inflorescences with widely separated bracts. Bracts ovate or elliptic,
(8-)10-13 X 2.5-5 mm, with prominent spur at base; spur narrow and + falcate.
Peduncles (4-)8-13 cm long. Articles 1.5-3 X 2~3 mm, with prominent reddish brown
reticulate venation, glabrous or rarely finely hairy, bristle-free or sometimes bristly
(mostly towards margins); bristles usually small and weak.

Selected specimens: Queensland. NORTH KENNEDY DISTRICT: Rockingham Bay, Dallachy (MEL). SouTH KENNEDY
District: 8 km SE of Eungella Dam, Apr 1978, Byrnes 3627 & Clarkson (BRI). LEICHHARDT District: Great
Dividing Range near Rolleston, Jun 1977, Crisp 3041 (MEL). Wipe Bay District: Fraser Island, Oct 1930,
Hubbard 4491 (BRD. BURNETT District: Nanango, Dec 1930, Watson (BRI). MORETON District: Peregian
Beach, Feb 1979, Batianoff 1340 & Elsol (BRI). New South Wales: Pimlico near Ballina, Mar 1894, Bauerlen
[NSW 183394] (NSW); Armidale, in 1871, Perrott (MEL); Barrangan beyond Mudgee, collector unknown (MEL);
Tenterfield, Jan 1871, Sturt (MEL); Tabulan, Nov 1904, Boorman [NSW 46741] (NSW); Little River, 12.5 km
ENE of Jenolan Caves, Jun 1971, Rodd 1614 (NSW); Black Mt, Canberra, Nov 1949, Gauba (AD,MEL); Bega,
Dec 1920, Rodway [NSW 46769] (NSW).

Distribution and habitat: From Rockingham Bay, Queensland, to Bega River, New South
Wales (Map 3); occurs usually in sandy soil in open forest and grasslands, on sand
dunes, low hills and ridges and in disturbed areas.

Zornia dyctiocarpa var. filifolia (Domin) S. Reyn. & Holland comb. nov.

Z. filifolia Domin, Biblioth. Bot. 89: 210 (1926). Type: Rocky Island, Yarrabah,
January 1910, K. Domin (syn: PR(PR 527341-42))

Z. filifolia var. subeglandulosa Domin, |.c., nom inval.
Distinguishing characters are as set out in the key above.

Stems very slender, few-branched, prostrate or + prostrate. Leaflets usually linear, +
concave, 1.2-2.7 cm long, 1-2(-2.5) mm wide. Inflorescences with few (1-5 rarely -7),
widely spaced, small, narrow bracts. Bracts narrowly elliptic, 5-7 * 1-2 mm: spur narrow
and + falcate. Peduncles 3.5-6 cm long. Articles 1.7-2 * 1.7—-2 mm, usually finely hairy,
bristles few, usually small and weak or absent.

Selected specimens: Queensland. COOK DistRIcT: Yorkey’s Knob near Cairns, Apr 1962, AfcKee 9037 (BRI,CANB);
Rocky Island, Yarrabah, Jan 1910, Domin (PR). NORTH KENNEDY District: Palm Island, Bancroft [AQ 239394]
(BRI). SOUTH KENNEDY District: Pind: Pindi, Feb 1959, Mathews 24 (BRI.

Distribution and habitat: Chiefly coastal Queensland, from near Mossman River to Pindi
Pindi (Map 4); occurs usually in sandy rocky soil in disturbed areas.

Note: Forms intermediate between the varieties are present around Brisbane and Coaldale,
northern New South Wales.

Z. dyctiocarpa var. filifolia is recognisable by its very slender, few-branched stems,
by the very narrow leaflets and bracts, and by inflorescences with widely spaced few
flowers. The plants resemble Z. ramosa in general appearance but the flowers are solitary
in that species. Some semi-prostrate forms of Z. muriculata are also difficult to distinguish
from this variety, but in Z. muriculata the inflorescences usually have more flowers and
overlapping bracts, and the fruits are usually glabrous and muriculate or with long
bristles.

Typification: The type of Z. dyctiocarpa DC. is Sieber 610, from New Holland, exact
locality unknown (not recorded on microfiche) and not Sieber 31 as cited by Mohlenbrock
(1961). It is not known why he stated the latter, from Barrangan beyond Mudgee NSW

Reynolds & Holland, Zornia 3]

(MEL 1554072) as an isotype, as the type is clearly indicated by De Candolle as Sieber
610. There is also uncertainty with regards to the collector of Sieber 31. It is doubtful
whether Sieber collected this specimen or that “No. 31” is his collecting number. The
name of the collector is not given on the two slips of paper attached to the MEL sheet
and the two handwritings thereon do not appear to match that of Sieber. Also, the “31
Zornia dyctiocarpa’ written on a slip of paper seems more likely to be part of an
‘dentification list that has been cut into pieces, rather than a collector’s number. The
handwriting on the blue label giving the place of collection is very similar to that of
Allan Cunningham who collected in that area in 1823 and who used similar blue paper
to record his collecting data. It is possible that he collected this specimen numbered 31.
Although we have not been able to locate the type, we are confident that Mohlenbrock’s
concept of the species is correct. The specimens cited, mostly annotated by him, agree
with the protologue and with the microfiche (at BRI) of Sieber 610. Moreover Sieber
collected in New South Wales where this plant is a common species and his specimen
(microfiche) is a good match for plants from New South Wales.

15. Zornia albiflora Mohl., Webbia 16: 65, f. 40, 48 (1961). Type: Northern Territory.
| mile [1.6 km] S of Elkedra H.S., 22 October 1956, G. Chippendale 3104 (holo:
BRI: iso: AD,CANB,MEL).

Z. nervata Mohl., Webbia 16: 130 (1961). Type: Queensland. GREGORY NORTH
District: 50 miles [80 km] SE of Camooweal, 12 December 1947, S.L. Everist
3388 (holo: BRI).

Hairy herbs 13-30 cm high with much-branched, erect or spreading stems. Petioles 0.4-
2.5 cm long, densely patent hairy; leaflets broadly elliptic or oblong, oblique, 0.6-2.1 X
0.4-1.3 cm, obtuse or subacute at apex, obtuse at base, sparsely hairy above, densely
hairy below with appressed hairs, epunctate, rarely with few obscure pellucid dots; jateral
nerves oblique, prominent; petiolules to 1 mm _ long. Stipules narrowly elliptic or
subrhombic, 5~9 X 1.5-3 mm, acute at apex, with a long, narrow, acute spur at base,
3-5-nerved: nerves usually prominent and reddish in colour; spur 1-3.5 mm long.
Inflorescences 4-7 cm long, (1-)3—10(—21)-flowered, with overlapping distichous bracts
+ separated towards base (internodes i/2-1(-2) times as long as bracts); peduncles 1.5-
4 cm long; bracts broadly ovate to suborbicular, 6.5-12.5 * 4-8 mm, acute at apex,
usually shortly spurred or subcordate at base, 4-7-nerved, sparsely hairy, ciliate, epunctate
or rarely with a few glandular dots; nerves and reticulate venation conspicuous, reddish
coloured, especially in older specimens; spur i-~3 mm long, truncate or obtuse, sometimes
minute and inconspicuous or almost absent. Calyx 3-3.5 x 1.5-2.5 mm, petals white or
yellow; standard 4-6 mm long. Fruit (1-)2-4(-5)-articulate; articles 2-3.5 X 2-2.) mm,
finely puberulent and with hairy bristles, finely reticulate veined usually with pale
reticulation the same colour as the articles; bristles fine, 0.5-1.5 mm long, retrorse hairy.

Selected specimens: Western Australia. Just NW of Wolf Creek Crater, Apr 1979, George 15286 (NT,PERTH);
McLarty Hills, Great Sandy Desert, Aug 1872, George 14646 (CANB,PERTH); Kimberley District, in 1887,
Nyulasy [MEL 1554131] (MEL). Northern Territory. Tanumbirini Stn, May 1947, Blake 17620 (BRI); SE corner
of entry to Lake Surprise, Lander River, Mar 1973, Maconochie 1669 (BRILDNA,NT); Tanami Desert, Lake
Surprise, Jun 1982, Gibson [NT 69629] (NT); 40 miles [64 km] N of Wauchope, Aug 1956, Lazarides 5845 (NT);
1 mile [1.6 km} S of Elkedra H.S., Oct 1956, Chippendale 3104 (AD,BRI,MEL); About 4 km N of central Mt
Stuart, Jun 1972, Maconochie 1415 (NT), 14 miles [22.4 km] NW of Aileron, Aug 1962, Nelson 417 (NT).
Queensland. GREGORY NorTH DISTRICT: 50 miles [80 km] SE of Camooweal, Dec 1947, Everist 3388 (BRI); 28
km E of Urandangie, May 1985, Neldner 1979 & Stanley (BRI); 2 km SE of Poodyea Point, Toko Ra., Sep 1977,
Purdie 1083 (BRD.

Distribution and habitat: Common species in Central Australia, from Great Sandy Desert,
Western Australia, to Toko Ranges, western Queensland (Map 2); occurs in sandy soll
on ranges, sandy ridges, sand dune and sand plains in deserts and semi deserts.

Z. albiflora is distinguished from closely related species by its erect, densely hairy
stems, broad hairy leaflets and broad congested bracts, the latter and stipules maturing
with reddish coloured nerves. This species is very similar to 2. prostrata. At times it Is
difficult to differentiate between the two species because of the presence of intermediate
forms. However, in Z. prostrata the stems are prostrate and the bracts are usually
distinctly separated, smaller and with a prominent spur, and the fruits are usually much
exserted, prominently reticulate veined and bristly.

Note: Z. albiflora Mohl. and Z. nervata Mohl. have been synonymised here because the
types are from the same taxon. The former is accepted for the species as the type of Z.

32 Austrobaileya 3(1): 1989

neryata 1s not covered by Mohlenbrock’s description of the species he described as Z.
nervata and is not conspecific with the other specimens he cited for it. Most plants
previously called Z. nervata Mohl. are therefore treated as belonging to a new species
we have named Z. prostrata.

16. Zornia prostrata S. Reyn. & Holland species nova, Z. albiflorae Mohl. et Z.
muellerianae Mohl. primo aspectu maxime similis quoad formam foliorum et
bracteorum. A Z. albiflorae caulibus prostratis et bracteis angustioribus plerumque
distinctis, differt. A Z. muellerianae imprimis foliolis, stipulis et plerumque bracteis
eglandulosis differt. Typus: Queensland. BURKE District: About 40 km W of
Doomadgee Aboriginal Station, Burketown to Wollogorang road, 6 May 1974, R.
Pullen 9095 (holo: BRI; iso: CANB).

[Z. nervata auct. non Mohl.: Mohlenbrock, Webbia 16: 130-131, f. 32, 88 (1961)
quoad descriptionem et distributionem.]

Herbs with prostrate, few-branched, epunctate, pubescent to glabrous stems 10-90 cm
long; hairs when present usually long, reflexed and dense. Petioles 5-25 mm long; leaflets
elliptic or ovate, rarely narrowly elliptic, 0.9-3 x 0.4-1.5 cm, obtuse or subacute and
mucronate at apex, obtusely suboblique at base, glabrous or puberulous, eglandular or
rarely sparsely pellucid-dotted on lower surfaces; lateral nerves prominent, sometimes
reddish coloured when dry, reticulate venation obscure; margins ciliate (especially in
young leaves); petiolules 0.5-1 mm long, hairy. Stipules narrowly elliptic or rhombic to
subovate or subfalcate, 0.4-2 cm long, 1-3 mm wide, acute or acuminate at apex, spurred
at base, 5—-8-nerved, usually epunctate: spur 1-8 mm long. Inflorescences 5-19.5 cm
long, (4-)8-21-flowered with bracts overlapping about 1/4 their length initially, later
separated (internodes 1/4-3 times as long as bracts); peduncles 1.5-7.5 cm long; bracts
ovate or elliptic, 0.6-1.8 cm long, 2.5-6.5 mm wide, acute at apex, with small or
prominent spur at base, 4—6-nerved, reticulate veined, ciliate, epunctate or with few
pellucid dots; spur 0.5-6 mm long, obtuse or acute. Calyx 3.5-5 X 1.5-2 mm: petals
yellow or white, standard 3.5-15 mm long. Fruits 2-5(-6)-articulate; articles 2-3.5 x
1.5-2.5 mm, finely hairy and with hairy bristles, with prominent reticulate venation
usually the same colour as the rest of the articles: bristles 0.5-1 mm long, finely hairy.

Z. prostrata is distinguished by the long prostrate stems with few branches, erect
inflorescences arising from axils of erect petioles and prostrate stems, usually separated
bracts, broad leaflets, hairy long bristly fruits with hairy bristles, and articles with
reticulate venation the same colour as the rest of the articles. It is at first sight very
similar to Z. albiflora Mohl. and Z. muelleriana Mohl. having similar leaflets and bracts,
and is no doubt very closely related to both these species. It differs from the former in
its prostrate habit and usually narrower separated bracts, and from the latter in the
absence of dense pellucid dots on leaflets, stems, stipules and bracts, and also in the
longer petioles which are usually longer than the leaflets, and in the usually broader

obtuse leaflets.

Note: Z. prostrata includes plants previously called Z. nervata Mohl. See discussion
under Z. albiflora above.

Etymology: The specific epithet refers to the prostrate stems of the species.

The species varies considerably in shape and size of leaflets and especially the
bracts. Two varieties are recognisable:

Flowers 4-9.5 mm long. Bracts 6-11 mm long; spur usually short or rudi-
mentary, 0.5-3 mm long. Inflorescences (4-)8-21-flowered. Fruits 2-
Gra THC OIA le ahh gen a a lon Pt in ete ce, var. prostrata
Flowers (10-)14-17 mm long. Bracts (11-)15-18 mm long; spur elongate, 5—
mm long. Inflorescences 4-9-flowered. Fruits 2- or
POOPIE ie Ts cholate ive nie fae eels Ey Pads, macrantha

Aornia prostrata var. prostrata

Distinguishing characters are as set out in the key above.

Reynolds & Holland, Zornia 33

Stems hairy with usually long reflexed hairs, rarely subglabrous. Leaflets usually broadly
ovate or elliptic, glabrous or sparsely hairy; lateral nerves prominent, sometimes drying
reddish coloured. Stipules narrowly elliptic to subovate, 4-10 X 1-3 mm; spur 1-3 mm
long. Inflorescences (4—)8-21-flowered, bracts overlapping to separated (internodes 1/4-
3(-4) times as long as bracts); peduncles 1.5-4 cm long. Bracts very variable, elliptic or
ovate, 6-11 x 2.5-5 mm, usually with prominent reddish coloured reticulate venation
when old; spur 0.5-—3 mm long.

Selected specimens: Western Australia. King Edward River crossing on road to Mitchell River Stn, May i975,
Symon 10249 (AD); 73 km W of Wyndham about 13 km S of Paradise Pool on Ernest River, Mar 1978, Lazarides
8634 (CANB,NT,PERTH), Cambridge Gulf, in 1886, Lucomus (MEL); 8 km SE of Kununurra, Mar 1978,
Paijamans 2409 (CANB). Northern Territory. Boggy Plain Ck near J abiru, Jul 1973, Adams 3050 (BRI,CANB,NT);
Cox River Stn, Jul 1977, Latz 7216 (NT); Pine Creek, Mar 1961, Chippendale 7586 (BRI,MEL,NT); Little Lagoon,
Groote Eylandt, Apr 1948, Specht 187 (AD,BRI,MEL);,; Keep River National Park, Feb 1981, Dunlop 5810
(CANB,DNA); Settlement Ck, Mar 1922, Brass 134 (BRD). Queensland. BURKE DisTRICT: Sweers Island, Henne
[MEL 1554122] (MEL); Old Corinda Outstation about 40 km W of Doomadgee Aboriginal Station, Burketown
to Wollogorang Rd, May 1974, Pullen 9095 (BRI,CANB); Iffley about 80 miles [128 km] SSE of Normanton,
May 1954, Everist 5378 (BRI). MircHELt District: Jericho, Mar 1946, Clemens [AQ309878] (BRI); approx 31
km N of Shirley H.S. on Aramac-Torrens Ck Rd, Jun 1977, McDonald 2621 (BRI).

Distribution and habitat: Common along the coast of northern Australia, from Broome,
Western Australia, to Gulf of Carpentaria and extending to central Queensland, including
offshore islands (Map 11); occurs usually in sandy soil in undulating country, on creek
levees and scree slopes, in open woodlands and grasslands.

Var. prostrata varies considerably throughout its range in attributes of leaflets,
stipules and bracts. Usually plants from the islands have very small, separated bracts,
yet some plants have wide bracts e.g. collections from Berry Springs, Northern Territory,
not unlike those of Z. albiflora, differ from that species in their prostrate habit. An
isolated group of plants from Western Australia also has pellucid dots on leaflets, stipules
and bracts, but the glands are inconspicuous and few and their presence is inconsistent,
varying within the same individual.

Zornia prostrata var. macrantha S. Reyn. & Holland var. nov. a var. prostrata S. Reyn.
& Holland bracteis et floribus magnis (fere duplo majoribus) et leguminibus
plerumque biarticulatis differt. Typus: Northern Territory. Katherine Gorge above
Edith Falls (14°11’S, 132°14’E), 25 February 1982, S. King 59 (holo: DNA).

Distinguishing characters are as set out in the key above.

Stems trailing, glabrous. Petioles 1.1-2.1 cm long; leaflets elliptic or ovate, 1.4-2.8 X
0.7-1 cm, obtuse at both ends, glabrous except ciliate margins, discolourous, nerves
indistinct. Stipules narrow and elongate, elliptic or rhombic, (1-)1.4-2 cm long, 2-2.5 mm
wide, acuminate at apex, with elongate spur at base; spur 6-8 mm long, acuminate or
obtuse. Inflorescences with separated, rarely overlapping bracts (internodes 1.5-2.5(-3.5)
times as long as bracts); peduncles 2.5-7.5 cm long; bracts ovate, (1.1-)1.5-1.8 cm long,
(4.5-)6-6.5 mm wide, acute at apex, usually with a very prominent spur at base, 5-
nerved, obscurely reticulate veined, rarely sparsely pellucid-dotted; spur 5-6 mm long,
acute or obtuse. Standard |.4-1.7 cm long.

Selected specimens: Northern Territory. Waterfall Ck, 2 miles [3.2 km] above falls, Apr 1969, Byrnes 1516
(CANB,NT); 23 miles [36.8 km] ESE of El Sharana Mine, Feb 1973, Lazarides 7837 (CANB,NT,PERTH);
Katherine Gorge above Edith Falls, Feb 1982, King 59 (DNA).

Distribution and habitat: Known only from Northern Territory (around Waterfall Creek
and Edith Falls) (Map 12); occurs in sandy soil on gentle slopes and tops of sandstone
escarpments.

Var. macrantha is easily recognisable by the large, ovate, separated bracts and
large flowers. The large broad bracts with prominent basal spur, present on some plants,
is very distinctive and further collections may indicate this variety is worth recognising
as a distinct species. Because of the presence of intermediate forms we are keeping it
under the very variable Z. prostrata for the present.

Etymology: The varietal epithet refers to the large flowers of this variety.

34 Austrobaileya 3(1): 1989

17. Zornia oligantha S. Reyn. & Holland species nova prope Z. prostratae S. Reyn. &
Holland sed inflorescentiis paucifloris (floribus 2-8 raro-12), bracteis ovatis
distinctis leguminibus glabris et articulis parvulis differt. Typus: Northern Terri-
tory. Smith Point, Cobourg Peninsula, about 140 km NE of Darwin, (11°O7’S,
132°08’E), 5 April 1977, R. Pullen 10617 (holo: CANB:; iso: CANB,DNA).

Prostrate herbs with usually sparsely hairy, slender stems to 31 cm long. Stems, petioles
and peduncles with sparse, antrorse, appressed or spreading hairs, epunctate. Petioles {-—
2.3 cm long; leaflets ovate to elliptic, (1-)1.7-1.9(-3) x (0.4-)0.6-1 cm, acute at apex,
obtuse and suboblique at base, glabrous or lower surfaces hairy with scattered appressed
hairs, sparsely pellucid-dotted below; lateral nerves oblique, obscure; petiolules to 1 mm
long, villous. Stipules narrowly elliptic or + triangular, 9.5-15 x 1-2 mm, acuminate at
both ends or spurred or truncate at base, 5-nerved, glabrous or with few hairs on margins;
spur |-3.5 mm long. Inflorescences 2.5~4.5(-12) cm long, 2-8(-12)-flowered with usually
very widely separated bracts (internodes 1-2.5 times as long as bracts); peduncles 1.5-
3.5 cm long; bracts broadly ovate, 6-9.5 x 3.5-4.5 mm, acute at apex, with short spur
at base, 5-nerved, ciliate, glabrous or with few appressed hairs, obscurely reticulate
veined, epunctate or with fine pellucid dots: spur 0.5-2.5 mm long, acute, rounded or
truncate. Calyx 3.5 mm long; petals yellow; standard to 9 mm long. Fruits (3 or) 5-7-
articulate, exserted; articles 1.5-2 X 1.5~1.8 mm, finely hairy (when young) to glabrous,
obscurely reticulate veined and sparsely covered with small yellow glands: usually with
few small bristles mostly at periphery, or non bristly; bristles to 0.5 mm long, glabrous
or finely hairy.

selected specimens: Northern Territory. Airstrip, Smith Point, 11°08’S, 132°09’E, Oct 1968, Byrnes 1084 &
Maconochie (AD,NT), Smith Point, Cobourg Peninsula, about 140 km NE of Darwin, Apr 1977, Pullen 10617
(CANB,DNA); Darwin, Apr 1964, Letts (NT); ditto, Jun 1964, Nelson 1029 (BRENT).

Distribution and habitat: Known only from Cobourg Peninsula and Darwin, Northern
Territory, but is common about Darwin (Map 7); occurs usually in sandy soil in lawns,
old dunes and beach ridges.

Z. oligantha is characterised by the usually few-flowered, fragile inflorescences,
prostrate slender stems; ovate, acute, thin leaflets; widely separated bracts and compar-
atively large flowers, and also by the small glabrous scarcely bristled or non bristly
articles of fruits.

It is closely related to and resembles Z. prostrata S. Reyn. & Holland and Z.
cantonensis Mohl. It differs from the former in the thin, ovate, acute leaflets with obscure
nerves and pellucid dots, also in the few-flowered (up to 8(-12)-flowered) inflorescences
and glabrous fruits with small articles. The leaflets in Z. prostrata are obtuse, prominently
nerved and usually eglandular; the inflorescences are up to 21-flowered and fruits are
hairy. From Z. cantonensis it differs in the 3- or 4- (or 5)-articulate, glabrous, scarcely
bristled or non bristly fruits. In Z. cantonensis the fruits are 5—7-articulate, hairy and
bristly.

Etymology: The specific epithet refers to the few-flowered inflorescences.

Imperfectly known taxa
Zornia sp. (1)

Herbs to 40 cm high with erect, glabrous, obscurely pellucid-dotted stems. Petioles 1.2~
1.7 cm long; leaflets narrowly elliptic, 1.7-3.5 cm long, 1-2.5 mm wide, acute at apex,
acute or obtuse at base, glabrous, conspicuously pellucid-dotted, usually drying blackish;
petiolules to 1 mm long. Stipules narrowly elliptic, 6-8 * 1-1.5 mm, acuminate both
ends, 5-nerved, pellucid-dotted; spur 1-1.5 mm long, acute. Inflorescences 3-12.5 cm
long, 2-9-flowered, with separated bracts: peduncles 1.7-3.5 cm long; bracts narrowly
elliptic, 7-10 X 2-2.5 mm, acute at both ends, 5-nerved, obscurely reticulate veined,
pellucid-dotted; spur 1-2.5 mm, acute or acuminate. Calyx 3-4 mm; petals yellow with
red lines; standard to 9 mm long. Fruits (2- or) 3- or 4-articulate: articles 25-3 x 1.7-—
2 mm, finely hairy, epunctate, covered with finely hairy bristles: bristles to 1 mm long.

Specimens examined: Queensland. Cook District: 19 km S of Paimer River Crossing, Peninsula Development
Rd, | km N of Cook shire boundary, 16°15’S, 144°43’E, Mar 1987, Clarkson 6642 & McDonald (BRI).

Reynolds & Holland, Zornia 35

Distribution and habitat: Known only from the above collection from Queensland; occurs
in woodlands among grasses.

Note: This specimen is not unlike an unnamed specimen at BRI from Papua New
Guinea namely Henty (NGF14337) from Leron River, Morobe District (6°20'S, 146°25°E),
Sep 1961, differing only in having fewer articles in the fruit (Henty has 5-7-articulate
fruits). They are probably both closely related to Z. papuensis Mohl. which, according
to the protologue description, differs from them in having glabrous, 5-7-articulate (as in
Henty) fruits but with glabrous bristles.

Zornia sp. (2)

Stems glabrous. Petioles 1.5 cm long; leaflets narrowly ovate, 2.7-4.5 cm long, 3-9 mm
wide, acute and mucronate at apex, glabrous, sparsely pellucid-dotted, lower surfaces
sparsely hairy; petiolules to 1.5 mm long. Stipules narrowly elliptic, ca 8 X 1.5 mm,
acuminate at both ends, 5-nerved, sparsely pellucid-dotted with yellow dots; spur 2.5
mm long. Inflorescence 4.5-11.7 cm long, 9-20-flowered with bracts overlapping especially
towards apex (internodes (0.5—)1.5-2.5 times as long as bracts); peduncles 1.3-4 cm long;
bracts narrowly elliptic, 8-9 x ca 2 mm, acute at apex, with or without a small spur at
base, 5-nerved, usually pellucid-dotted with yellow dots, sparsely hairy, spur to | mm
long, obtuse. Standard 5~7 mm long. Fruits 4- or 5-articulate; articles subglobose, ca 2
x 1.5 mm, finely hairy, rarely with few small glands, bristly; reticulate venation fine,
reddish coloured: bristles slender, same colour as the reticulate venation, to | mm long,
finely hairy.

Specimen examined: Queensland. NoRTH KENNEDY District: King Ranch, Tully District, Mar 1979, Middleton
[AQ330876] (BRD).

Distribution: Known only from the above collection.

This plant is a good match for plants from Brazil identified as Z. /atifolia Smith
and agrees quite well with Mohlenbrock’s description of that species. Until more
collections are available for study we cannot be certain of its identity. If the plant 1s Z.
latifolia Smith, then it is probably an escape from cultivation.

Acknowledgements

The authors are grateful to the Directors of the following herbaria (for the loan
of their herbarium material and types) - AD, CANB, DNA, G, L, MEL, NT, PERTH
and PR. They also extend their thanks to Les Pedley for the latin diagnoses, Rod
Henderson for his comments on the manuscript, David Halford and Brian Hacker for
their assistance and Dr Judy West for helping to search for the type of Z. dyctiocarpa
DC. at K, BM, G.DC and G. The senior author also appreciates the financial support
given by the Australian Biological Resources Study, Federal Department of The Arts,
Sport, The Environment, Tourism and Territories.

References
BAILEY, F.M. (1913). Comprehensive Catalogue of Queensland Plants. Brisbane: Government Printer.
BENTHAM, G.W. (1864). Zornia. Flora Australiensis 2: 228-229. London: Lovell Reeve & Co.
DOMIN, K. (1962). Zornia. Bibliotheca Botanica 89: 208-211, t. 25.
MILNE-REDHEAD, E. (1953-55). Zornia in Tropical Africa. Boletum da Souldade Boteriana, 27-29: 79-104.
MOHLENBROCK, R.H. (1961). A monograph of the Leguminous genus Zornia. Webbia 16: 1-141.
MOHLENBROCK, R.H. (1965). Zornia. Journal of South African Botany 31(2): 100-109.

VAN MEEUWEN. M.S.. NOOTEBOOM, H.P. & VAN STEENIS, C.G.G.S. (1961). Preliminary Revisions of
some Genera of Malaysian Papilionaceae |. Retmwardtia 5(4): 419-456.

VERDCOURT, B. (1979). A manual of New Guinea Legumes: Zornia. Papua New Guinea, Depariment of
Forests. Botany Bulletin 11: 375-380.

Accepted for publication 2 May 1989

36 Austrobaileya 3(1): 1989

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y, &
@
2
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i
t\ \ ter
|
{ ar ( ;
ro af — f
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Maps 1-6. |. BZ. muelleriana subsp. muelleriana, @ Z. muelleriana subsp. congesta. 2. WZ. floribunda, @ Z.
albiflora, 3. Z. dyctiocarpa var. dyctiocarpa 4, @ Z. dyctiocarpa var. filifolia. 5. @ Z. muriculata subsp. muriculata.
6. @ Z. muriculata subsp. angustata.

Reynolds & Holland, Zornia 37

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9 10
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Maps 7-12 7. ® Z. adenophora, @ Z. oligantha, 8. WZ. chaetophora, ® Z. stirlingii. 9. W@ Z. areolata, ® Z. pallida.

10. @ Z. disticha, @ Z. maritima. 11. WZ. prostrata var. prostrata. 12, @ Z. prostrata var. macrantha, @ Z.
prostrata—-miuelleriana intermediates.

ow at th Ho th i Hw

ICI I A Ha tl a th th a a a Mat i

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38 Austrobaileya 3(1): 1989

Map 13. A Z. acuta, @ Z. pedunculata, @ Z. ramosa.

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Austrobaileya 3(1): 39-44 (1989) 39

TWO NEW SPECIES OF EUCALYPTUS (MYRTACEAE)
FROM CENTRAL QUEENSLAND

A.R. Bean
PO Box 397, Nambour, Qld 4560
and M.I.H. Brooker

CSIRO Division of Plant Industry, GPO Box 1600, Canberra, ACT 2601

Summary

Eucalyptus xanthope, a new bloodwood, and Eucalyptus decolor, a new ironbark, are described and illustrated.
Both species have a restricted distribution in sub-coastal central Queensland.

Taxonomy

Eucalyptus xanthope Bean & Brooker, species nova affinis Eucalypto intermediae R.
Baker a qua cortice interiore aspero flavescenti, ramulis laevibus, plantulis
pubescentibus tantum ad nodum tertium, foliis adultis angustioribus et florescentia
dissimili differt. Typus: Queensland. PORT CURTIS DISTRICT: Bruce Highway,
0.5km south of Glen Geddes siding, 23°02’S, 150°16’E, 14 February 1988, A.R.
Bean 753 (holo: BRI).

A tree to 20 m tall with flaky bloodwood bark on the trunk and branches greater than
5 cm diameter. Underbark yellow; newly exposed bark yellow and often producing a
blotchy appearance, fading slowly to brown then grey; terminal branches (less than c. 5
cm diameter) smooth-barked. Cotyledons reniform, to 9 mm X 12 mm. Seedling leaves
opposite for many pairs, shortly petiolate, not peltate, narrow lanceolate, to 11 cm X
1.6 cm, concave, discolourous. Hairs present on leaves and stems until the 2nd or 3rd
pair of leaves; subsequent leaves and stems glabrous. Juvenile leaves glabrous, similar
in shape, but somewhat larger than seedling leaves. Adult leaves alternate, petiolate,
lanceolate, up to 19 cm X 2.5 cm, strongly discolourous, main lateral veins set at a wide
angle to the midrib and terminating at an intramarginal vein which 1s removed from
the leaf edge by a single line of areoles; venation densely reticulate; oil glands small,
scattered, appearing singly within the areoles if present. Inflorescences apparently terminal,
compound. Peduncles angular, to 10 mm long, surmounted by 7 buds. Pedicels up to 6
mm long. Buds ovoid to clavate, smooth, to 8 mm X 5 mm. Opercula low hemispheric,
both shed at anthesis. Stamens white, fully inflexed, all fertile. Fruits pedicellate, ovoid,
grey or brown, speckled or uniform in colour, to 18 mm X 14 mm, disc descending.
Valves deeply enclosed. Seed brown with a long terminal wing. Fig. 1.

Specimens examined: Queensland. PORT Curtis District: 11.5 km from Marlborough towards Rockhampton,
May 1987, Bean 566 (BRI); 1.5 km west of Glen Geddes siding, between Rockhampton and Marlborough, Oct
1985, Bean 333 (BRI,NSW); Glen Geddes about 25 mls [40 km] N.W. of Rockhampton, Sep 1968, Everist 8008
(BRI); 0.3 km south of Glen Geddes siding, Oct 1987, Brooker 9772 & Bean (AD,BRI,CANB,MEL,NSW); Bruce
Highway, 0.5 km south of Glen Geddes Siding, 23°02’S, 150°16’E, Feb 1988, Bean 753 (BRI); 11 km from
Yeppoon towards Rockhampton, Dec 1985, Bean 342 (BRI,NSW).

Distribution and habitat: Eucalyptus xanthope has a quite restricted distribution between
Rockhampton and Marlborough. The main occurrence is along the Bruce Highway
between Canoona and Marlborough, and there is a population on the Rockhampton-
Yeppoon road (Map 1). It grows on low hills in shallow soils. Its occurrence coincides
with outcrops of serpentinite, an uncommon type of ultramafic rock, which contains a
high proportion of magnesium (as silicate), usually 30-40% (expressed as MgO). Ultramafic
rocks, including serpentinite, produce notoriously infertile soils the world over (Beadle
1981). The distribution and origins of ultramafic rocks in the Rockhampton area is
discussed at length by Murray (1969).

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40

Austrobaileya 3(1L); 1989

cm

Fig. 1. Eucalyptus xanthope: A. buds. B. fruits.

39

Bean & Brooker, Eucalyptus 4

Associated eucalypt species are EF. fibrosa F. Muell. subsp. fibrosa, E. papuana F.
Muell. and E. erythrophloia Blakely. Other prominent associated species include Racos-
perma leptostachyum (Benth.) Pedley, Triodia sp. and Hakea trineura F. Muell.

Flowering period: February—April.

Affinities: The bloodwood bark, the transverse leaf venation, the apparently terminal
inflorescence and the mop-type stigma (Boland & Sedgely 1986) confirm FE. xanthope as
belonging to the informal Z. subgenus Corymbia (Pryor & Johnson 1971), while the
winged seeds indicate its placement in the informal E. section Rufaria -(E. series
Terminaliptera Maiden (Chippendale 1988)). It is morphologically close to E. intermedia
R. Baker from which it differs in the rough yellow underbark, smooth terminal branches,
seedlings hairy to the third node only, narrower adult leaves and different flowering
period. It also resembles that species in the commonly speckled, ovoid fruits and the
smooth ovoid buds. The two species are readily separable in the field and Table 1 lists
the morphological differences.

Table 1. Comparison of E. intermedia and E. xanthope

Character E. intermedia E. xanthope
Rough-bark colour grey outside, brown brown to grey
underneath outside, yellow underneath
Rough-bark extent persistent throughout terminal branches smooth
Canopy dense sparse, somewhat
pendulous
Length/breath ratio 3-5 5-9
for adult leaves
Flowering period Nov-—Jan Feb-Apr
Seedlings hairy throughout glabrous beyond Node 3

In addition to the above, £. intermedia and E. xanthope have different ecological
requirements. &. intermedia, which requires a relatively high rainfall, is not common in
the Rockhampton area (sites include Mt Archer and Yeppoon). It 1s unlikely that it
would be able to tolerate the hostile edaphic conditions imposed by serpentinite (see
above) or the somewhat lower rainfall experienced in the Rockhampton—Mariborough
area.

E. brachycarpa D. & S. Carr is another similar species. E. brachycarpa differs
from E. xanthope due to its grey or reddish bark, smaller adult leaves, narrower seedling
leaves and urceolate fruits.

Etymology: The name refers to the blotchy yellow rough bark. The older, outer dead
bark is shed unevenly in patches revealing the fresher yellow underbark. (Greek xanthos-
yellow and ope—hole).

Eucalyptus decolor Bean & Brooker, species nova, arbor (“ironbark’’) affinis Eucalypto
paniculatae Smith a qua cortice trunci atrocineraceo, ramulis laevibus, folzis
plantularum angustioribus et foliis adultis et fructibus parvioribus differt. Typus:
Queensland. PORT CURTIS DISTRICT: Mt Castletower, 24°10’S, 151°17’E, 9 August
1986, A.R. Bean 480 (holo: BRI).

A tree to 25 m tall with dark grey, furrowed tronbark on the trunk and largest branches;
small and medium sized branches smooth, white or pink to rich coppery. Cotyledons
reniform, to 4mm X 7 mm. Seedling leaves petiolate, opposite for 3 or 4 pairs, narrow-
lanceolate, to 9.2 cm X 1.8 cm, green, discolourous, tips obtuse, with a short point.
Juvenile leaves alternating, petiolate, narrow-lanceolate, to 11.5 cm x 1.8 cm, discol-
ourous, dark green above, much paler below. Adult leaves alternate, petiolate, lanceolate,
to 12 cm X 1.8 cm, strongly discolourous, dark green above, much paler below, not

4? Austrobaileya 3(1): 1989

glossy; side veins terminating at an intramarginal vein which is removed from the leaf
edge by a single line of areoles; reticulation dense; oil glands few, obscure, discrete within
the areoles or apparently associated with veinlets. Inflorescences apparently terminal,
compound. Peduncles angular, to 11 mm long, surmounted by 7 buds. Pedicels angular,
to 5 mm long. Buds diamond-shaped, to 8 mm X 4 mm. Operculum conical or rostrate,
narrower than the hypanthium at the join. Stamens white, inflexed, outer filaments
sterile. Fruits pedicellate, ovoid to hemispherical, to 6 mm X 6 mm, staminophore
broad, prominent. Disc obscure, valves 4, rarely 5, enclosed, slightly below or at rim
level. Fig. 2.

Specimens examined: Queensland. Port CuRTisS District: Mt Castletower, Aug 1986, Bean 480 (BRI); Oaky
Creek, near Mt Castletower, Oct 1987, Brooker 9766 & Bean (AD,BRILCANB,MEL,NSW); slopes of Mt Stanley,
west of Bororen, Feb 1988, Bean 749, 750 (BRI,NSW). BURNETT DISTRICT: about 2 km from Coongara Rock
campsite, towards Biggenden, 25°39’S, 152°00’E, May 1988, Bean 810 (BRI,NSW).

Distribution and habitat: Eucalyptus decolor is known only from the Many Peaks Range
south of Gladstone, and in the ranges south of Biggenden. (Map 2). It grows on hilly
to mountainous terrain between altitudes of 300 and 600 metres. Soils vary from white
sands to grey loams derived from granite and are always shallow. Associated species
include EF. trachyphloia F. Muell., &. major (Maiden) Blakely, E. acmenoides Schauer,
and EF. maculata Hook., and at the highest altitudes, EF. andrewsii Maiden.

Flowering period: unknown; flowers have been collected in October and February.

Affinities: The ironbark bark, early shedding of the outer operculum, inflexed stamens,
the presence of staminodes and the pinhead stigma place this species in the informal E.
series Paniculatae (Pryor & Johnson 1971) (E. series Rhodoxyla (Blakely) Chippendale

: @ Marlborough Byfield

—23°S

YEPPOON

O 15Skm
a

ROCKHAMPTON YEE

150 E

Map 1. Distribution of Eucalyptus xanthope.

Bean & Brooker, Eucalyptus 43

(Chippendale 1988)). This series comprises about 15 species distributed from southern
New South Wales to the Tropic of Capricorn in Queensland. Of the members of this
group, E. decolor is perhaps closest to E. paniculata, from which it differs by the darker
grey bark on the trunk, smooth outer branches, smaller leaves, narrower seedling leaves
and smaller fruits. Both species are unusual in possessing strongly discolourous adult
leaves. Another similar and related species is E. melanoleuca S.T. Blake. E. decolor
differs from both of these species by the characters outlined in Table 2. The strongly
discolourous adult leaves distinguish E. decolor from all other ironbarks indigenous in
Queensland.

Table 2. Comparison of E. decolor, E. paniculata and E. melanoleuca

Character E.. decolor E. paniculata E. melanoleuca
Rough bark smaller branches rough-barked smaller branches
extent smooth throughout smooth
Seedling leaves to 9.2 X 1.8 cm to 8.0 X 5.0 cm to 5.0 X 1.0 cm
Adult leaves discolourous, discolourous, concolourous,

to 12.0 X¥ 1.8cm to 15.0 X 2.2 cm to 15.0 * 2.5 cm
Fruit hemiusperhical obconical to ovoid, to 6

to ovoid, to pyriform, to x 5 mm

6 X 6 mm 9X 7mm

O

f

1

‘

\

fy

"
7 N ~~

8
., V

1

~S \

0
\
»
. N

\!
J

Fig. 2 Eucalyptus decolor. A. buds. B. fruits.

44 Austrobaileya 3(1): 1989

Ktymology: [The name refers to the adult leaves, which are pale on the under-surface
(Latin decolor - discoloured or faded).

Acknowledgements

We are grateful to Jan Sked for preparing the drawings, and to David Champion
for providing geological information.

References
BEADLE, N.C.W. (1981). The Vegetation of Australia. Cambridge: Cambridge University Press.

BOLAND, D.J. & SEDGELY, M. (1986). Stigma and style morphology in relation to taxonomy and breeding
systems in Lucalyptus and Angophora (Myrtaceae). Australian Journal of Botany 34: 569-584.

CHIPPENDALE, G.M. (1988). Eucalyptus, Angophora (Myrtaceae). In George, A.S. (ed.), Flora of Australia Vol.
19. Canberra: Australian Government Publishing Service.

MURRAY, C.G, (1969). The Petrology of the Ulitramafic Rocks of the Rockhampton district, Queensland.
Queensland Department of Mines, Brisbane, Publication No. 343: {-9.

PRYOR, L.D. & JOHNSON, L.A.S. (1971). A Classification of the Eucalypts. Canberra: Australian National
University Press.

Accepted for publication 7 December 1988

‘AN 152°E

caine 0 3Okm
24°S
A.
A
Miriam
Vale
»>sBUNDABERG

Biggenden IGH

Map 2. Distribution of Eucalyptus decolor.

Austrobaileya 3(1): 45-49 (1989) 45

A NEW SPECIES OF MARSDENIA R. BR.
(ASCLEPIADACEAE) FROM EASTERN AUSTRALIA

J.B. Williams

Botany Department, University of New England, Armidale N.S.W. 2351

Summary

Marsdenia liisae J. Williams, a new species from north-eastern New South Wales related to 4. glandulifera C.
White of Queensland, is described and figured. Notes on habitat, distribution and conservation status are given.

Introduction

This paper complements the series Studies on the Australasian Asclepiadaceae
(Forster 1987; Forster & Thongpukdee 1988) by providing a name and description for
the sole New South Wales Marsdenia species that is known to be distinct but which is
as yet unnamed. Flowering material of this species was first collected by the writer in
the New England National Park, west of Bellingen, New South Wales, in February 1974
and later on the Nightcap Range north of Lismore in December 1975, during a survey
of rainforest climbing plants of New South Wales (Williams & Harden 1980). It is
similar to M. rostrata R. Br. but has much larger flowers and several leaf and other
floral differences. A further excellent collection of flowering material was made near
Tyringham, New South Wales, in February 1976 by Ms M.L. Lapinpuro, who included
a partial description and line drawings of the species in her thesis on the Asclepiadaceae
of north-eastern New South Wales (Lapinpuro 1976). Subsequently the writer published
a short description in flora format as Marsdenia “Unnamed species” (Williams 1984).
Since 1976 several more collections have been made but only three further flowering
populations have been found. Curiously, examination of collections at NSW, BRI and
MEL revealed no material of this species, apart from the above-cited collections by the
writer and Lapinpuro.

The floral description of the species is based partly on material preserved in spirit
and on fresh material, as well as on herbarium specimens.

Marsdenia liisae J. Williams, species nova affinis M. glanduliferae C. White sed pedicillis
tenuioribus longioribusque (9-25 mm longis), pedunculis 8-40 mm longis, lobis
corollae in alabastro + rectis, apice leviter spiraliter convoluto, sepalis ciliatis et
lobis corollae ciliatis, corollae fauce et in tubo breviter pubescenti; parte basalt
coronae alata, cucullata carnosa et parte supera subtereti vel compressa incurva,
folliculis angustatis, lanceolatis-fusiformibus, 10-13.5 cm X 2-2.5 cm, et glandibus
2-16, elevatis, minoribus, ca 0.4 mm longis ad basim costae laminae differt.
Typus: New South Wales. NORTH CoasT SUBDIVISION: 9 km along Deervale loop
road from its western junction with Ebor—Dorrigo road, February 1988, J. Williams
88078 (holo: BRI; iso: K,NSW,NE).

Marsdenia “Unamed species” of J.B. Williams in N.C.W. Beadle, Students Flora
of North Eastern New South Wales, Part 5: 732, t. 322k (1984).

Semi-woody vine to 5 m high. Stems glabrous or sparsely puberulent when young, green
or purplish, twining, 1-7 mm thick with copious milky latex; older stems with pale
fissured corky bark; shoot tips pubescent. Leaves petiolate. Lamina chartaceous, rather
thick, oblong-ovate to oblong, ovate or ovate-deltoid, glabrous or almost so, moderately
dark green and glossy above, pale green below, 4-18 cm long, !.5-8 cm wide; margins
undulate; apex acuminate, often abruptly, to a rather fine point; base deeply to shallowly
cordate or rounded-truncate, rarely broad-cuneate; midvein green or purple, secondary
veins prominent, moderately raised below, 5-9 on either side of the midvein, making
looping interconnections; coarsest reticulate veins distinct, others obscure;
2-16 erect glands ca 0.4 mm long present on the upper surface of the lamina at base of
midvein. Petiole 13-55 mm long, green or purple, terete but mostly grooved above,
slabrous or sparsely puberulent above. Inflorescences simple, lax, pedunculate, 2-8-

A6 Austrobaileya 3(1): 1989

Fig. 1. Marsdenia liisae. A. part of flowering shoot with simple umbels x 0.5. B,C. leaves showing variation in
shape and size X 0.4, D. base of lamina, adaxial view, showing cluster of squamellae X 4. E. flower bud showing
contorted corolla lobes spirally twisted at the apex and each with an abrupt projection at the base X 2. F. corolla
lobe removed from bud (abaxial view) X 2. G. corolla lobe removed from bud (adaxial view) X 2. H. single
flower from above X |. E. single flower from below X |. J. adaxial view of single sepal showing ciliate margins
and basal glands X 3. K. pollinarium X 18. L. base of flower dissected to show staminal corona and long-exserted
bifid style (i = incurved free part of corona segment, h = hooded basal part of corona segment) X 3, M,N. fruits
x 0.5. O. seeds with long coma X 0.7. All drawn from type collection except M (M, Williams 75159); A~D from
dry specimens, E-O from fresh or preserved specimens.

Williams, Afarsdenia litsae 47

flowered umbels: peduncles solitary, interpetiolar, rarely forked, glabrous, 8-40 mm long;
pedicels slender, glabrous, 9-25 mm long. Sepals broad-lanceolate to ovate, obtuse, 3.3-—
6.5 mm long, 2-5.5 mm wide, glabrous except for the ciliate margins, minute glands
(squamellae) 0.3-0.5 mm long scattered in groups of 2 or 3 at base of sepals inside,
numbering ca 20-25 per calyx. Corolla creamy-yellow, rotate, 8-9 mm long, 20-43 mm
diameter; lobes spreading not reflexed, linear-lanceolate, obtuse, twisted near apex, 9-
20 mm long, 2-5 mm wide, glabrous externally, internally shortly pubescent near the
base: margins ciliate, strongly recurved; in bud corolla lobes straight, not spirally twisted
except near the apex, each with an abrupt lobe at the base; tube broad-ovoid, 5.5-8 mm
long, 5-7 mm diameter, glabrous outside, shortly pubescent inside and at the throat;
corolline corona absent. Staminal column 8-16 mm long, 3.3-5 mm diameter; anthers
with an obtuse membranous appendage at the apex; corona greenish, of 5 double
segments, the lower part of each segment 2-3 mm long, adnate to the base of the column,
with a hood-shaped fleshy wing, the upper part free, slender, subterete or somewhat
compressed, incurved, white, 2.3-2.6 mm long. Style-head conical, white, much longer
than the stamens, the apex bifid, exserted up to 4 mm beyond the corolla tube; ovaries
slabrous. Pollinia ascending, without pellucid margins, oblong-ovoid, yellow, ca 0.7 mm
long and 0.3 mm wide; translators slender, 0.4-0.5 mm long; corpusculum dark red, ca
0.35 mm long and 0.3 mm wide. Follicles lanceolate in outline or fusiform, turgid, 10-
13.5 cm long, 2-2.5 cm diameter. Seeds compressed, ovate-lanceolate, 9-11 mm X ca
4.5 mm; coma 30-40 mm long. Fig. 1.

Selected specimens: New South Wales. NorTH Coast: Whian Whian State Forest, junction of Nightcap track
and Gibbergunya Range road, 29 km N of Lismore, Dec 1975, Williams 75159 (flowers, BRISMEL,NE,NSW), 8
km east of Tyringham on Grafton-Armidale road, Feb 1976, L. Lapinpuro s.n. (flowers, BRI,NE,NSW): Bruxner
Park, 9 km NW Coffs Harbour, Dec 1980, Williams 80255 (NE,NSW), Grasstree Ridge, New England Nat. Park,
Feb 1974, Williams 74025 (flowers, BRI,NE,NSW); Black Scrub Ridge, Bellinger River State Forest, Nov_1979,
Floyd 1394 (NE); upper Hastings River ca 30 km NNW of Yarras, Mar 1981, Wiliams 81037 (NE,NSW); Banda
Banda Beech Preserve, Dec 1980, Williams 80263 (flowers, BRILMEL,NE,NSW).

Distribution: The species has been found only in the North Coast district of New South
Wales. The distribution map (Map 1) indicates a concentration in three more or less
disjunct areas, the higher portion of the Nightcap Range north of Lismore, the Dorrigo
Plateau and adjacent ranges, and, further south, the headwaters of the Hastings River.
There are several areas of apparently suitable habitat in Queensland.

Habitat: M. liisae occurs over an altitude range of 300-1050 metres, in sites with a
mean annual rainfall of ca 1400-2000 mm, and on deep soils derived from a wide range
of rocks including rhyolite, basalt and granodiorite as well as argillite and other fine-
srained metamorphics. It has been found in both tall eucalypt forests and in rainforests
of various types including subtropical, warm-temperate and cool-temperate, often along
roadsides. The most common associated trees are the rainforest species Ceratopetalum
apetalum, Doryphora sassafras, Callicoma serratifolia and Nothofagus moorei, and the
eucalypts E. microcorys, E. saligna and E. pilularts.

Phenology: This species flowers from December to February. Fruiting specimens have
been collected in late December and in February.

Notes: Among Australian species Marsdenia liisae most nearly resembles M. glandulifera
C. White and M. rostrata R. Br., especially in the very long + conical bifid style-head
and strongly rotate corolla. It also resembles M. glandulifera in the very large flowers
and the usually large, cordate leaves. It is readily distinguished from M. glandulifera by
the lax, simple umbels with long slender peduncles and pedicels, the ciliate sepals and
petals, the presence of a fine, short tomentum within the corolla tube, at the throat and
on the base of the corolla lobes, the flower buds with + straight not conspicuously
spirally twisted corolla lobes, the large but slender, lanceolate-fusiform (not ovoid)
follicles 10-13.5 cm long and the 2-16 smaller, less conspicuous squamellae at the base
of the lamina. The geographic ranges of M. liisae and M. glandulifera are widely separated
and their habitats are different. Vegetatively (4. liisae might be confused with the larger-
leaved forms of M. rostrata but the leaf apex in M. liisae is finely pointed, the lamina
is thicker and more deltoid and the margins are strongly undulate; the lateral veins make
narrower angles of 30-50° (not 45-75°) with the midvein. The flowers with corollas 2-
4 cm across are far larger than those of M. rostrata and the tomentum in the corolla
tube and throat is different, lacking the thick pointed retrorse hairs and not forming 5
tufts as in that species. The long slender follicles contrast with the broad ovoid follicles
of M. rostrata. The flower buds of M4. Hisae are very distinctive with five abrupt

48 Austrobaileya 3(1): 1989

projections in the lower part. The corona of M. /iisae is also distinctive, each segment
having a hood-shaped, winged adnate basal part and an incurved free upper part. The
umbels of M. /iisae are deciduous, showing no tendency to develop perennial elongated
peduncles as do those of M. glandulifera.

Some variation in flower size and in the form and size of leaves has been noted.

The leaf blades often vary considerably in size and shape on the same plant, with
the upper laminas on flowering shoots being relatively small and not at all cordate at
the base. Specimens from the northern population on Nightcap Range have somewhat
smaller flowers than the others. Rarely two umbels are found on a short common
peduncle.

28 a ~ «628
aoe
OLismore
O
Grafton
30 30
@
e
Oo. ., QYCoffs Harbour
Dorrigo
OArmidale
@
50 km.
eae eee
oKempsey
, @ ’ a
a 8 |
NO Sa

Map 1. Distribution of Marsdenia liisae from herbarium specimens.

Williams, Afarsdenia ftisae 49

Conservation status: The species is presently known from some thirteen populations; all
the collections examined were made in the period 1972-1988. Several of the gatherings
are from single plants and no extensive populations have been found. Since several
collections were made along roadsides it seems that M. /iisae is not disadvantaged by
moderate disturbance. Nine of the populations are within National Parks, Nature
Reserves, or forestry Flora Reserves. On present knowledge its status is assessed as 3RC,
ie. rare but not vulnerable, using the risk coding of Leigh, Briggs and Hartley (1981).

Vernacular name: . liisae has been known as Large-flowered Milk Vine in New South
Wales (Williams & Harden 1980).

Etymology: Named in honour of Ms Luisa Lapinpuro of Armidale, who made an early
collection of this species, gave a preliminary account of its features and has provided

considerable information on the morphology of Australian Asclepiadaceae (Lapinpuro
1976).

Acknowledgements

Mr N. Byrnes kindly provided comments on the distinctiveness of M. liisae when
specimens were sent to BRI in 1976. I thank the Directors of the Queensland Herbarium,
Brisbane, the National Herbarium of New South Wales, Sydney and National Herbarium
of Victoria, Melbourne for providing access to material.

References

FORSTER, P.I. (1987). Studies on the Australasian Asclepiadaceae, Il. A new combination in Gymnema R. Br.
Austrobaileya 2: 401-404.

FORSTER, P.I. & THONGPUKDEE, A. (1988). Studies on the Australasian Asclepiadaceae, III. A new species
of Cynanchum L. and a new name in Marsdenia R. Br. Austrobaileya 2: 451-457.

LAPINPURO, MARJA LIISA (1976). Studies in the taxonomy, ecology and anatomy of the Asclepiadaceae of
north-eastern New South Wales. Unpublished. B. Sc. Hons thesis, Armidale: University of New England.

LEIGH, J., BRIGGS, J. & HARTLEY, W. (1981). Rare or threatened Australian Plants. Australian National
Parks & Wildlife Service Special Publication No. 7. Canberra: Australian National Parks & Wildlife Service.

WILLIAMS, J.B. (1984). Asclepiadaceae. In Beadle, N.C.W., Students Flora of North Eastern New South Wales.
Part V: 719-736. Armidale: University of New England.

WILLIAMS, J.B. & HARDEN, G.J. (1980). Rainforest climbing piants. Armidale: University of New England.
Accepted for publication 2 March 1989

Austrobaileya 3(1): 51-62 (1989) 51

KAILARSENIA TIRVENGADUM EMEND. PUTTOCK
(RUBIACEAE: GARDENIEAE) IN AUSTRALIA

C.F. Puttock

University of New South Wales, Kensington, NSW 2033

Summary

Gardenia jardinei F, Muell. ex Benth., G. ochreata F. Muell. and G. suffruticosa R. Br. ex Benth. are revised and
transferred to Kailarsenia Tirvengadum emend. Putteck. Full descriptions, notes and a key to these species are
provided. Gardenia kershawii Bailey, G. macgillivraei Benth. and G. ochreata var. parviflora F. Muell. are analysed
using multiple regression analysis. These taxa are reduced to synonymy of Kailarsenia ochreata (F. Muell.) Puttock.

Following the revision of Gardenia Ellis and Randia Houst. ex L. in West Africa
(Keay 1958) there has been a piecemeal regional dismemberment of these two large
heterogeneous assemblages. Unfortunately several of the resultant small genera have
been poorly circumscribed (e.g. Tirvengadum & Sastre 1979; Tirvengadum 1983) and
thus the assessment of relationships of some taxa outside local regions has been difficult.
This paper considers four of the five taxa removed from Gardenia by Puttock (1988)
viz G. jardinei F. Muell. ex Benth., G. kershawii Bailey, G. macgillivraei Benth. and G.
ochreata F. Muell. It also accounts for the only other Australian species belonging to
this group, G. suffruticosa R. Br. ex Benth., a Northern Territory endemic.

These species are similar to Gardenia sens. strict. in possessing sympodial branch-
ing, conical stipules and unilocular ovaries with many ovules. They differ, however, 1n
producing monad pollen and polygamous flowers, and in having leaves with intersecon-
dary venation (Hickey 1973). This combination of characters places them in Rothmannia
Thunb. (sensu Bremekamp 1957 and Yamazaki 1970), or in Kailarsenia Tirvengadum.

Tirvengadum (1983) gave only a brief diagnosis of Kailarsenia, distinguishing it
from Rothmannia sens. strict. on the basis of rheophytic habit, tubular 2-lobed persistent
stipules, fasciculate flowers with linear lanceolate or tentaculate calyces, small fruits and
testa cells thickened but without pits or ornaments. This diagnosis provides little accurate
information concerning the type species Kailarsenia tentaculata (J.D. Hook.) Tirven-
gadum and the other five species placed in the genus. The evolution of the rheophytic
habit is considered at best useful at the specific and subspecific level in the Gardenieae
(Robbrecht 1988). Rothmannia sens. strict. has interpetiolar stipules that are short,
triangular and free to the base. In Kailarsenia tentaculata the stipule is an elongated,
chartaceous, conical sheath, terminating with two minute interpetiolar limbs. The expand-
ing bud tears the sheath along two intrapetiolar lines of weakness to form two sheathing
lobes that persist on the stem until the abscission of the leafbases. Inflorescences are
clearly variable within genera of the Gardenieae and even within the six species currently
attributed to Kailarsenia. The inflorescence is basically cymose but the axes may be
reduced to give the appearance of “fasciculate flowers”, as in K. tentaculata. Several
species have been described with or as often having solitary flowers (e.g. K. lineata
(Craib) Tirvengadum and K. godefreyana (O. Kuntze} Tirvengadum). Tirvengadum
described the fruits of Kailarsenia as small. However, at least one species, K. stenosepala
(Merr.) Tirvengadum, has fruits to 2.5 cm long, which is in the second largest fruit-size
class (III) described for the Gardenieae, a class only exceeded in the Gardeniinae by
some species of Rothmannia (Robbrecht & Puff 1986). The majority of the Kailarsenia
species are within Robbrecht and Puffs intermediate (1-2 cm) fruit-size class (J). The
exotestal cells are thickened along the radial and inner tangential walls in K. tentaculata
with the thickening of the inner tangential walls distinctly pitted (= “perforations” of
Robbrecht & Puff 1986, see fig. 20e4). In this species the exotestal cells around the hilum
and the rest of the perimeter of the seed are markedly radially elongated compared with
those on the flat surfaces (see Robbrecht & Puff 1986, fig. 17e-g). These features of the
seed-coat appear to contradict Tirvengadum’s description (without pits or ornaments).

5? Austrobaileya 3(1}: 1989

Hooker (1880) described the flowers of his Gardenia tentaculata as polygamous,
but according to Robbrecht and Puff (1986) and my own observations on limited
herbarium material, they are hermaphrodite. The condition in the other five south-east
Asian species has not been investigated.

The Australian species agree with the two characters that appear to be unique to
Kailarsenia, the nature of the stipule and the seed-coat morphology. In all three Australian
species the stipule is an elongated, chartaceous sheath with minute, terminal, interpetiolar
limbs (figs th,q, 5g). The seed-coat in Gardenia jardinei and G. ochreata is considerably
well developed around the hilum. The inner tangential walls of the exotestal cells of G.
jardinei have almost identical pitting to that found in Kailarsenia tentaculata, while the
pits in Gardenia ochreata are much larger giving a prominent reticulum. The Australian
species also closely agree with other character states found within Kailarsenia: unarmed
trees and dwarf shrubs, ovaries unilocular, fruits many seeded, seeds embedded in
placental mass, pollen 3- or 4-colporate, colleters lanceolate, flowers solitary or in few-
flowered cymes, calyces linear, corolla tubes tubiform, corolla lobes usually shorter than
corolla tube, and anthers and style scarcely exserted beyond the throat of the corolla
tube. In addition, all three Australian species show some degree of polygamy, a character
described for K. tentaculata by Hooker (1880) but as yet unconfirmed in this or other
south-east Asian species. Therefore, on the basis of the essential agreement of vegetative,
floral and fruit characters, and in particular the stipular and seed-coat morphologies, the
three Australian species are now transferred to Kailarsenia. It is necessary to emend
Kailarsenia to include the diversity of the characters of the six original taxa and the
Australian species. Hence, a new and full generic description is provided below.

All Australian species have been observed in the field. Descriptions and meas-
urements are based on living and dried material and are applicable to both. The
terminology used in this paper for bark, leaf venation and floral characters follows
Puttock (1988). The conservation status has been coded using the ranks proposed by
Leigh, Briggs and Hartley (1981).

Taxonomy

Kailarsenia Tirvengadum emend. Puttock. Type: K. tentaculata (J.D. Hook.) Tirvengadum

Dioecious, gynodioecious, or ?monoecious, facultatively deciduous, thornless dwarf shrubs
or smail trees with sympodial branching. Bark layered, subrhytidome deep green. Wood
hard and close grained, not brittle, cream. Leaves opposite or ternate, often unequal,
petiolate; lamina entire; secondary venation camptodromous with intersecondary veins
and reticulate tertiary veins; domatia commonly present in secondary/midvein angles.
Stipules connate, fused into a long sheath with 2 or 3 minute, terminal, interpetiolar
limbs; sheath membranous or chartaceous, splitting with bud expansion to form larger
lobes, usually persisting until leafbase abscission. Numerous lanceolate to cylindrical
colleters present in the basal part between stipule and stem, producing a small quantity
of viscid resin. Flowers in terminal, 2-8-flowered cymes, or solitary. Flowers herma-
phrodite, or functionally male and female and dimorphic, 5- or 6(-8)-merous, pedicellate.
Hypanthium globular, smooth, without ridges continuous with the calyx lobes. Calyx
coriaceous; tube cylindrical or tubiform with linear, erect or reflexed lobes. Corolla pale
green in bud, white at anthesis, turning yellow with age; tube cylindrical, tubiform or
crateriform; lobes patent, narrowly ovate to elliptical, usually shorter than the tube.
Anthers sessile, linear, more or less medifixed; tips of pollen-bearing anthers (male and
hermaphrodite flowers) exceeding the tube by several millimetres, sterile anthers (female
flowers) enclosed. Pollen grains single, radially symmetric, isopolar, 3- or 4-colporate,
exine smooth. Style usually clavate; stigmata 2(-4), connate, sometimes with reflexed
lobes, emarginate, exserted by several millimetres in flowers with sterile anthers, rarely
exceeding the corolla tube in male flowers. Disc annular. Ovary unilocular with 2 or 3
parietal placentas; ovules numerous, partially embedded in the placenta. Fruit globular
to ellipsoid drupe or berry, more than 10 mm diameter, usually crowned by persistent
calyx; pericarp thin and parenchymatous with a membranous endocarp, or thick and
fibrous with a bony putamen. Seeds numerous, lenticular, embedded in the firm placental
pulp. Seed-coat thicker around the hilum, which occupies one-third to one-half of the
perimeter of the seed; exotestal cells with radial and inner tangential walls sclerified:
thickening of inner tangential wall pitted or forming a reticulum.

Puttock, Kailarsenia 53

A genus of about eleven species; three endemic to tropical Australia; about eight
species in tropical Asia, but none known from New Guinea.

Key to the Australian species of Kailarsenia

i. Dwarf shrubs with underground stems (geofrutex); leaves lanceolate or, 1f
obovate, less than 3 cm long; mesocarp parenchymatous, endocarp
membranous (NT) .............................. K. suffruticosa

Small trees; leaves elliptical to obovate, more than 5 cm long; mesocarp
fibrous; endocarp brittle (NEQ) .. ............... iS ane een 2

2. Leaves glabrous, 5-12 cm long, 3~7 cm wide, drying black, secondary
veins 8-12 each side of the midvein; petioles 1-4 mm long; stipules

Se NEE Sam csr: See hs ps Ret eee my cases BNL, Pe ke cope bse eens, Se a A oe we
Leaves tomentose to sub-glabrous, 5-25 cm long, 3-13 cm wide, drying
various shades of brown, secondary veins 12-18 each side of the
midvein; petioles 3-12 mm long, stipules 8-11 mm long ...... K. ochreata

K. jardinei

Kailarsenia jardinei (F. Muell. ex Benth.) Puttock, comb. nov.

Gardenia jardinei F. Muell. ex Benth., Fl. austral. 3: 410 (1867). Type: Queensland.
NorTH KENNEDY DISTRICT: Mt Elliot, undated, Da/llachy s.n., (lecto (here chosen):
K!: isolecto: MEL!(MEL 598350)).

F. Mueller, Fragm. 7: 46 (1869); F.M. Bailey, Syn. Queensl. fl. 222 (1883).

Dioecious, columnar tree to 15 m tall; trunk at breast height to 27 cm diameter. Bark
to 25 mm thick, with shallow irregular tessellations, grey; outer bark layered and with
brownish orange blaze; inner bark blaze fawn to cream; bark of branches silver-grey or
ereen, glabrous; lenticels irregular to circular protrusions; young internodes occasionally
with minute hyaline hairs. Leaves ternate or rarely opposite; petioles 1-4 mm long,
glabrous or minutely hairy, greyish green; lamina ovate to elliptical with obtuse apex
and base, 5-12 cm long, 3-7 cm wide, glabrous, glossy deep green above, dull deep
ereen below, thinly coriaceous; secondary veins 8-12 pairs, at 45-55° to the midvein,
raised above and below; tertiary venation translucent; well-formed shallow depressions
in secondary/midvein angles with woolly tuft of long hyaline hairs. Stipules 8-10 mm
long, minutely hairy or glabrous outside; colleters cylindrical, 0.3-0.45 mm long, 0.15-
0.2 mm wide. Female or hermaphrodite flowers unknown; male flowers 5- or 6-merous,
solitary or in 2—4-flowered cymes. Pedicels 3-5 mm long, glabrous. Hypanthium 5 mm
long, glabrous. Calyx glabrous; tube tubiform, 2-3 mm long; lobes linear, 3-5 mm long,
reflexed or rarely erect. Corolla tube cylindrical, 20-40 mm long, 2-3 mm diameter at
the base increasing to 4 mm diameter in the upper part, glabrous; lobes elliptical to
narrowly ovate, 15~30 mm long, 5-10 mm wide, glabrous. Pollen-bearing anthers 8-10
mm long, attached 5-7 mm from their apices, inserted 3-4 mm below the sinuses of
the corolla lobes, their apices exceeding the tube by 2-3 mm. Style 8-10 mm long, not
reaching the anthers, glabrous; stigmatic lobes 2(3), 2-3 mm long, reflexed. Placentas 2
or 3, bearing 60-70 ovules each. Fruit usually solitary, ellipsoid, 25-40 mm long, 20-
40 mm diameter, smooth, crowned by persistent calyx remnants; pedicels 2-4 mm long;
exocarp pale green whilst developing, yellowish green when mature; mesocarp fibrous,
3-5 mm thick; endocarp brittle, 0.5-1.0 mm thick; placental mass cream. Seeds 3.2-4.9
mm diameter, 0.9-1.8 mm thick; hilum occupying 0.37-0.43 of perimeter of seed; seed-
coat pale brown; exotestal cells with pitted thickening of inner tangential walls. Fig. 1
A-I.

Selected specimens: Queensland. NORTH KENNEDY District: Mt Elliot, undated, Dallachy sn. (K,MEL); Mt
Elliot, [1858], Fitzalan s.n. (MEL (MEL 103682, MEL 103683)); 1.5 km E of Splitters Ck, NE of Mt Roundback,
19°59’S, 148°00’E, Jan 1983, Puttock UNSW 14494 (K,UNSW); Port Denison, undated, Dallachy sn. (MEL (MEL
578349)); Mt Dryander, undated, Fitzalan sn. (MEL (MEL 103681)); Cannonvale, Nov 1959, Jones 1347
{(BRI,CANB); between Airlie Beach and Shute Harbour, 20°16’S, 148°46’E, Nov 1983, Puttock UNSW 15822
(UNSW); between Shute Harbour and Airlie Beach, 20°17’S, 148°46’E, Jan 1983, Puttock UNSW 14260

(BISH,BR,K,MEL,UNSW). SouTH KENNEDY DIsTRICT: scrubs nr Mackay, Jan 1926, Langfudy [14] (BRD. (23
specimens examined).

Distribution and habitat: Endemic to the coastal slopes and plains between Mount Elliot
(19°30’S) and Mackay (21°10/S) from sea level to 100 metres altitude but recent collections
only known from Mt Roundback near Bowen, and Conway National Park and neigh-

54 Austrobaileya 3(1): 1989

bouring islands (Map 1). It is found in the open woodland and deciduous vine thickets
behind mangroves on shale and alluvial soils in the Conway National Park.

Phenology and pollination biology: Flowering from November to February, flowers sweetly
perfumed; fruits maturing between March and August. Many small scarab beetles visit
flowers, but no lepidoptera have been observed (pers. obs. Conway NP).

Affinities: Closely related to K. ochreata, from which it differs by its glabrous, glossy
leaves with shorter petioles and fewer secondary veins and shorter stipules, as well as
its essentially glabrous fruits.

Conservation status: This species is locally common near Strathdickie, and between Airlie
Beach and Shute Harbour. Much of its original habitat in this area is now given over
to sugarcane, and other pockets of natural vegetation are under threat of urban devel-
opment. However, small populations will survive in the refuges of Conway National
Park and on the lower slopes of Mount Dryander. Near Bowen (formerly Port Denison)
it can be found sporadically along the banks of creeks draining Mount Roundback. Much
of this area is improved pasture, and very few trees remain even along the gullies.
Conservation status 2K.

Vernacular name: None known.

Etymology: The isolectotype sheet at MEL which was annotated by Mueller states that
the species was named after Mr [F.L.] Jardine, the Police Magistrate at Albany Bay,
Cape York.

Typification and notes: The only sheet of the specimens cited in the protologue and
known to have been seen by Bentham (Mount Elliot, Dallachy) is here chosen as the
lectotype. Protologue specimens from both localities are lodged at MEL.

Although Bentham (1867) described the fruit as having three parietal placentas,
this 1s extremely rare; commonly there are two. As in the other two Australian species,
floral dimorphism probably occurs, but only pollen-producing flowers are known despite
extensive searching in January and November 1983.

Kailarsenia ochreata (F. Muell.) Puttock comb. nev.

Gardenia ochreata F. Muell., Fragm. 1: 55 (1858). Type: Queensland. NorrTH

KENNEDY DistRIct: In less fertile grassy places along the Burdekin River, [October
1856], Mueller s.n., (lecto (here designated): K!(lower right hand element); isolecto:
MELI(MEL 598352)).
Essay on the plants collected by Eugene Fitzalan during Lieut. Smith’s Expedition
to the Estuary of the Burdekin 11 (1860); G. Bentham, FI. austral. 3: 409 (1867);
F.M. Bailey, Syn. Queensl. fl. 222 (1883), Queens. fl. 3: 756 (1900), Compreh.
catal. 241, pl. 215 (1913); K. Domin, Biblioth. Bot. 22(89): 620, pl. 193 (1929).
B.P.M. Hyland, Common Rainforest Trees 2nd ed. 95 (1981).

Gardenia ochreata var. parviflora F. Muell., Fragm. 7: 46 (1869). Type: Queensland.
LEICHHARDT DISTRICT: Issacs R., [1844], Leichhardt 34 (holo: MEL!).

Gardenia macgillivraei Benth., Fl. austral. 3: 409 (1867). Type: Queensland. Cook
Disrricr: Cape York, November 1849, MacGillivray s.n. (Bot. 509), (lecto (here
designated): K(upper element on sheet of flowering material)!).

F. Mueller, Fragm. 7: 46 (1869); F.M. Bailey, Syn. Queensl. fl. 222 (1883), Queensl.
fl. 3: 756 (1900).

Gardenia kershawii Bailey, Queens]. Agric. J., n.s. 2: 75, pl. 37 (1914). Type:
Queensland. Cook District: Claudie River, ‘undated, Kershaw (lecto (here des-
ignated): BRI).

Dioecious or ?gynodioecious, columnar tree to 10(-20) m tall; trunk at breast height to
15 cm diameter. Bark to 15 mm thick, smooth, flaky or shallowly tessellated, silver to
dark grey; outer bark layered, sranular with a pinkish brown blaze; inner bark blaze
cream; branches silver-grey to rusty brown, glabrous to tomentose; lenticels scattered,
tangentially-elongated to circular protrusions. Leaves ternate or occasionally opposite,
tomentose to glabrous; petioles 3-12 mm long, greyish green; lamina obovate to elliptical
with an acute apex and an obtuse or decurrent base, 5-25 cm long, 3-13 cm wide, pale

Puttock, Katlarsenia 55

ee Heiricy,
Shi ‘i “ ek =

at

bY ss

af
SNES

SS

LN,
eat |

he
waht
el aria

Fe ani
rhs aE

oy r a”

eT a

- Mehogine

ry
4
oat y

pllatiee te
oa

p i *
en tet ee eB LE al Ge ae eT
ed ee ce

na Ne a

ieee Le Se nL

Fig. 1. Kailarsenia jardinei: A. flowering branchlet < 0.67. B. mature fruit x 1. C. T.S. of mature fruit x 1. D.
style X 3. E. anther, 2 views X 3. F. T.S. of polleniferous anther X 6. G. seed, 2 views X 3. H. stipule crowning
branch tip X 5. L embryo X 10. K. ochreata: J. flowering branchlets X 0.67. K. style X 3. L. anther, 2 views X
3. M. T.S. of polleniferous anther X 6. N. T.S. of mature fruit X 1. O. Mature fruit x |. P. seed, 2 views X 3.
Q. stipule crowning branch tip X 5. R. embryo X 10. (A, Puttock UNSW14260; D-F, Puttock UNSW15822;
B,C,G,1, Puttock UNSW15809; H, Puttock UNSW14494; J, Wyatt s.n.; K-M, Clarkson 5032; N,O,P,R, Puttock
UNSW13326 & Wilson, Q, Puttock UNSW15833).

56 Austrobaileya 3(1): 1989

green above, dull pale green below, chartaceous to thinly coriaceous; secondary veins
{2-18 pairs, at 45-60° to the midvein, not raised above, raised below; tertiary venation
translucent; shallow depressions 1n secondary/midvein angles with a dense tuft of long
hyaline hairs. Stipules 5-12 mm long, tomentose to subglabrous; colleters lanceolate,
0.3-0.5 mm long, 0.12-0.17 mm wide, accompanied by hyaline hairs. Flowers (5)6-
merous, solitary or occasionally in 2~4-flowered cymes; pedicels 6-12 mm long, tomentose
or glabrous. Hypanthium 5-12 mm long, tomentose to glabrous. Calyx tube tubiform,
5-8 mm long; lobes linear, 6-12 mm long, reflexed or erect, tomentose to glabrous.
Corolla tube cylindrical, 20-55 mm long, 2-3 mm diameter at the base increasing to 4-
5 mm diameter in the upper part, glabrous or sparsely pubescent outside, hairy to
subglabrous inside; lobes narrowly elliptical, 17-40 mm long, 6-13 mm wide, glabrous.
Pollen-bearing anthers 12-14 mm long, attached 5-7 mm from their apices, inserted 2-
3 mm below the sinuses of the lobes, exceeding the tube by 3~6 mm. Sterile anthers 6-
8 mm long, attached 2~3 mm from their apices, included within the corolla tube. Style
18~58 mm long, hairy to glabrous, exceeding the corolla tube by 3-6 mm in plants with
sterile anthers, but reaching only to the middle of the anthers 1n pollen-producing flowers;
stigmatic lobes (2)3(4), 6-14 mm long, connate. Placentas (2)3(4). Fruit usually solitary,
ovoid, 22-50 mm long, 15-35 mm diameter, smooth, glabrous or sparsely hairy; pedicels
6-12 mm long; calyx persistent as a collar or reflexed ring bearing remnants of the lobes;
exocarp pale green whilst developing, yellowish green when mature, 4-6 mm thick:
mesocarp fibrous, 3-5 mm thick; endocarp brittle or hard, 0.5-2.5 mm thick, yellow;
placental mass cream. Seeds 3.1-6.2 mm diameter, 1-1.3 mm thick; hilum occupying
0.34-0.45 of perimeter of seed; seed-coat pale brown; exotestal cells with reticulate
thickening of inner tangential wails. Fig. 1 J-R.

Selected specimens: Queensland, Cook DISTRICT: Cape York (between carpark and beach), 10°41’S, 142°31’E, Jul
1984, Puttock UNSW 16933 & King (BRI,;CANB,UNSW); Cape York, Nov 1849, MacGillivray s.n. [Bot. 509]
(K); Ca pe York, undated, A7// {31 (K); South Kokialah Ck, c. 40 km E of Aurakun, 13°20’S, 142°32’E, May
1982, Clarkson 4392 (K, NSW ,ORS); Claudie R., undated, Kershaw (BRI); Melville Ra. 14°] 15'S, 144°30'E, sep
1970, Hyland 4817 (BRI,QRS),; Kimba—Palmerville rd, 15°35'S, 144°03’E, Nov 1983, Clarkson 5032 (BRI, UNSW);
Trevethan Ra. nr Black Gap, Cooktown Dev. rd, 15°39’ S, 145°13’ E, Dec 1983, Puttock UNSW 15929
(BRLCANB,NSW,UNSW); slopes of Mt Surprise, Dec 1983, Puttock UNSW 15833 (CANB,NSW,UNSW). NorTH
were District: In less fertile grassy places along the Burdekin River, [1856], Afweller s.n. (KR,MEL); Magnetic

19°10’S, 146°50’E, Apr 1970, Wyatt s.n. (BRD; Alligator Ck, 19° 27S, L46°57’E, Jun 1982, Puttock UNSW
13326 & Wilson (BRI, CANB UNSW); Burdekin Expedition, [Cape Upstart, 1859, Fitzalan] il (MEL). SOUTH
KENNEDY District: Burdekin crossing, nr “Glendon”, 20°39’S, 147°10’E, Sep 1950, Smith 4626 (BRI); c. 10 km
from Mt Coolon towards Collinsville, Nov 1978, Stanley 78377 & Ross (BRI). LEICHHARDT District: Isaacs R.,
[1844], Leichhardt 34 (MEL); SW end of Lake Elphinstone, 21°33’S, 148°14VE, Jan 1983, Puttock UNSW 14241
(BISH,BR,BRI,DNA,K,L,UNSW); “Tay Glen” nr Cotherstone, E of Clermont, Sep 1959, Thomson s.n. (BRD.
{128 specimens examined).

Distribution and habitat: The species occurs in Queensland from the tip of Cape York
(latitude 10°42’S) to Clermont (latitude 22°40’S); Map 1. In the northern part of Cape
York it is commonly found in semi-evergreen mesophyll vine forests on alluvial soil
with 1300-1600 mm annual rainfall. In the southern part of its range it 1s restricted to
semideciduous vine forests, deciduous vine thickets and occasionally open woodland,
srowing almost exclusively in association with granite outcrops or on soils derived from
granite with less than 1000 mm annual rainfall.

Phenology and pollination biology: Flowers are present from October to February
(occasionally aseasonally), sweetly perfumed, visited (?pollinated) by papilionoid lepi-
doptera in the late afternoon (pers. obs. Cape York); fruits mature between March and
August and ripen on the ground. Reported to be eaten by aborigines (Mueller 1860).

Vernacular names: Scented Gardenia bush, wild Gardenia (Williams 1979),

Etymology: Ochreata (from ocreatus, greaved, 1.e. fitted with armour for the ankles and
shins) refers to the sheathing nature of the oblong stipules (Mueller 1858).

Typification and notes: Mueller’s type collection of Gardenia ochreata was made during
the latter stages of the Gregory Expedition (October 1856), from one of numerous pockets
of semideciduous vine forest along the Burdekin valley between “Jervoise” and the
confluence of the Burdekin and Suttor Rivers. The best of the type material is at K: of
the four elements on the single sheet there, the flower-bearing element in the lower right
is designated lectotype. Although the fruit was described in the protologue, none are
with the type material. Authentic type material 1s labelled ““Burdekin”’, not to be confused
with other collections labelled “Burdekin Expedition” collected by Fitzalan at Cape
Upstart in 1859. |

Puttock, Kailarsenia 57

Mueller (1869) described G. ochreata var. parviflora for a collection made by
Leichhardt from Isaac River. Only one specimen is known and this is regarded as the
holotype. Mueller distinguished this variety by the corolla lobes being “only 1/2” long”
and “‘equalling the tube’; the corolla tube of var. ochreata being “up to 2” long”.

Gardenia kershawii Bailey was described from a collection of mixed material (see
Puttock 1988). The lectotype is conspecific with Kailarsenia ochreata.

Gardenia ochreata var. parviflora and G. kershawii have only ever been applied
to their type specimens.

Gardenia macgillivraei Benth. was described from two collections from the vicinity
of “Somerset” at the top of Cape York Peninsula. The upper element on MacGillivray’s
sheet of flowering material at K is chosen as the lectotype. Bentham (1867) did not
describe the anthers. Mueller (1869) added the description of the anthers from an
unspecihed collection.

G. macgillivraei has been separated from G. ochreata on the basis of leaf shape,
size, venation and tomentum, bark type, flower and fruit size, and distribution. The
northern material is also highly variable in endocarp thickness. To investigate the limits
between these populations, additional collections were made in the region between
northern Cape York and the South Kennedy District (11°S and 16°S). From north to
south trees become smaller, leaves become smaller with fewer veins and more densely
hairy or tomentose, flowers and fruits become smaller, bark becomes tessellated, and
the habitat becomes restricted to granite soils and outcrops, and to regions of lower
rainfall. The trend in the leaf size and hairiness 1s illustrated in Figure 2 using specimens
collected at approximately one degree intervals of latitude.

200 J

Vi K O P
10 13 16 19 22
Fig. 2. Raw values of lamina length (CJ) and width (O) for herbarium specimens of Kailarsenia ochreata collected

closest to 40’S of each degree of latitude within its range. Open symbols = + glabrous; half closed symbols =
scattered hairs; closed symbols = tomentose. M,K,O, & P indicate vicinity of type localities (see Map 1).

58 Austrobaileya 3(1): 1989

To test the hypothesis that there is continuous and gradual variation between the
two extreme forms, all available specimens with mature fruit (47) were scored for 8
characters:

. number of veins each side of the midrib;

. pubescence on the abaxial surface: dense, scattered, glabrous;
. petiole length;

. lamina length;

. lamina width;

. pedicel length;

. fruit length;

. fruit width.

The data obtained were normalised and standardised and then subjected to a multiple
regression analysis. The transformed data were reduced to a single value (Y) for each
specimen. The scattergram of the Y values plotted against latitude falls within a narrow
band with gradually increasing Y values to the south with no descernible disjunctions
(Fig. 3). The multiple regression was highly significant (P > > 0.001). A similar analysis
was performed on the 22 flowering specimens available, replacing corolla tube and lobe
lengths for characters 7 and 8. Again, this revealed no clear discontinuity that would
indicate existence of separate gene pools (Fig. 4), (P ~ 0.001). It 1s concluded, therefore
that the specimens represent a single gene pool in which there is clinal variation in
several characters that correlates with latitide. Hence Gardenia macgillivrael, G. kershawii
and G. ochreata var. parviflora are reduced to synonymy with Kailarsenia ochreata and
without subspecific ranks. The separate specific status previously accorded the northern
and southern populations is an artifact of the marked differences between the extreme
forms of the cline and lack of collections in intervening regions.

OO IA Un Be Gb it

The floral dimorphism in this species is uniform throughout its distribution. Plants
bearing flowers with sterile anthers and long styles have been observed to produce mature
fruits. The ovaries of flowers with pollen producing anthers and short styles are well
developed, having numerous ovules, but it has not been established whether they ever
develop fruits and fertile seeds.

2 @
4 , @ @
O @ @ 1
-4 ree ~ .
10 12 14 16 18 20 22 24

Fig 3. Scattergram of values obtained from multiple regression analysis of individual fruit-bearing trees of
Kailarsenia ochreata plotted against latitude. For characters scored see text. Y is the standardized form (with
mean = 0 and standard deviation = 1) of the multiple regression estimate of latitude based on robust transformations
of the observations.

Puttock, Kailarsenia $9

2
1 P & &
O ‘
“1 © ® : &
10 12 14 16 18 20 22 24

Fig. 4. Scattergram of values obtained from multiple regression analysis of individual flowering trees of Kailarsenia
ochreata plotted against latitude. For characters see text. Y is the standardized form (with mean = 0 and standard
deviation = 1) of the multiple regression estimate of latitude based on robust transformations of the observations.

Kailarsenia suffruticosa (R. Br. ex Benth.) Puttock comb. nov.

Gardenia suffruticosa R. Br. ex Benth., Fl. austral. 3: 410 (1867). Type: Northern
Territory. DARWIN AND GULF: Point S, Gulf of Carpentaria, 28 January 1803,
Brown [3455], (lecto (here designated): BM!(fruiting element on left of sheet);
isolecto: K!, two sheets).

F.M. Bailey, Syn. Queensl. fl. 221 (1883); F.M. Bailey, Catal. pl. Queensland 22
(1890), Queensl. fl. 3: 756 (1900); Specht, Rec. Amer. Austral. exped. Arnhem
Land 306 (1958).

Gynodioecious or ?dioecious, dwarf shrub (geofrutex) to 0.3 m high. Underground
branches 10-20 mm diameter; aerial branches to 10 mm diameter; bark smooth, fawn:
outer bark layered, granular, with a pinkish brown blaze; inner bark with cream blaze;
branchlets pubescent; lenticels absent; subrhytidome deep green. Leaves opposite or
ternate; petioles 1-3 mm long, greyish green; lamina lanceolate to oblong-obovate with
an acute apex and an obtuse or decurrent base, 8-12 cm long, 1.5-—2 cm wide, pale green
above, dull pale green below, chartaceous, bullate and pubescent, or flat with scattered
hairs below; secondary veins 6-13 pairs, at 40-50° to the midvein, sunken above, strongly
raised below; tertiary venation translucent; shallow depressions in secondary/midvein
angles on the abaxial surface with or without hyaline hairs. Stipules 5-8 mm long,
pubescent outside; colleters lanceolate, 0.25-0.35 mm long, 0.12-0.15 mm wide, accom-
panied by hyaline hairs. Flowers 6(7)-merous, solitary or 1n 2—4-flowered cymes; pedicels
6-15 mm long, tomentose. Hypanthium 5 mm long, pubescent. Calyx tube cylindrical,
3~5 mm long; lobes linear, 5-10 mm long, erect, tomentose. Corolla tube cylindrical to
tubiform, 18-25 mm long, 3-4 mm diameter at the base increasing to 7-8 mm diameter
in the upper part, pubescent outside, tomentose on the upper part inside, glabrous below;
lobes narrowly elliptical, 20-23 mm long, 10-13 mm broad, glabrous. Pollen-bearing
anthers 9-11 mm long, attached 5-7 mm from their apices, inserted 4-5 mm below the
sinuses of the corolla lobes, the tips exceeding the tube by 1-3 mm; sterile anthers 7-9
mm long, otherwise identical to pollen-bearing anthers. Style 20-28 mm long, exceeding
the corolla tube by 2-6 mm, glabrous; stigmatic lobes 2(3), 9-14 mm long, connate.
Placentas 2(3). Fruit solitary (only immature known) ovoid, 20-25 mm long, 12-15 mm
diameter, smooth with scattered hairs; calyx persisting as a collar or reflexed ring bearing
remnants of the lobes; pedicels 10-18 mm long; pericarp (not fully mature) succulent,
about 2 mm thick; endocarp membranous. Mature seeds unknown. Fig. 5.

60 Austrobaileya 3(1): 1989

Selected specimens: Northern Territory, DARWIN AND GULF District: Pickataramoor, Melville I., 11°45’,
130°53’E, Dec 1977, Angeles s.n. (DNA); Gunn Pt area, 12°13’S, 131°03’E, Nov 1978, Rankin 1562
(CANB,DNA,MEL,PERTH); 6.1 km S of Old Point Stuart ruins, {2°21’S, 131°49’E, Nov 1980, Waterhouse UNSW
9857 (CANB,UNSW); 3.9 km E of Hades Flat, SE of Jabiluka Outhter, [2°40’S, 132°53’E, Dec 1980, Waterhouse
UNSW 10923 (BR,K,UNSW), 12°05’S, 133°41’E, Jun 1974, Pullen 9484 (CANB); Point S, [Port Blane, Groote
Eylandt,] Gulf of Carpentaria, Jan 1803, Brown 3455 (BM,K); Emeraid R. rd, Groote Eylandt, 14°01’S 136°27’E,
Feb 1977, Waddy 635 (DNA); Hemple Bay, Groote Eylandt, May 1948, Specht 342 (BRI,CANB,MEL,NSW);
Yirrkala, 12°12’S, 136°47’E, Aug 1948, Specht 896 (CANB). (16 specimens examined).

Distribution and habitat: Endemic to the coastal plains and islands of the Darwin and
Gulf District of the Northern Territory (Map 1). Of sporadic occurrence in open woodland
on well drained sandy flats.

Phenology: Flowering mostly from October to February, but occasionally aseasonal,
flowers sweetly perfumed; fruiting early in the dry season (from March).

Affinities: K. suffruticosa resembles K. ochreata and K. jardinei in its ocreate stipules
and corolla morphology, but differs from both in its lanceolate-oblong leaves, broader
corolla tube, suffruticose habit, parenchymatous mesocarp and membranous putamen.
In fruit characters and habit, K. suffruticosa has a closer affinity to the type species, K.
tentaculata, from Malaysia.

Conservation status: This species is rarely collected, being generally obscured by tall
grass for much of the year. It regenerates well after fire. It is probably under threat from
srazing in some areas but may be well represented in Kakadu National Park (conservation
Status 3K).

Vernacular name: None known.
Etymology: Named for 1ts low spreading habit.

Typification and notes: Of the three sheets of type material available, only that at BM
has a precise location and date of collection; the fruiting element on the left of the BM
sheet 1s designated lectotype.

Fig. 5. Kailarsenia suffruticosa: A. flowering branchlets < 0.67. B. style x 3. C. anther, 2 views X 3. D. T.S. of
polleniferous anther < 6. E. fruit (not completely mature) xX !f. F. T.S. of fruit x 1. G. Stipule crowning branch
tip X 5. (A, Waterhouse UNSW9857: B-D,G, Waterhouse UNSW10923; E\F, Waddy 635).

Puttock, Kailarsenia 61

This species requires further collecting to determine the significance of two leaf
forms present in the material. Around Darwin to the west of Arnhemland and on Groote
Eylandt the leaves are flat and bear scattered hairs on the veins; near the Arnhemland
escarpment the leaves are bullate and densely pubescent.

Few flowering collections exist in herbaria and detail of the floral dimorphism 1s
incompletely known. Some specimens have short, sterile anthers and long styles which
appear to be receptive (females). Other specimens have longer anthers producing pollen,
although, as in K. ochreata, the ovaries are scarcely smaller than in flowers with sterile
anthers. However, unlike K. ochreata, the style is usually as long as, or longer than the
corolla tube, and several pollen producing plants have been found with young fruits.
Hence these flowers appear to be hermaphrodite.

“0

Map 1. Distribution of Kailarsenia species in northern Australia. A X. jardinei, @ K. ochreata, WK. suffruticosa.
O, P, M, & K indicate the localities of the type specimens of Gardenia ochreata, G. ochreata var. parviflora, G.
macegillivraei and G. kershawiti respectively.

Acknowledgements

1 would like to thank the directors of BM, BR, BRI, CANB, CBG, DNA, E, JCT,
K, L, MEL, NSW, QRS for making material available for study; J. Clarkson (DPI,
Mareeba) for valuable dried and pickled material; B. Hyland for use of facilities at QRS;
R.S. Vickery for advice on regression analyses; C.J. Quinn and E. Robbrecht for comments
on the manuscript; R.J. King, S. Paterson, S.J. St George, P.G. Wilson for assistance
with field trips; the Linnean Society of NSW for a Joyce W. Vickery Research Fund
travel grant; and to the late J.T. Waterhouse for encouragement in the early stages of
my work on the Gardenieae.

References
BENTHAM, G. (1867). Flora australiensis. Vol. 3. London: Lovell Reeve & Co.

BREMEKAMP, C.E.B. (1957). Some new species of Acanthaceae and Rubiaceae from Laos (Indochina). Koninklijke
Nederlandse Academie van Wetenschappen. Proceedings Series C. 60: 1-14.

ae i Na . (1973). Classification of the architecture of dicotyledonous leaves. American Journal of Botany 60:
17-33.

62 Austrobaileya 3(1): 1989

HOOKER, J.D. (1880). Flora of British India. Vol. 3(7). London: Loveli Reeve & Co.
KEAY, R.W.J. (1958). Randia and Gardenia in West Africa. Bulletin du Jardin botanique de l’Etat. Bruxelles 28:

LEIGH, J., BRIGGS, J. & HARTLEY, W. (1981). Rare or threatened Australian Plants. Australian National
Parks and Wildlife Service Special Publication No. 7. Canberra: Australian National Parks and Wildlife
Service.

MUELLER, F. (1858). Fragmenta phytographiae Australiae. Vol. |. Melbourne: Government Printer.

MUELLER, F. (1860). Essay on the plants collected by Eugene Fitzalan during Lieut. Smith’s Expedition to the
Estuary of the Burdekin. Melbourne: Government Printer.

MUELLER, F. (1869). Fragmenta phytographiae Australiae. Vol 7. Melbourne: Government Printer.
PUTTOCK, C.F. (1988). A revision of Gardenia (Rubiaceae) from north-eastern Queensland. Austrobaileyva 2:
433-449.

ROBBRECHT, E. (1988). Tropical woody Rubiaceae. Opera Botanica Belgica. Vol. 1. Meise. 272 pp.

ROBBRECHT, E. & PUFF, C. (1986). A survey of the Gardenieae and related tribes (Rubiaceae). Botanische
Jahrbticher 108: 63-137.

TIRVENGADUM, D.D, (1983). New taxa and name changes in tropical Asiatic Rubiaceae. Nordic Journal of
Botany 3: 455-469,

TIRVENGADUM, D.D. & SASTRE, C. (1979). La signification taxonomique des modes de ramification de
Randia et genres afines. Mauritius Institute Bulletin 8: 77-98.

WILLIAMS, K.A.W. (1979). Native plants of Queensland Vol. |. Brisbane: Cranbrook.
YAMAZAKI, T. (1970). A revision of the genus Randia L. in Eastern Asia. Journal of Japanese Botany 45: 337-~
341.

Accepted for publication 3 March 1989

Austrobaileya 3(1): 63-67 (1989) 63

THE GENUS ANCANA F. MUELL. (ANNONACEAE) IN
AUSTRALIA

L.W. Jessup

Queensland Herbarium, Meiers Road, Indooroopilly, Qid 4068

Summary

Two species of Ancana F. Muell. occur in Australia, viz A. stenopetala F. Muell. and A. hirsuta which is described
as new. Possible relationships with other genera in Annonaceae are briefly discussed.

The genus Ancana F. Muell. was described to accommodate a single species A.
stenopetala F. Muell. This situation remained unchanged until Fries (1953) transferred
this species to Fissistigma Griffith. The treatment of Ancana as congeneric with Fissis-
tigma has been maintained in later classifications of the Annonaceae (Fries 1959;
Hutchinson 1964).

Early in my study of Australian Annonaceae it became apparent that because of
the differences given below this seneric transfer by Fries could not be upheld. This view
was reinforced by the discovery of a second species in Queensland referable to Ancana.

Ancana species are shrubs with solitary axillary flowers and the ovaries are
surmounted by a markedly differentiated globose, sessile stigma, narrowly and eccentr-
ically attached. Fissistigma species are climbers with few- to several-flowered terminal
or leaf-opposed inflorescences and the obloid or clavate stigma is broadly attached and
not clearly differentiated from the ovary. More fundamental differences in chromosome
number (2n = 18 in Ancana, In = 16 in Fissistigma) and pollen structure have been
documented by Morawetz (1988) and Waha and Morawetz (1988).

Measurements of flowers and fruit unless otherwise noted in this account are
taken from material preserved in spirit. Petal width was measured at the widest point
and petal thickness was measured midway along the petal length, 1.¢. “midlength”’.

ANCANA
Ancana F. Muell., Fragm. 9: 27 (1865). Type: Ancana stenopetala F. Muell.

Evergreen understorey shrubs. Flowers hermaphrodite, solitary, axillary. Sepals 3, valvate,
free or very shortly connate at the base. Outer petals 3, valvate. Inner petals 3, valvate,
similar in size to outer petals, srooved on inner surface near the base. Torus hemispherical,
pilose between stamens and carpels. Stamens broadly wedge-shaped, anthers nearly
sessile. Top surface of connective flat, oblique, concealing the anther cells, produced
adaxially and upward on the innermost stamens. Carpels several, free; stigma capitate,
obliquely attached to either side of the suture. Mature monocarps subglobular to allantoid,
usually umbonate at the apex, the shape and length depending on the number of seeds
set. Seeds transversely ellipsoid with a circumferential groove.

Distribution: The genus is an Australian endemic consisting of two allopatric species.

Key to Species

1. Outer petals 5-5.5 mm wide and at midlength 0.5-0.7 mm thick. Inner

petals 3.5-4 mm wide and at midlength 0.8-1.2 mm thick. Mature
MMOTOCHIDS BIBOFOUS. 5 ew 5 Se Se eh ps 1. A. stenopetala
Outer petals 8-9 mm _ wide and at midlength ca 0.2 mm thick. Inner
petals 7-9 mm wide and at midlength ca 0.2 mm thick. Mature
monocarps hirsute hone eee ale khe tii eewins Saas 2. A. hirsuta

*

1989

Austrobaileya 3(1)

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Jessup 837

Jessup, Ancana 65

1. Ancana stenopetala F. Muell., Fragm. 5: 27 (1865); Unona ancana F. Muell., Fragm.
§:27 (1865), nom. inval. in synon., Unona_ancana F. Muell. ex Bailey, Suppl. to
Syn. Queensl. fl. 3: 5 (1890), nom. illegit.; Fissistigma stenopetala (F. Muell.) R.E.
Fries, Ark. Bot. 3(2): 36-37 (1953). Type: Tweed River, C. Moore (n.v.).

Shrubs to 5 m, flowering at 0.5 m. Shoots with antrorsely appressed, shining, brown,
simple, thick-walled hairs, glabrescent. Twigs soon glabrous. Leaf laminae lanceolate,
oblanceolate or narrowly ovate, acuminate, 5-10(-14) cm_* 1.5-3(-4) cm, glabrous on
adaxial surface, scattered appressed hairs on abaxial surface, glabrescent, base obtuse,
rounded or broadly acute; midvein shallowly channelled above; secondary veins mostly
12-18 pairs, with some intersecondaries, raised on both surfaces; tertiary venation
reticulate, raised. Petioles 2.5-3 mm long, with some persistent hairs. Pedicels 3-4 mm
long with 2 or 3 ovate suprabasal bracts 1.5-2.5 mm long. Sepals broadly ovate or
triangular, acuminate or acute, 4-4.) mm X ca 3 mm, appressed pubescent on outside.
Outer petals very narrowly ovate, convex on outside, concave on inner side, 30-435 mm
x 5-5.5 mm at 4-6 mm from base, 0.5-0.7 mm thick at midlength, pubescent outside
with appressed brown hairs 0.3-0.5 mm long and semi-erect pale hairs ca 0.1 mm long,
pubescent on inner side with semi-erect pale hairs ca 0.1 mm long. Inner petals nearly
linear, 30-48 mm X 3,5-4 mm near base tapering to 2.5 mm near apex, 0.8-1.2 mm
thick at midlength; inner surface shallowly concave near base and with about 5 or 6
glabrous longitudinal grooves extending for about 1/3 the length of the petal, towards
apex flat or slightly convex and covered with short tortuous hairs; outer surface pubescent
with straight appressed and tortuous hairs, convex near base, flat or slightly concave
towards apex. Stamens ca 35-45, 1.5-2 mm long, anther cells 1-1.5 mm _ long. Carpels
usually 5, ovary ca 1.5 mm long with appressed hairs ca 0.5 mm long. Ovules 5 or 6.
Ripe monocarps up to 4.5 x 2 cm, warted, glabrous, on stipes up to 5 mm long. Seeds
usually 2-5. Fig. 1.

Specimens examined: Queensland. MORETON District: Tambourine Mt, May 1945, Blake 15822 (BRI); ditto,
Apr 1947, Clemens (BRI); Upper Coomera—Canungra road ca 25 km from Upper Coomera, Mar 1976, Stanley
80 (BRI); Burleigh Heads, Dec 1917, White (BRI); Currumbin, Mar 1916, Longman (BRI). New South Wales.
NortTH COAst: Tweed River, Fawcett [AQ210452] (BRI,MEL); Mt Warning, Dec 1981, Jessup 457 (BRI); Terania
Ck, 10 km NNE of The Shannon, Feb 1980, Coveny 10627 (BRI); North Ck, Ballina, Jan 1892, Bauerlen 737
(BRI).

Distribution and habitat: This species occurs from Tambourine Mountain, Queensland
to near Coffs Harbour, New South Wales. It is often locally common in fairly restricted
areas. It is found in upland, lowland and littoral types of complex notophyll vine forest
on soil derived from or enriched with basalt.

’. Ancana hirsuta Jessup species nova

Frutex usque 2.5 m altus. Innovationes pilis antrorsis adpressis nitido-cinnamomeis,
simplicibus, parietibus crassis obsitae. Ramunculi hirsuti. Laminae foliorum lanceolatae,
oblanceolatae vel ellipticae, plerumque acuminatae, 5-10(-14) cm longae et 1.3-3(-5)
cm latae, supra praeter costam glabrae, infra pilis dispersis adpressis obsitae, basi obtusae
rotundatae vel late acutae. Petioli 2-2.5 mm _ longi, hirsuti. Pedicelli 2-4 mm long}
pubescentes, bracteis late ovatis 1.5-2.2 mm longis. Sepalae late ovatae, abrupte acu-
minatae, 4.5-5.5 mm longae et 4-4.8 mm latae, extus pubescentiae, intus glabrae. Petalae
exteriorae anguste ovatae, attenuatae, 30-40 mm longae et 8-9 mm latae, extus adpresse
pubescentes, intus breviter et molliter pubescentes. Petalae interiorae anguste ovatae,
attenuatae, 30-48 mm longae et 7-9 mm latae, extus pubescentes, intus prope basin
concavae, glabrae, multisulcatae, versus apice planae vel cum marginibus incurvis,
pubescentae. Stamina 45-60, 1.2-2 mm longa, antherae 0.8-1.6 mm longae. Carpella 5-
11(-16), ovaria !.8-2 mm _ longa, pilis adpressis, 0.5 mm longis obsita. Ovula 4-6.
Monocarpia matura (in siccitate) ad 2 cm longa et | cm lata, hirsuta, pilis ca | mm
longis, cinnamomeis. Semina I-35. Typus: Queensland. Cook DISTRICT: Henrietta Creek,
Palmerston Highway, 17°36’S 145°40’E, November 1982, Jessup 512 (holo: BRI; 1so:
BRICBG,K,L,MEL,NSW,QRS,U).

Shrub to 2.5 m, flowering at less than 0.5 m. Shoots with antrorsely appressed, shiny
brown, simple, thick-walled hairs. Twigs hirsute. Leaf lamina lanceolate, oblanceolate or
elliptic, usually acuminate, 5—10(-14) cm x |.3-3(-5) cm, glabrous on adaxial surface
except for pilose midvein, scattered appressed hairy on abaxial surface, base obtuse,
rounded or broadly acute; midvein shallowly channelled above, raised below; secondary
veins mostly 13-16 pairs, with some intersecondaries, scarcely raised above, more

66 Austrobaileya 3(1): 1989

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| : A. | . B. of flower with | outer and
removed X 3. C. inner surface of inner petal x 3. D. carpel X 12. E. L.S. carpel x 25. F. outer enon te

inner stamen X 12. H. stamen with dehisced anthers x 12. 1 I
Jessup 706; D-H, Jessup 512: I, Jessup 478 nearly mature monocarps in dry state X 2. A-C,

Jessup, Ancana 67

prominently raised below; tertiary venation reticulate to percurrent. Petiole 2—2.5 mm
long, hirsute. Pedicel 2-4 mm long, pubescent, with 2 or 3 broadly ovate bracts 1.5-2.2
mm long. Sepals broadly ovate, abruptly acuminate, 4.5—5.5 mm X 4-4.8 mm, pubescent
on outside, glabrous on inner side. Outer petals narrowly ovate, attenuate, convex on
outside, concave on inner side, 30-40 mm X 8-9 mm at 5—7 mm from base, ca 0.2
mm thick at midlength, appressed hairy outside, softly pubescent on inner side. Inner
petals narrowly ovate, attenuate, 30-48 mm X 7-9 mm at 4-6 mm from base, ca 0.2
mm thick at midlength; outer surface pubescent, convex near base, elsewhere flat; inner
surface concave near base, glabrous, multigrooved, the grooves becoming fewer and
wider distally and extending for about 1/4-1/3 the length of the petal, towards apex flat
or with incurved margins, pubescent. Stamens 45-60, 1.2-2 mm long; anther cells 0.8-
1.6 mm long. Carpels 5-11(-16), ovaries 1.8-2 mm long with appressed hairs ca 0.5
mm long. Ovules 4-6. Ripe monocarps (when dried) up to 2cm X | cm, hirsute, the
hairs ca 1 mm long, brown. Seeds 1-5. Fig. 2.

Specimens examined: Queensland. Cook District: Cedar Bay, N of Bloomfield R, Oct 1972, Webb & Tracey
13359 (Dick 8) (BRI); ditto, Jan 1973, Webb & Tracey 13360 (Dick 24) (BRI); near entrance to Mossman Gorge
National Park, near Mossman R, Dec 1982, Jessup 544, 545 (BRI), ditto, Dec 1984, Jessup 706 (BRI); Crawfords
Lookout to Tchupalla Falls Track, Palmerston National Park, Feb 1982, Jessup 478 & Tracey (BRI,QRS), Henrietta
Ck, Palmerston Highway, Nov 1982, Jessup 512 (BRI,CBG,K,L,MEL,NSW,QRS,U).

Distribution and habitat: This species has been found only in three refugia in north
eastern Queensland: at Cedar Bay, along the Mossman River and the North Johnstone
River. It occurs on lowlands and footslopes in complex mesophyll vine forest on soil
derived from basalt or a mixture of acidic and basic rock types.

Discussion

The relationships of Ancana with other genera are still unclear. Waha and Morawetz
(1988) have shown that the genus shares a disulcate pollen character with Fitzalania
and Haplostichanthus as well as the 2n = 18 chromosome number.

The genera Meiogyne, Polyaulax, Guamia and Oncodostigma all have a warted
or grooved inner surface to the inner petals near the base, axillary flowers, flat, oblique
connectives to the stamens and a subglobular stigma all similar to those in Ancana.
Ancana differs from Meiogyne, Guamia and Oncodostigma in having fewer ovules per
carpel. Meiogyne has predominantly Early Phase venation while Guamia, Oncodostigma
and Polyaulax have predominantly Middle Phase venation (Klucking 1986). The venation
of Ancana is also largely Middle Phase. Po/yaulax differs from the other genera by
having much shorter and thicker petals.

Acknowledgements

Mr W. Smith prepared the line drawings. Mr R. Henderson offered helpful criticism
of the manuscript.

References

FRIES, R.E. (1953). Verstreute Beobachtungen der Familie Annonaceae. Arkiv for botanik utgivet av k. svenska
vetenskapsakademien. Stockholm. 3(2): 36-37.

FRIES. R.E. (1959). Die Naturlichen Pflanzenfamilien 17a: }—47 1.

HUTCHINSON, J. (1964). The Genera of Flowering Plants. Dicotyledons I: 71-108. London: Oxford University
Press.

KLUCKING, E.P. (1986). Leaf Venation Patterns. Volume i. Annonaceae. Berlin, Stuttgart: J. Cramer.

MORAWETZ, W. (1988). Karyosystematics and evolution of Australian Annonaceae as compared with Eupom-
atiaceae. Himantandraceae and Austrobaileyaceae. Plant Systematics and Evolution 159: 49-79.

WAHA. M. & MORAWETZ W. (1988). Pollen evolution and systematics in Annonaceae with special reference
to the disulcate Australian endemic genera. Plant Systematics and Evolution 161: 1-i2.

Accepted for publication 23 January 1989

Austrobaileya 3(1): 69-78 (1989) 69

SECAMONE R. BR. (ASCLEPIADACEAE: SECAMONOIDEAE)
IN AUSTRALIA

P.J. Forster
Botany Department, University of Queensland, St Lucia, 4067
and K. Harold

6 Woodland Rise, Wakefield, WF2 9DL, England

Summary

Secamone is represented in Australia by a single variable species, S. elliptica R. Br. which also occurs in Malesia.
Lectotypes are designated for §. elliptica and the synonyms S. ovata R. Br. and S. papuana Warb. S. lanceolata
Blume, S. micrantha (Decne) Decne, S. attenuata Decne, S. multiflora Decne and S. flavida Schltr. are newly
placed in synonymy. An illustrated taxonomic account with notes on variation, distribution, ecology, habitat and
conservation status is given.

Introduction

Taxa included within the subfamily Secamonoideae (Asclepiadaceae) are charac-
terised by the anthers being dithecal and tetrasporangiate, with two pollinia in each
theca (Schill & Jakel 1978). Thus the Secamonoideae differ from the two other subfamilies
of Asclepiadaceae in having 4 pollinia per pollinarium, as opposed to 2 in the Asclepia-
doideae and an aggregation of granular pollen in the Periplocoideae.

Floral development in species of Secamone has been extensively investigated,
primarily by Safwat (1962) but also by Puri and Shiam (1966). Safwat (1962) outlined
in detail how the bilocular anthers of the Asclepiadoideae (as Cynanchoideae) are a
result of phylogenetic suppression of the two outer locules of each anther, whereas in 4-
celled anthers in the Secamonoideae no suppression has occurred. In Secamone, the
entire style-head is glandular in comparison to the 5 glandular regions found in the
Asclepiadoideae.

The genus Secamone R. Br. was first validly described in Brown (1810a) and not
in Brown (1810b) as listed in Farr et a/. (1979). Brown (1810b) was issued as a preprint
of Brown (1811) and was intended to be simultaneously published with Brown (1810a),
but the Prodromus predates the preprint by some seven days (Mabberley 1985). As
listed in Farr et a/. (1979), Phillips (1951) lectotypified Secamone with S. emetica (Retz.)
R. Br. ex Schultes based on Periploca emetica Retz. The combination was not effectively
published in Secamone until 1820; however it is quite clear that Brown (1810b) included
the taxon it refers to within his concept of Secamone, despite his not formally transferring
the name. Hence it is not considered necessary to relectotypify with a type specimen of
one of the names published in Brown (1810a).

Species of Secamone are widely distributed in Africa, India, south-east Asia and
New Guinea with two species (S. e/liptica R. Br. and S. ovata R. Br.) having been
described from Australia (Brown 1810a). No critical account of Secamone in Australia
has been provided since that of Bentham (1869) who accepted both of Brown’s species.
Bentham’s treatment was copied by Bailey (1900). Bentham commented that S. ovata,
the second of the two species described by Brown in his Prodromus, was doubttully
distinct, differing from S. e/liptica only in the broader, more ovate and obtuse or scarcely
acute, shortly petiolate leaves.

Materials and Methods

Herbarium material at BRI, BRIU, CANB, DNA, JCT, LAE, MEL, NSW (partial
holdings only), NT, PERTH, QRS and photographs or microfiches of types at BM, K,
G and P were examined. Information on material at B and L were obtained from the
Directors or relevant staff of those institutions. Material was collected or procured

70 Austrobatleya 3({1}: 1989

throughout the species’ range in Australia. In the citation of specimens, preference has
been given to fertile collections that demonstrate the variation in the species and cover
its geographic range. Material preserved in spirit is indicated by an *. No index to
collectors has been included as most material in Australian herbaria has now been
determined.

Taxonomic Treatment

Secamone R. Br., Prod. 464 (1810). Type: Periploca emetica Retz. (S. emetica (Retz.) R.
Br. ex Schultes) (lecto, fide Phillips 1951).
Asclepiadeae 44 (1810); Mem. Wern. Nat. Hist. Soc. 1: 55 (1811); Blume, Bijdr.
1049-1051 (1826); Decne, Ann. Sci. Nat. Bot. 343 (1838); Decne in DC., Prod.
8: 500-504 (1844); Mig., Fl. Ned. Ind. 2: 471-474 (1856); Wight, Contr. bot.
India 60 (1834); Endl., Gen. pl. 8: 589 (1838); Walp., Ann. bot. syst. 3: 48 (1852):
Benth., Fl. austral. 4: 327-328 (1869); Benth. in Benth. & J.D. Hook., Gen. pl.
2: 146 (1876); J.D. Hook., Fl. Brit. India 4: 12-13 (1885); K. Schum. in Engl. &
Prantl, Nat. Pilanzenfam. 4(2): 261-263 (1897); Schitr., Bot. Jahrb. Syst. 50: 89-
90 (1914); Merrill, Enum. Philipp. fl. pl. 3: 342-343 (1923); Ridley, Fl. Malay
Penins. 2: 375 (1923); Backer & Bakhuizen van den Brink, Fl. Java 2: 254 (1965);
H. Huber, Rev. Handbk. Fl. Ceylon 4: 88-89 (1983).

Derivation of Name: From the Arabic name for one species.

Woody lianes with white latex. Stems slender to thickened with corky outgrowths, twining
or trailing, glabrous. Roots fibrous. Leaves flattened in cross-section, coriaceous or
herbaceous, opposite, margins entire, glabrous (occasionally pubescent), extrafloral nec-
taries lacking or present at lamina base, petiolate with small stipular structures at base.
Cymes appearing at nodes between the pair of leaves, |-many-flowered. Sepals 5, distinct,
generally with glands at base of lobes. Corolla deeply 5-lobed, subrotate, petals contorted
in bud, right hand edges overlapping. Staminal corona of 5 laterally compressed lobes,
adnate to the base of the staminal column. Stamens inserted at corolla-tube base, connate:
anthers with short incurved terminal membrane. Pollinaria each with 4 pollinia; pollinia
smooth, erect, 2 in each anther theca, globose or ellipsoid; corpusculum rounded; caudicles
flattened and very short. Gynostegium flat-topped to umbonate with obtuse or capitate
style-head enclosed by stamens; ovaries free, glabrous. Follicles lanceloid to ovoid or
angular, smooth or roughened; triangular to semi-terete in cross-section, often paired.
Seeds flat, ovate, brown, comose at germinating end only.

One species in Australia. Worldwide ca 20 species.

secamone elliptica R. Br., Prod. 464 (1810). Type: Carpentaria, Groote Islands [Queens-
land], 4 January 1803, R. Brown 8 [Iter Australiense 2871] (lecto (designated
here): BM(piece directly in middle of sheet); isolecto: BRI(AQ333111),
MEL(MEL1553064)).
Sprengel, Syst. veg. 1: 837 (1825); Decne in DC., Prod. 8: 504 (1844); Benth., FI.
austral. 4: 327 (1869); Bailey, Cat. pl. Queens]. 30 (1890); Queensl. fl. 3: 997-998
(1900); Compr. cat. Queensl. pl. 325 (1913); Maiden & Betche, Census New South
Wales pl. 174 (1916); Williams & Harden, Rainforest Climbing pl. 39 (1980),
Jacobs & Pickard, Pl. New South Wales 70 (1981); Williams, Students fl. N.E.
New South Wales 5: 721-722 (1984); Williams, Native pl. Queensl. 2: 258-259
(1984); Green, Census vasc. pl. Western Austral. 145 (1985); Dunlop, Checklist
vasc. pl. Northern Territory 9 (1987).

Secamone ovata R. Br., Prod. 464 (1810) synon. nov. Type: Coen River, Carpentaria
[Queensland], 6 November 1802, R. Brown 9 [Iter Australiense 2870] (lecto
(designated here): BM(2 sheets); isolecto: CANB).

Sprengel, Syst. veg. 1: 837 (1825); Benth., Fl. austral. 4: 328 (1869); Bailey, Cat.
pl. Queensl. 30 (1890); Queensl. fl. 3: 998 (1900); Compr. cat. Queensl. pl. 325
(1913).

Secamone lanceolata Blume, Bijdr. 1051 (1826) synon. nov. Type: Java (fide Bak-
huizen van den Brink 1950).
Bakhuizen van den Brink, Blumea 6: 371 (1950); Backer & Bakhuizen van den
Brink, Fl. Java 2: 254 (1965).

"~~ ‘i 0 hn ee ee te te ee wwe een ‘en nne PWM EB WY By WB a a Wwe MY MM wt Be Met Haw Ba HI Why my Hy Hy yy Hp Hi Hy Hy My Hy Dy Hy yy HHH yy Hy Hh hy Hy HI yy HH y Hy HH HH, I, yy MH HH hy Hy mm peng ii ni iin I I EM I MM My Be a tH Ma Ma Ba BB Me WW Bh wt a Ba Mh MB Mh aM MMM WB WP WWM Bo Mao wh WM a Ma Mh MW WW Wie Pon eee 2 ME Yon OnE OEE Yoo OEE REE 1 A Pe Ph 17 ns Me PO Ph Ph i Pn Mah A a Py By Py Pm Mo Mo i Py ath eM Pe Ma ty BI IM

yer abba Et at att a AEN A EE A oe tate NG, BASEN Ie diowaath abd thse aul MM bits ec Anta nab toma nate entation lant dita a tS EASA le at Nt

SAE RRA ats Bb AE a ie a BS a an a Ne BM ian tie abana ak

hs Bh

71
iew
ing

lv

. apica

o
x
te
ie
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=)

—

le
E

i corona with style-head protud

fk
rai, “es . “ a, et Ten, O8
ve i PART 4 dt baat an re ppcttl =
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7 Sf

d glands at sepal bases x

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o¢ "5

+e
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7
4

iew of stam
(c = corpuscie) X 70. All from Forster 988.

Ing OVaTFICS an

When

ae
b 4
* Poa yim ll

> w

hoot with little branched cymes X |. B. leaf X |

i corona X 40. G. side v

Ind

iptica
A. flowering s
| view of stam

iptica

as

»

=
+

. apica

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ete te at

~~?

inarla in rotating sequence

Fig. 1. Secamone elliptica:
D. flower with gynostegium and corolla removed show

Forster & Harold, Secamone ell
of flower X 10. F
x 40. H. Pol!

72 Austrobaileya 3(1): 1989

Secamone micrantha (Decne) Decne, Etud. Ascl. Ann. Nat. (1838), in DC., Prod.
8: 501 (1844) synon. nov. Tylophora micrantha Decne, Nouv. Ann. Mus. Hist.
Nat. 3: 377 (1834). Type: Timor, 1821, Spanoghe s.n. (syn: G, 3 sheets (photo!);
isosyn: K(photo!), MEL(MEL1553191); Timor, Zippel s.n. (syn: G(microfiche!);
isosyn: K(photo!),P(photo!); Java, Zollinger s.n. (syn: P(photo!); [Java] Soerakarta,
1802-1818, 7. Horsfield s.n. Gsosyn: K(photo!)).

Mia., Fl. Ned. Ind. 2: 472 (1856); Craib, Bull. Misc. Inform. 1911: 415 (1911);
Cost. in Lecomte, Fl. Indo-Chine 4: 41 (1912); Ridley, Fl. Malay Penins. 2: 375
(1923); Tsiang, Sunyatsenia 4: 57 (1939).

Secamone attenuata Decne in DC., Prod. 8: 501 (1844) synon. nov. Type: Philippine
Islands, Island of Mindoro, H. Cuming 1536 (holo: G; 1so: BM,K(2 sheets,
photo!),P(2 sheets, photo!).

Mia., Fl. Ned. Ind. 2: 472 (1856); Vidal, Phan. Cuming. philipp. 127 (1885),
Revis. pl. vasc. filip. 187 (1886); Merrill, Enum. philipp. fl. pl. 3: 342 (1923).

Secamone multiflora Decne in DC., Prod. 8: 501 (1844) synon. nov. Type: Philippine
Islands, Province of Cagayan, Luzon, H. Cuming 1284 (iso: BM,
BRI(AQ333115),KQ sheets, photo!),L,MEL(MEL1553192),P(2 sheets, photo!)).
Mig., Fl. Ned. Ind. 2: 472 (1856); Vidal, Phan. Cuming. philipp. 127 (1885),
Revis. pl. vasc. filip. 187 (1886); Merrill, Enum. philipp. fl. pl. 3: 342 (1923).

Secamone papuana Warb., Bot. Jahrb. Syst. 18: 206 (1894). synon. nov. Type: “Dutch
New Guinea”, 1888, O. Warburg 21322 (lecto (here designated): BRI; isolecto:
A(n.v.)).

Secamone flavida Schitr., Bot. Jahrb. Syst. 50: 89-90 (1914) synon. nov. Type:
Nordéstl. Neu-Guinea: Hane in den Waldern am Minjem-Thor, February 1908,
R. Schlechter (B f).

Cynanchum dichasiale O. Schwarz in Feddes Repert. Spec. Nov. Regni Veg. 24: 94
(1927). Type: Bleeser 586 (syn: B, n.v.; isosyn: NSW); Darwin, Bleeser 244 (syn:
B, n.v.; isosyn: MEL).

Blake, Proc. Roy. Soc. Queensland 59: 168 (1948).

[Secamone emetica auct. non (Retz.) R. Br. ex Schultes: F. Muell., Fragm. 5: 161
(1866). |

Woody liane. Stems green when young, with age there develops grey, hard, fissured bark;
up to 10 cm diameter, up to at least 10 m long, twining, much branched. Leaves
herbaceous, glabrous or with sparse indumentum of uniseriate hairs; lamina elliptic,
narrow-elliptic, lanceolate or ovate, 20-60 mm long, 6-35 mm wide, tip acuminate or
acute, base acute or rounded; petiole 3-8 mm long, 0.5-1 mm diameter. Cymes with
few to many branches, borne on first 4 or 5 nodes below apex, up to 2 cm wide; peduncle
2-3 mm long, 0.75 mm diameter, finely pubescent with uniseriate hairs. Flowers 2 mm
long, 2-5 mm diameter; pedicels 2-4 mm long, | mm diameter, finely pubescent with
uniseriate hairs. Sepals 0.75-1.5 mm long, 0.75-1.5 mm wide, ovate, finely pubescent,
0-3 glands present at base. Corolla lobes reflexed, with a concave base, obtuse, glabrous,
greenish yellow, 1.5-2 mm long, 1.25-1.5 mm wide. Staminal corona 1.25-1.5 mm long,
1.25~1.5 mm diameter; corona lobes adnate to base of anthers, curving inwards, subulate-
cylindrical, 0.25-0.75 mm long, 0.2-0.25 mm diameter. Staminal column 1.5 mm long,
| mm wide; anther appendages 0.25 mm long, 0.25 mm wide; slit between anther wings
0.5-0.75 mm long. Style-head projecting past anther connectives, flattened globose, 0.25-
0.5 mm long, 0.3 mm diameter. Ovaries 0.5 mm long, 0.5 mm wide (both). Pollinaria
0.20 mm long, 0.20 mm wide, golden; pollinia (singly) 0.12-0.13 mm long, 0.07-0.08
mm wide; corpusculum 0.10-0.11 mm long, 0.07-0.10 mm wide; caudicles 0.10 mm
long, 0.02 mm wide. Follicles terete-ovoid, 40-100 mm long, 7-13 mm diameter; seed
ovate, tan, 7-10 mm long, 2-3 mm wide; coma 23-35 mm long.

Selected specimens: Philippine Islands. MINDANAO: Buayan, Cotabato Province, Apr 1932, Ramos & Edano
85131 (BRI). Timor. Port Timor, E of Manotutu, Dec 1953, van Steenis 18007 (CANB). Papua New Guinea.
Tarara, Wassi Kussa, Dec 1936, Brass 8501 (A,BRI); Tavai Ck area, ca 43 miles [71 km] SE of Port Moresby,
9°44’S, 147°26’E, May 1967, Pullen 6878 (A,;CANB,L,LAE); Yule Is, 8°50’S, 146°32’E, Womersley & Simmonds
NGF7107 (CANB,LAE). Australia. Western Australia. Broome Harbour, [7°58’S, 122°14E, Jan 1971, Allan 597
(CANB,PERTH): Walsh Pt, Port Warrender, Admiralty Gulf, 14°34’S, 125°S1’E, Jan 1982, Kenneally 7824
(CANB,PERTH). Northern Territory. East Pt, Darwin area, 12°24’S, 130°56’E, Jan 1988, Stobo [AQ408478]
(BRI*); Channel Pt, 13°10’S, 130°08’E, Nov 1982, Wightman 329 & Dunlop (CANB,DNA*,MEL,PERTH).
Queensland. Cook District: Mapoon -— lily lagoon, S of Cullen Point (Port Musgrave), 11°59’S, 14i°52’E, Jun
1981, Mforton AM1265 (BRI,MEL): Eastern Beach beyond Pearling Stn, Paki Is, 10°44’S, 142°13’S, Jan 1987,
Guuness 2100 (BRI*). NortH KenNepy District: 40 Mile Scrub, 18°O1’S, 144°45’E, Jan 1972, Hyland 5888

Forster & Harold, Secamone elliptica 73

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Fig. 2. Secamone elliptica: A. trip of flowering stem & two other separate cymes demonstrating much branched

nature X 1. B. leaf X 1. C. single flower X 10. D. flower with gynostegium and corolla removed showing ovaries

and glands at sepal bases X 20. E. apical view of flower X 10. F. apical view of staminal corona X 40. G. side

bei of ae corona with style-head protuding x 40. H. pollinaria in rotating sequence < 70. All from
unness

PEER EA SERB aE

74 Austrobaileya 3(1): 1989

(BRI,CANB,QRS), Fletcher Ck, 2 km SW of Mt Keelbottom, 19°49’S, 146°0S’E, Mar 1988, Forster & Bolton 3620
(BRI*). MITCHELL District: near Jericho, Apr 1946, Clemens [AQ216898] (BRI). LEICHHARDT District: Apis
Ck, ca 40 km N of Marlborough, 22°40’S, 149°35’E, Apr 1975, Craven 3145 (BRI,CANB); Wolfang Peak, 22°33%S,
147°50’E, Mar 1988, Forster 3609 (BRI). Port Curtis District: Dan Dan S.F., 24°LO/’S, ISP°OS’E, Dec 1987,
Gibson [AQ408480] (BRI*). BURNETT District: 4.5 km NNE of Monogorilby, 26°01’S, [SP°OVVE, Dec 1981,
Forster 988 (BRI*). Wipe Bay District: Mt Glastonbury, 26°14’S, 152°29'E, Jan 1987, Forster & Sharpe 2875
(BRI*). MorRETON District: Flinton Hill, Worlds End Pocket, 27°31’S, 152°45’E, Dec 1987, Forster 3309 (BRI*).
New South Wales. Sandiland Ranges, Nov 1904, Boorman (NSW). New Caledonia. Magenta, Noumea, Dec 1960,
McKee 7835 (CANB); Ouen Toro, hill at S end of Noumea, Jan 1982, McPherson 4560 (BRI; MO n:.¥.).

Distribution, habitat and ecology: S. e/liptica is widely distributed in the Philippine
Islands, Malesia, New Guinea, in northern and eastern tropical to subtropical Australia
(Map 1), and New Caledonia. S. el/liptica occurs as a woody liane in the canopy of
numerous rainforest types, including semi-evergreen vine thicket, araucarian microphyll
vine forest, araucarian notophyll vine forest, notophyll vine forest and low microphyll
vine forest, on a variety of soil types including red krasnozems, red euchrozems, black
earth and quartz sands, and on different types of rock outcrops. It is commonly used as
a host plant by larvae of Euploea core corinna (MacLeay) (Common Crow Butterfly)
(Sankowsky 1975), which feeds on a wide range of Asclepiadaceous hosts (Scheermeyer
& Zalucki 1985; Forster 1987).

Seed germinates within 3-4 days of planting if fresh and the seedlings (Fig. 4B)
make rapid growth, reaching 50-80 cm in height during the first growing season.

Phenology: Flowering occurs from October to April, fruiting from March to June.

Notes: The type sheet of S. el/liptica at BM has two labels in Brown’s handwriting
attached. One has “Cynanchum glabellum, Brown No. 8, Carpentaria main opp...
Groote Islands” and the other has “Cynanchum, Brown No. 9, Coen River’’. At the
base of the sheet is written ““Secamone elliptica R. Br.” in Brown’s handwriting. There
is no way of telling whether the sheet is a mixed collection of material from two different
localities as there are several discrete pieces of material attached. Hence the piece directly
in the middle of the sheet has been selected to preserve the application of Brown’s name
to an elliptic leaved specimen.

The localities for Cuming’s type collections of S. attenuata and S. multiflora are
taken from the listing of Merrill (1915). The isotype sheets at K of S. attenuata have
‘Prov. Batangus, Luzon’ and of S. multiflora have ‘Prov. Albay, Luzon’ on the labels.
According to Merrill, all of Cuming’s collections were distributed with labels stating
“Ins. Philippinae 1841” although many did not actually come from The Philippines.
According to Merrill (1903), complete sets of Cuming’s collections exist at B (presumably
now destroyed), BM, K and P, with partial sets at G, L and a number of other herbaria.

Warburg (1894) in describing S. papuana did not provide a formal description,
however his sentence “Corona squamae 5, tubo stamineo nune brevis-sime nune alte
adnatae, a latere compressae, apice liberae rectae vel incurvae.” may be construed as a
diagnosis. No specimen was cited and the above collection of Warburg’s which is labelled
“Secamone papuana Warb.” is selected as lectotype.

S. elliptica is a very variable species particularly with respect to leaf length, width
and shape as demonstrated by Figures 1-3. There appears to be no strong geographical
correlation of this leaf variation within Australia, although a greater proportion of
herbarium records from the southern parts of the range tend to have elliptic to narrow-
elliptic leaves. Examination of any population in the field generally reveals branches
with leaves that vary from narrow-elliptic to elliptic to broadly lanceolate-elliptic or
ovate, depending on the conditions of growth. Similarly there is no geographic correlation
of plants with either solitary or compound cymes as can be seen from Figures la and
4a, both from plants in the southern part of the range with elliptic leaves. Some minor
variation in floral characters such as corona size, corona lobe size and shape, pedicel
length, sepal size, corolla size and calycine gland number occurs (Figures 1-3) but this
is no more than occurs in many taxa of Asclepiadaceae and shows no geographical or
ecological correlation. Hence there is little basis on which to recognise 5. ovata, 5S.
elliptica, S. micrantha, S. multiflora, S. attenuata, S. lanceolata, or S. flavida as distinct
species. The name S. elliptica is applied here because, of the earliest two names available
(the other being S. ovata), it has been more commonly used.

Conservation status: The species is widespread and not endangered in any way.

Forster & Harold, Secamone elliptica

Fig. 3. Secamone elliptica: A. \eaf (tip lost) * 1. B.

RR ge ll car qa,

a
‘anor
")

4
wed A

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single flower X 10. C. flower with gynostegium and corolla

removed showing ovaries and glands at sepal bases X 20. D. apical view of flower X 10. E. apical view of staminal

corona X 40, F. side view of staminal corona with
Stobo AQ440519.

style-head protuding X 40. G. pollinarium x 70. All from

CeO Oe hi eee GOO COOL RCCL CoC Ck COCO LCCC LLCO LCi ee LoL ec CL COL iL Le ce ee ee ee Te eC CTC CCIM. TC TCIiCrrrrrerrrerrrr rc rere rrrerrrcrerrrrrrrr rc Trerrirrrrrirreriirrrrr iter eT eee ee erere rere rrr errr errr err rrrny

rei et

‘enna te

716 Austrobaileya 3(1): 1989

Fig. 4. Secamone elliptica: A. tip of flowering stem with elliptic leaves and much branched cyme X I. B. seedling
x 1. C. follicles X 1. D. seeds X 1. A, Forster 3309; B, Forster 310: Stony Ck, 25°29’S, 131°54’E, C-D, Forster
533: Mt Blandy, 25°24’S, 151°46’E.

Acknowledgements

The Directors of BRI, CANB, DNA, JCT, LAE, MEL, NT, PERTH and QRS
allowed full access to specimens, either at their institutions or on loan. Partial holdings
were allowed on loan from NSW. The Director of L provided information on type
material at that institution. Dr B. Leuenberger (B) provided information on type material
of R. Schlechter held at that institution. Mr L. Pedley (BRI) arranged for the loan of
specimens. Dr G.P. Guymer (BRI) and Dr J.G. West (CANB) whilst Australian Botanical
Liaison Officers at Kew arranged for photographs to be taken of various types. Special
collections were made or arranged by Mesdames C. Cox, A. Gunness, J. Stobo, R.
Tingey and Mr N. Gibson. Some collections were made on various field trips with
Messrs L.H. Bird and M.C. Tucker and Dr M.P. Bolton. The Australian Biological
Resources Study provided partial funding in 1988. Mrs E.M. Ross (BRI) provided
constructive comments on the manuscript. All of this assistance is gratefully acknowledged.

References
BAILEY, F.M. (1900). Secamone. In The Queensland Flora 3: 997-998. Brisbane: Government Printer.
BENTHAM, G. (1869). Secamone. In Flora Australiensis 4: 327-328. London: L. Reeve & Co.
BROWN, R. (1810a). Prodromus Flora Novae Holiandiae et Insulae van Diemen. New York: J. Cramer.

BROWN, R. (1810b). On the Asclepiadeae. [a natural order of plants separated from the Apocineae of Jussieu].
London: R. Brown. [A preprint of Brown [811.]

Forster & Harold, Secamone elliptica

ORT a =
t 7 |

ir :
; 4
: |
|

| ;
|

en
Mr a -
wees
jt 2 ed
ne 2 4

, 7

=
yaaa
ee
T
a2 a
nm :
a “
‘ ee el
enn, el
ee al
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35°

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be t 3 A
ae" *

Map 1. Distribution of Secamone elliptica in Australia mapped at 1:250 000 grid scale.

BROWN, R. (1811). On the Asclepiadeae, a natural order of plants separated from the Apocineae of Jussieu
Memos of the Wernerian Natural History Society 1: 12-78.

FARR, E.R., LEUSSINK, J.A. & STAFLEU, F.A. (eds) (1979), Index Nominum Genericorum Plantarum. Utrecht/
The Hague: Bohn, Schelterma & Holkema/ Dr. W. Junk Publishers eV.

FORSTER, P.I. (1987). New host records for Euploea core corinna (Macleay) (Nymphalidae). Journal of the
Lepidopterist's Society 41: 354-355.

MABBERLEY, J. (1985). Jupiter Botanicus - Robert Brown of the British Museum. Braunshweig: J. Cramer.

MERRILL, E.D. (1903). Botanical work in the Philippines. Bureau of Agriculture, Bulletin No. 4. Manila: Bureau
of Printing.

MERRILL, E.D. (1915). Genera and species erroneously credited to the Philippine Flora. The Philippine Journal
of Science, C. Botany. 10: 171-194.

PHILLIPS, E.P. (1951). The Genera of South African Flowering Plants. 2nd Edition. Cape Town: Government
Printer.

PURI, V. & SHIAM, R. (1966). Studies in floral anatomy. VII]. Vascular anatomy of the flower of certain species
of the Asclepiadaceae with special reference to corona. dgra University Journal of Research 15: 189-216.

SAFWAT, F.M. (1962). The floral morphology of Secamone and the evolution of the pollinating apparatus in
Asclepiadaceae. Annals of the Missouri Botanical Gardens 49: 95-119.

SANKOWSKY,. G. (1975). Some new food plants for various Queensland butterflies. -ustralian Entomological
Magazine 2: 55-56.

SCHEERMEYER, E. & ZALUCKI, M.-P. (1985). Food plant records of Euploea core corinna (W.S. Macleay)
with some notes on larval coloration. Australian Entomological Magazine 11: 87-90.

qT]

78 Austrobaileya 3(1): 1989

SCHILL, R. & JAKEL, U. (1978). Beitrag zur Kenntnis der Asclepiadaceen-Pollinarien. Tropische und subtropische
Pflanzenwelt 22: 1-122.

WARBURG, O. (1894). Plantae Hellwigianae. Beitrag zur Flora von Kaiser Wilhelms-Land. Botanische Jahrbticher
fiir Systematik Pflanzengeschichte und Pflanzengeographie 18: 184-212.

Accepted for publication 10 March 1989

Austrobaileya 3(1): 79-99 (1989) 79

STUDIES IN AUSTRALIAN GRASSES: 4%.
TAXONOMIC AND NOMENCLATURAL STUDIES
IN AUSTRALIAN ANDROPOGONEAE

B.K. Simon

Queensland Herbarium, Meiers Road, Indooroopilly, Qid 4068

Summary

Eight new species, namely Cymbopogon dependens, Eulalia annua, Ischaemum albovillosum, [. tropicum, Mnesithea
pilosa, Schizachyrium mitchelliana, Thaumastochloa heteromorpha and Vetiyeria rigida are diagnosed and described.
New combinations are Bothriochloa bladhii subsp. glabra based on Andropogon glaber Roxb., Dichanthtum sericeum
subsp. Aumilius based on Dichanthium humilius J. Black, Dichanthtum sericeum subsp. polystachyum based on
Andropogon sericeus var. polystachyus Benth. and Hyparrhenia rufa subsp. altissima based on Hyparrhenta altissima
Stapf. Synonyms created for the first time are Andropogon affinis R. Br., A. chrysatherus F. Muell., A. sericeus
var. mollis Bailey (and names based on these basionyms) under Dichanthium sericeum (R. Br.) A. Camus,
Andropogon sericeus var. geniculatus Bailey and A. sericeus forma micranthus Domin under Dichanthium sericeum
subsp. Aumilius and Hyparrhenia quarrei Robyns under H. Airta (L.) Stapf. Dimeria chloridiformis (Gaudich.)
Schumann & Lauterb. is recognized as the correct name for D. ciliata Merr., Eulalia aurea (Bory) Kunth for E.
fulva (R. Br.) Kunth and &. geniculata Stapf, Mnesithea rottboellioides (R. Br.) Koning & Sosef for Coelorachis
rottboellioides (R. Br.) A. Camus, Mnesithea granularis (L.) Koning & Sosef for Hackelochioa granularis (L.)
Kuntze, Mnesithea annua (Lazarides) Koning & Sosef for Heferopholis annua Lazarides, Mnesithea formosa (R.
Br.) Koning & Sosef for Rottboellia formosa R. Br. and Polytrias diversiflora (Steudel) Nash for P. amaura (Buse)
Kuntze.

As a result of research undertaken for the preparation of the account of Andro-
pogoneae for the Flora of Australia, a number of new species require formal description
and a few nomenclatural changes are necessary. In the following account they are treated
alphabetically by genus. The descriptions of the new species are extracted from the
database of the Andropogoneae compiled for the Flora using the DELTA system (Dallwitz
& Paine 1986).

New Taxa and New Combinations

BOTHRIOCHLOA Kuntze

Bothriochloa bladhii subsp. glabra (Roxb.) B. Simon, comb. nov.

Andropogon glaber Roxb., Fl. ind. 1: 271 (1820); Amphilophis glabra (Roxb.) Stapf
in Prain, Fl. trop. Afr. 9: 174 (1917); Bothriochloa glabra (Roxb.) A. Camus, Ann.
Soc. Linn. Lyon, n.s. 6: 164 (1931). Type: Bengal, India, W. Roxburgh (holo:
BM!(photo BRI,K); itso: K!(photo BRD).

Specimen examined: Queensland. MORETON DISTRICT: Wendy Allison Park, Brisbane, Apr 1977, Bisset B184 &
B185 (BRI).

Work on the genetic barriers of Bothriochloa bladhii (under B. intermedia) (de
Wet & Harlan 1966; Faruqi 1969) has established that this species is capable of crossing
with other species of Bothriochloa and species of the related genera Dichanthium and
Capillipedium. This promiscuous character of B. bladhii has led some workers to use
the biological species concept to unite the three genera taxonomically (de Wet & Harlan
1966). However I have previously given reasons why I do not consider the wisdom of
such a move with respect to Bothriochloa and Dichanthium (Simon 1982), namely the
morphological differences of spikelet shape, the presence or absence of a translucent
mid-line in the rachis internodes and pedicels and differences in phytogeography. The
species Bothriochloa glabra has been sunk under B. bladhii (Faruqi 1969; Clayton &
Renvoize 1982) following the trend of regarding B. bladhii as a very variable species.
Another point of view is that B. glabra is a hybrid between B. bladhti and Capillipedium
parviflorum and furthermore that C. spicigerum 1s the product of a back-cross between
B. glabra and C. parviflorum (de Wet 1987). In view of the fact that B. glabra does not
occur naturally in Australia I feel these assumptions cannot be made. However I also

* continued from Austrobaileya 2(3): 281-283 (1986)

80 Austrobaileya 3(f): 1989

feel that information is lost by placing A. glabra into synonymy with &B. bladhti as the
introduced B. glabra has slightly smaller spikelets (3-3.5 mm long) than the native B. bladhii
(3.5-4 mm long) and always has pitted spikelets whereas this character 1s a fairly rare
feature in B. bladhii. For this reason I propose the rank of subspecies for B. glabra.
Although this entity has been introduced into cultivation a number of times there appears
to be only one record of it having become naturalised. This was in a Brisbane suburban
park near an old plant introduction field station.

CYMBOPOGON Sprengel

Cymbopogon dependens B. Simon, species nova afhnis C. ambiguo A. Camus et C.
procero (R. Br.) Domin sed foliorum laminis ad 3 mm latis, filiformibus vel
perangustis, culmis subtilibus et dependentibus differt. Typus: Katherine Gorge,
Northern Territory, 8 March 1964, 4. Lazarides 7036 (holo: BRI'(2 sheets BRI
139135 & BRI 139136); iso: CANB!,DNA!K'(photo BRDI),MEL!). Fig. 1.

Culms 90-120 cm tall, 3-5-noded below the inflorescence, delicate and drooping,
internodes smooth, glabrous; nodes glabrous. Leaf sheaths longer than the internodes,
smooth, glabrous, not rolling up at maturity. Ligules to 1 mm long. Collar glabrous.
Leaf blades gradually narrowed to the base, flat, smooth above, glabrous, 15-30 cm long,
0.5-2.5 mm wide. Inflorescence occupying more than 1/5 the culm length, 17-25 cm
long, 3-5 cm wide, + interrupted, 3-5-noded, branched to the 3rd degree; inflorescence
branches mostly exceeding the internodes. Spatheoles very narrowly elliptic, glabrous,
smooth, 2-4 mm long, 1.5-—2 mm wide. Peduncle hairy towards the tip, 8-11 mm long,
1/3-1/2 as long as the spatheole. Racemes distinctly hairy, with hairs 4-7 mm long, not
reflexed, 20-25 mm long, 3-5-jointed. Joints and pedicels linear, concave ventrally,
convex on the back, 2-2.5 mm long, ciliate on the margins and villous on the back,
nearly equal. Sessile spikelet 4.5-5 mm long, 5-6 times as long as wide, very narrowly
ovate-elliptic, awned; longer callus hairs to 1.2 mm long. Lower glume flat or concave
on back, ca | mm long, 2-keeled throughout, with 0-4 intracarinal nerves, flat to shallowly
concave. Upper glume 3-nerved. Lower lemma oblong, a little shorter than the glumes.
Upper lemma 3/4 as long as the lower lemma, lobed to I/2 the length. Awn 17-20 mm
long. Column 6-8 mm long. Pedicelled spikelet neuter, narrow linear to narrowly ovate,
ca 2 mm long (rudimentary).

Specimens examined: Western Australia. GARDNER District: Hidden Valley, Kununurra, Pullen 10.84!
(CANB,NSW,PERTH,WIR). Northern Territory. DARWIN AND GULF: Woolaning, Dunlop 5925 & Craven
(CANB,DNA,K,L); 19 mi NE of Katherine, Feb 1965, Hilson 333 (BRI,CANB,DNA,K,L,NSW,US); 4 mi W of
Pine Creek, Lazarides 237 & Adams (CANB,NT); Katherine Gorge, Mar 1980, Simon 3667, Cousins & Grosvenor
(BRI). CENTRAL SoutH: 26 mi NE of Docker River Settlement, Laiz 872 (CANB,DNA).

This species is distinguished from the related C. ambiguus and C. procerus by its
small stature and decumbent or weakly erect habit. It inhabits sandstone cliffs in various
isolated regions of the Northern Territory and Western Australia. The above three species
are closely related, differing only 1n size and direction of culm growth. Some plants of
C. procerus are extremely robust, growing to heights of up to 2.5 m with leaf blades to
{5 mm wide whereas plants of C. dependens may have culms as short as 40 cm and
leaf blades never more than 3 mm wide. The quantitative differences may be a reflection
of the ecology of the habitats where the forms occur. C. procerus grows only in the
monsoon tropical areas of northern Australia; C. ambiguus 1s found in more arid regions
of the Australian subtropics, although collections have also been made in the tropics;
C. dependens has been collected from both tropical and subtropical regions. The differences
in spikelet morphology between C. ambiguus and C. procerus given by Blake (1974)
were not reflected in specimens examined when the Flora of Australia account was being
prepared. It is also interesting that Blake regarded the fragile specimens represented by
Lazarides 7036 and Wilson 333 (placed here with C. dependens) as a diminutive form
of the robust species C. procerus and not of the shorter species C. ambiguus. The close
relationship between C. ambiguus and C. procerus is reflected by the fact that the name
C. exaltatus, a taxonomic synonym of C. procerus, has been misapplied to C. ambiguus
in the past. In order to find the true relationship between these entities experimentation
is required. In the meantime I have decided to recognize them at species rank.

In general the taxonomic situation in Australian species of Cymbopogon is not as
clear as that presented by recent workers (Blake 1974, soenarko 1977). There appear to
be a number of situations of blurring of species boundaries and the existence of hybrids

Simon, Australian Andropogoneae

. One sheet of holotype of Cymbopogon dependens.

82 Austrobaileya 3(1): 1989

has been postulated. Examples of these have been cited e.g. hybrids between C. refractus
and three other species (C. obtectus, C. bombycinus and C. queenslandicus) cited by
Blake (1974), hybrids between C. ambiguus and C. bombycinus cited by Soenarko (1977)
and a postulated hybrid between C. ambiguus and C. queenslandicus (S.T. Blake 8353A)
from Castle Hill, Townsville, annotated by Blake in the Queensland Herbarium. The
boundary between C. bombycinus and C. obtectus is not clearly demarcated and there
are populations that are intermediate in the interzone between the wetter and drier
regions in which the two species occur. Some specimens which fall into this category
have curled-up old leaf-blades of C. bombycinus but spikelets of C. obtectus (Lloyd
NSW183537 (NSW) from Coonabarabran, N.S.W., Crisp 1542 (AD,CBG,NSW,L,US)
from near West Wyalong, N.S.W., Blake 10742 (BRI,NSW) from Noondoo, Queensland,
Chippendale 1738 (DNA,NSW) from Hamilton Downs, N.T. and 7. & J. Whaite 4159
(NSW) from near Geraldton, W.A.).

DICHANTHIUM Willimet

The Dichanthium sericeum complex, known commonly as Queensland blue grasses,
has been a problem group for some time with regards to its taxonomy (Blake 1969; de
Wet & Harlan 1962; de Wet & Richardson 1963; de Wet & Singh 1964). The similarity
between the entities was recognized by de Wet and Singh (1964) who stated that “these
species appear to form a closely related complex characterized by fasciculate, sessile or
subsessile racemes’. In a taxonomic study of the group Blake (1969) went to some
lengths to justify the recognition of three species D. sericeum, D. affine and D. tenuiculum.
Sega re-examination of type material has resulted in my perceiving the situation to

e as follows:

1. Blake’s “D. affine” really consists of two entities, one of which is a delicate annual
species found in arid inland areas and formerly recognized as D. humilius and
the other, to which the name D. affine has been given, is really a more slender
state of D. sericeurm and found together with the latter species in the black soil
plains of the subtropics.

2. The lectotypes of Dichanthium sericeum and D. affine are from a mixed gathering on
the Paterson River, New South Wales and mounted on a single sheet at BM
(Blake 1969). There is a discernible difference on the sheet between the inflo-
rescence of D. affine (labelled ““A”’ on the sheet) and the other six of D. sericeum,
the former having shorter racemes and awns and smaller spikelets. However the
fact that they were collected in the same gathering may either be a reflection of
phenotypic plasticity or the result of genetic variation of a Mendelian nature. At
this stage therefore I am not giving taxonomic recognition to this more slender
form, but including it within the variability of D. sericeum subsp. sericeum.

3. The tropical entity with many racemes was known as Dichanthium superciliatum
until Blake (1969) made the new combination D. tenuiculum based on an earlier
basionym. When I examined the type of Andropogon tenuiculus Steudel the
presence of only 2-4 racemes led me to believe Blake was wrong in placing this
entity together with the many-racemed D. superciliatum. At the time I thought
the specimen belonged with typical D. sericeum and I placed a det. slip on the
type with this name (at that stage I was calling it D. sericeum var. sericeum),.
However recent field work in tropical Australia and an examination of more
specimens has revealed that raceme number is not a good diagnostic character
for this tropical entity, nor is the other morphological character of the hairs being
arranged in a subapical fringe (de Wet & Harlan 1968) as both taxa display this
character. I have found more reliable distinguishing characters to be habit and
the way the hairs of the subapical fringe are arranged. The tropical entity is
usually a robust annual with the spikelet hairs being arranged in a distinct erect
fashion from the spikelet surface whereas typical D. sericeum 1s always a perennial,
sometimes fairly delicate, and the spikelet hairs are not as distinctively erect from
the spikelet surface. The type of Andropogon tenuiculus has no base and so it
was not possible to tell its habit; the erect hairs and the fact that it was collected
from the tropical Philippines however, has persuaded me to follow Blake’s original
placing of this entity. A physiological difference between the two has been noted
(Tothill 1977, 1981) in that the tropical entity flowers under short-day conditions
late in the season whereas D. sericeum sens. Strict. is day-neutral. Tothill postulates

Simon, Australian Andropogoneae 83

speciation of the latter from the former as an evolutionary development away
from short-day contro! of flowering in a similar manner to that in some other
members of the Andropogoneae in their southern migration.

4, Although the taxa of the D. sericeum complex are fairly well genetically isolated (de
Wet & Harlan 1968), there exist in herbarium collections specimens of an
intermediate nature which are difficult to place with certainty into one taxon or
another. In view of this dilemma it seems to me that the most satisfactory
nomenclatural solution at this stage is to treat the whole complex as one species,
with three subspecies in the manner outlined formally. At an earlier stage in my
studies I selected the varietal rank to separate “D. affine” (in the Blake sense)
from D. sericeum, whereas the decision to use the subspecies rank for D. tenuiculum
was taken at the outset.

With the establishment of D. affine as a synonym of D. sericeum subsp. sericeum,
the full synonymy for D. sericeum becomes:

Dichanthium sericeum (R. Br.) A. Camus, Bull. Mus. Hist. Nat. (Paris) 27: 549 (1921);
Andropogon sericeus R. Br., Prodr. 201 (1810). Type: Paterson R., N.S.W., &.
Brown [6178, excl. part A] (lecto: BM!(photo BRI,K); isolecto: E!(photo
ag ara & fragment BRI), fide S.T. Blake, Proc. Roy. Soc. Queensland
80: 67 (1

Dichanthium sericeum subsp. sericeum

Andropogon affinis R. Br., Prodr. 201 (1810); Sorghum affine (R. Br.) Kuntze, Revis.
gen. pl. 2: 791 (1891); Dichanthium affine (R. Br.) Camus, Bull. Mus. Hist. Nat.
(Paris) 27: 549 (1921). Type: Paterson R., N.S.W., R. Brown [6178, part A] (lecto:
BM!(photo BRI,K); isolecto: E!(fragment BRI, photo BRLK),K\(photo BRI), fide
S.T. Blake, /oc. cit.).

Andropogon iat ated F. Muell., Linnaea 25: 443 (1853). Type: Crystal Brook,
S.A., F. Mueller s.n. (syn: MEL!(fragment & photo BRD),P!(fragment BRI, photo
BRLK),S!(fragment BRI, photo BRI,K),W!(photo BRI); towards Rocky Creek,
S.A., F. Mueller s.n. (syn: MEL!(photo BRI),P!,S!).

Andropogon acutiusculus Hackel in DC., Monogr. phan. 6: 975 (1889); Dichanthium
acutiusculum (Hackel) A. Camus, Bull. Mus. Hist. Nat. (Paris) 27: 549 (1921).
Type: Port I Keppel Bay [Between Curtis Island and Facing Island, Qld], &.
Brown [6178 in part] (holo: W, n.v.(photo BRI); iso: BM!(photo BRI,K),BRI',E!,K!
(photo BRI),MEL!,P!(photo BRI)).

Andropogon sericeus var. mollis Bailey, Queensland Agric. J. 30: 316 (1913); Dichan-
thium sericeum var. molle (Bailey) de Wet & Harlan, Phyton 18: 13 (1962). Type:
Gindie, E.W. Bick 12 (holo: BRI!; iso: K!(photo BRI),MEL’).

Andropogon sericeus forma glaberrimus Domin, Bibhoth. Bot. 85: 267 (1915). Type:
Barcaldine, Qld, March 1910, K. Domin s.n. (syn: n.v.), Mareeba, Qld, K. Domin
s.n. (syn: K!(photo BRI).

Andropogon sericeus forma puberulus Domin, Biblioth. Bot. 85: 267 (1915). Type:
Chillagoe, Qld, February 1910, K. Domin s.n. (?holo: K!(photo BRI).

Andropogon sericeus forma ciliatus Domin, Biblioth. Bot. 85: 267 (1915). Type:
Jericho, Qld, February 1910, K. Domin s.n. (?holo: K!(photo BRI)).

Dichanthium caricosum subvar. racemosum Roberty, Boissiera 9: 165 (1960). Type:
Jericho, Qld, R. Simmons 69 (holo: G!(fragment BRI, photo BRIK); so: NSW).

[Andropogon annulatus auct. non Forssk.: F. Muell., Fragm. 8: 123 (1873)].

Dichanthium sericeum subsp. humilius (J. Black) B. Simon, comb. nov.

Dichanthium humilius J. Black, Trans. & Proc. Roy. Soc. South Australia 60: 164
(1936); Andropogon annulatus var. ?humilis Benth., Fi. austral. 7: 531 (1878).
Type: Charlotte Waters, N.T., March 1875, &. Giles s.n. (holo: MEL!(photo BRI)).

Andropogon sericeus var. geniculatus Bailey, Queensland Agric. J. 26: 128 (1911).
synon. nov. Type: Winton, Qld, February 1914, T.G. Wright s.n. (holo: BRI; iso:
CANB!,K!(photo BRI),MEL!).

84 Austrobaileya 3(1): 1989

Andropogon sericeus forma micranthus Domin, Biblioth. Bot. 85: 267 (1915) synon.
nov. Type: Between Winton and Barcaldine, March 1910, K. Domin s.n. (holo:
PR fide Blake (1969), n.y.).

Specimens examined: Western Australia. COOLGARDIE District: 40 km E of Mt Vetters Station, Kalgoorlie, Apr
1975, Mitchell sn. (NSW,PERTH). CARNARVON District: 94 km N of Minilya, Mar 1980, Simion 3783 & Stretch
(BRI). Austin District: Wiluna, July 1937, Afelville s.n. (PERTH). HaLt District: Fox River Station, Ma
1981, Glover FR 7 (BRI). Northern Territory. BARKLAY TABLELAND: Rockhampton Downs, Apr 1968, Nicholls
790 (BRI,DNA). CENTRAL NorRTH: Claraville Homestead, Jan 1971, Latz 1152 (BRI,DNA). CENTRAL SOUTH:
Andado Station, Apr 1977, Latz 6775 (BRI,DNA). South Australia. LAKE Eyre: Wangianna, Sep 1941, Cleland
s.n. (BRI). FLINDERS RANGES: 16 km NNE of Wooltana, Apr 1966, Eichler 18637 (AD,BRI). Queensland. BURKE
District: 40 mi SSW of Richmond, Jun 1954, Lazarides 4458 (BRI,CANB). GREGORY NORTH DISTRICT:
Currawilla, Jun 1949, Everist 3910 (BRI,K). New South Wales. NorTH WESTERN PLAINS: ‘Near Fort Bourke, Apr
1976, Simon 2959 (BRILL.NSW,K).

_ Dichanthium sericeum subsp. polystachyum (Benth.) B. Simon, comb. nov.

Andropogon sericeus var. polystachyus Benth., Fl. austral. 7: 530 (1878). Type:
Burdekin River, Qld, FE. Bowman s.n. (lecto (here designated): K!(photo BRI):
isolecto: MEL!(photo BRI)).

Andropogon tenuiculus Steudel, Syn. Pl. Glum. 1: 371 (1854); Dichanthium tenui-
culum (Steudel) S.T. Blake, Proc. Roy. Soc. Queensland 80: 69 (1969). Type:
Luzon, Philippines, H. Cuming 1398 (holo: P!\(photo BRI); iso FI, 7.v.(fragment
& photo BRI), W, n.v.(photo BRJ)).

Andropogon superciliatus Hackel, Bot. Jahrb. Syst. 6: 239 Sauce Sorghum super-
ciliatum (Hackel) Kuntze, Revis. gen. pl. 792 (1891); Dichanthium superciliatum
(Hackel) A. Camus, Bull. Mus. Hist. Nat. (Paris) 27: 550 (1921). Type: Near
Atapupu, Timor, Naumann s.n. (holo: B, fide Blake (1969) 1.Y.).

Specimens examined: Papua New Guinea. Port Moressy District: Port Moresby, 1893, FE. Cowley 16 (BRI).
Australia. Western Australia. FITzGERALD DisTRict: Kimberley 88 Outcamp, road to Mindewinde, Apr 1988,
Simon 4048 & Sands (BRI,CANB,DNA,K,MEL,NSW,PERTH). GARDNER District: 100 km SE of Kununurra,
Mar 1979, Petherami 322 (BRI,K,PERTH). Hatt District: Fox River Station, Mar 1981, Glover FR | (BRI.
Northern Territory. VICTORIA RIVER: Kidman Springs, Apr 1980, Andrew 616 (BRLCANB,DNA), DARWIN AND
GULF: Austral Downs, May 1947, Blake 17972 (AD, BRI,CANB,DNA,K,MEL,NSW,PERTH). BARKLY TABLELAND:
Newcastle Waters, May 1975, Latz 5989 (BRI,DNA). CENTRAL NORTH: 16 Mile Creek, N of Alice Springs, Feb
1971, Nelson 2085 (BRI,DNA). Queensland. Cook District: 11 km E of Meripah, May 1987, Clarkson 7158 &
Simon (BRI,LMBA,NSW). BurKE DISTRICT: Hughenden, Jun 1934, Blake 6248 (BRILCANB,NSW,PERTH).
GREGORY NorTH District: Tranby, May 1936, Blake 11429 (BRI). NORTH KENNEDY District: Antil Creek,
Mar 1973, Henderson 1422 (BRI). SouTH KENNEDY DISTRICT: Mackay, Nov 1963, Glover s.n. (BRI). MITCHELL
District: Cameron Downs, Mar 1943, Pearson 163 (BRI).

EULALIA Kunth

EKulalia annua B. Simon, species nova affinis FE. aureae (Bory) Kunth sed habitu annuo,
culmis caespitosis infirme interdum decumbentibus differt. Typus: 7 km N of
North Kennedy R. on Peninsular Development Road, Cape York Peninsula,
Queensland, 28 April 1983, J.R. Clarkson 4802 (holo: BRI!; iso: CANBI,
K!,MBA!,NSW!,QRS)). Fig. 2.

Habit annual. Culms 10-45 cm tall, decumbent to weakly tufted, sometimes decumbent,
glabrous at base, simple; nodes glabrous. Ligule a hair-fringed membrane, ca 0.1 mm
long. Blades 3~7 cm long, 1-3 mm wide. Inflorescence subdigitate, or digitate. Peduncles
hairy. Racemes 2 or 3, with dark brown hairs, 7—-20-jointed, 1.5-4 cm. Joints and pedicels
the same length, densely fulvous, 1.8 mm long. Spikelet dorsally compressed, lanceolate-
linear; 2.5-3 mm long, ca 0.6 mm wide: callus ca 0.3 mm long. Lower glume very
narrowly truncate, lanceolate-linear, rounded on back, membranous to thinly chartaceous,
with long fulvous hairs on lower half on back (larger towards tip), reddish brown, 5-
nerved (very faint). Upper glume 3-nerved (faint), as long as lower glume, linear-
lanceolate, thinly fulvous on back. Lower lemma absent. Upper lemma stipe-like, ca 0.5
mm long. Awn 18-20 mm long.

Specimens examined: Northern Territory, DARWIN AND GULE: Munmarlary, May 1978, Dunlop 4805
(BRI,CANB,DNA,K,NT); Munmarlary, Apr 1973, Latz 3870 (BRI,;CANB,DNA,L); Kakadu National Park,
Lazarides 8842 (CANB,MEL,NSW):; Hempe Bay, Groote Eylandt, May 1948, Specht 384 (BRI.MEL,NSW,PERTH);
Melville Island, Apr 1986, Johnson 4218 (BRI); Darwin, Allen 183 and 466 (BRI): Darwin, 1881, Burkitt s.n.
(BRI,MEL). Queensland. Cook District: Merepah, May 1987, Clarkson 7154 & Simon (BRI,DNA,K,L,
MBA,NSW,PERTH,QRS); Lakefield National Park, May 1987, Clarkson 6964 & Simon (BRI.K,L,MBA,NSW,QRS):
30 km N of Mareeba, May 1975, Simon 2652A, Clarkson & Staples (BRI,CANB,K); Bamaga, May 1981, Alcorn
8172 (BRI); ditto, 8172 (BRI); York Downs, May 1981, Morton 1209 (BRI,MEL). BURKE DISTRICT: Normanton,
Gulliver [BRI280496] (BRI,MEL).

Simon, Australian Andropogoneae 5

PP PPE

Onecabeate een. — ayes! cadena ot oe -

BRISSB

2

Fig. 2. Holotype of Eulalia annua.

86 Austrobaileya 3(1): 1989

This annual grass has been in herbarium collections for some time, having been
wrongly identified as Eulalia leschenaultiana (Decne) Ohwi, applicable to a decumbent
Asian perennial species and Pseudopogonatherum collinum (Balansa) A. Camus, a
synonym of P. contortum (Brongn.) A. Camus. The genera Eulalia and Pseudopogon-
atherum have recently been combined (Clayton & Renvoize 1986) on the basis that the
character of rachis toughness, implied by them to have been used as the diagnostic
character to keep the genera apart, breaks down. However at least three other characters
known to separate Pseudopogonatherum from Eulalia are the distinctly awned upper
glume (awnless in Eulalia), both spikelets of each pair being pedicelled (one sessile in
Eulalia) and the convex lower glume (concave in Eulalia). The new species is typically
Eulalia in all these characters.

HYPARRHENIA Andersson ex Fourn.

Hyparrhenia rufa subsp. altissima (Stapf) B. Simon, comb. nov.

Andropogon altissimus Hochst. ex A. Braun, Flora 24: 277 (1841), non Raspail
(1825), nec Colla (1836); Hyparrhenia altissima Stapf in Prain, Fl. trop. Afr. 9:
307 (1918). Type: Ethiopia, plants cultivated at Karlsruhe from seed collected by
Schimper (holo: n.v.; iso: K!(photo BRI)).

Andropogon fulvicomus var. approximatus Hochst. in Sched., Schimp. iter abyssicum
exs. 2: 928 (1842). Type: holo: 7.v.; iso: K(photo BRI).

Specimens examined: Queensland. Cook District: Malanda, Jun 1963, Johnston [AQ262567] (BRI). NortTH
KENNEDY District: Near Ingham, Jun 1952, Everist 5172 (BRI). New South Wales. NortH Coast: Maitland,
Vane {NSW67/401] (NSW); Newcastle, MfcLennan {NSW183563] (NSW); Raymond Terrace, Mar 1981, Lloyd 368
(BRI); Belmont, Jun 1986, Lioyd 412 (BRI); ditto, May 1986, Lioyd 413, (BRI,CANB); ditto, Jun 1986, Lloyd
415 (BRI, PERTH); Doyalson, Jun 1986, Lloyd 411 (AD,BRI); Jun 1986, ditto 414 (BRI,NSW); Sandgate, NW
of Newcastle, Story 7588 (CANB, 3 sheets).

According to the most recent revision of Hyparrhenia (Clayton 1969) there are
four species in Australia — H. rufa, H. hirta, H. quarrei and H. filipendula. The first three
are thought to be naturalized exotics, whereas H. filipendula has a distribution pattern
exhibited by some other Old World andropogonoid species, namely Themeda triandra
and Heteropogon contortus. Clayton (1969) regards H. rufa as a very variable species
and he lists ten characters with “no obvious discontinuities.”” Within Australia however,
the forms with villous peduncles appear to have a more pallid colour of the spikelet
indumentum, as well as a different distribution from the more common form with
glabrous peduncles and reddish brown spikelet indumentum. For these reasons taxonomic
recognition at subspecific rank seems appropriate to me. The subspecies is allopatric
from the typical subspecies (within Australia) and is known to occur in the Newcastle
area of New South Waies and in northern coastal Queensland.

Hyparrhenia quarrei Robyns is separated from H. hirta by Clayton on the basis
of the racemes being deflexed and he cites two specimens from Australia, one from
Western Australia and the other from New South Wales. Examination of these specimens
has revealed them to fall within the circumscription of H. hirta and that raceme deflection
is probably a function of age rather than by anything fixed genetically. The type of H.
quarrei does not show raceme deflection very well anyway, being a young specimen, and
supports my view of placing H. quarrei in synonymy.

Hyparrhenia hirta (L.) Stapf in Prain, Fl. trop. Afr. 9: 315 (1918). Andropogon hirtus
L., Sp. PL: 1046 (1753). Type: Italy, Burser 1.119 (holo: UPS, fide Clayton 1969,
n.V.).
Hyparrhenia quarrei Robyns, Fl. Agrost. Belge 1:171 (1921), synon. nov. Type: Zaire,
Kafubu, P. Quarre 199 (holo: BR!(photo BRLK)).
Robyns, Bull. Jard. Bot. Brux. 8: 234 (1930).

ISCHAEMUM L.

Ischaemum albovillosum B. Simon species nova affinis J. fragili R. Br. sed gluma superna
spiculae sessilis acuminata et hirsuta, gluma inferna spiculae sessilis hirsuta, palea
inferna absenti differt, et affinis J. roseotomentoso Phipps sed pilis albis, gluma

8/

AUSTRALIA PROJECT

EA

=

Simon, Australian Andropogoneae
Fig. 3. Holotype of Jschaemum albovillosum

88 Austrobaileya 3(1): 1989

inferna spiculae sessilis sine dente centrali differt. Typus: Hamersley Railway line,
between Dampier and Mt Tom Price, Western Australia, October 1968, Campion
sn. [PERTH 00349925] (holo: PERTH!(photo BRI)). Fig. 3.

Habit perennial. Culms to 40 cm tall, densely to loosely tufted, simple: nodes usually
bearded. Ligule ca | mm long (imperceptibly hair-fringed). Blades 5-12 cm long; 2-5
mm wide. Inflorescence single. Racemes !10-12-jointed, 6-8 cm long. Joints shorter than
pedicels, with ciliate margins, ca 5 mm long. Sessile spikelet dorsally compressed, elliptic,
ca 10 mm long, ca 2 mm wide. Lower glume not rugose, coriaceous, glabrous outside,
hairy towards apex inside, erose, 9-nerved, 2-keeled, without wings at apex. Upper glume
d-nerved, ca 9 mm long, acuminate, hairy inside and on keel outside, coriaceous. Lower
lemma hyaline. Lower palea absent. Upper lemma stipe-like, ca 5 mm long, membranous.
Awn to 30 mm long. Upper palea absent. Anthers ca 1 mm long. Grain ca 2.5 mm
long. Pedicelled spikelet ca 10 mm long, elliptic, developed. Lower glume coriaceous,
hirsute with long tubercle-based hairs especially on keels and subapical arch, 15-17-
nerved. Upper glume 2/3 as long as lower glume, glabrous, acute, 11-nerved. Lower
lemma ca 6 mm long. Upper lemma ca 5 mm long.

Thus far the new species is only represented by the holotype.

This very distinctive species resembles J. roseotomentosum from Zimbabwe in
that it has a single raceme, the lower floret of the sessile spikelet is sterile and the
corresponding spikelets are about the same size. J. albovillosum differs from I. roseoto-
mentosum in that the tomentum of the young inflorescence is white and not mauvish
pink and the lower glume of the sessile spikelet does not have broad wings with a central
tooth at the apex. Both species differ from J. fragile in the larger spikelets and the silky
tomentum.

Ischaemum tropicum B. Simon, species nova affinis J. po/ystachyo Presl sed inflorescentiis
racemis duobus vel tribus, culmis erectis differt, et affinis J. afro (J. Gmelin)
Dandy sed inflorescentiis racemis duobus vel tribus, spiculis parvioribus
differt, et affinis J. timorensi Kunth sed _ spiculis grandioribus differt.
Typus: NW of Katherine, Northern Territory, 26 April 1947, S.T.Blake 17475
(holo: BRI; iso: CANB!,DNA!,K).LI,MEL! |NSW!). Fig. 4.

Habit perennial. Culms 80-120 cm tall, densely to loosely tufted, shortly rhizomatous,

simple, nodes hairy. Ligule ca 2 mm long (rim of hairs). Leaves glabrous; blades 8-20 _
cm long, 2-4 mm wide. Inflorescence digitate. Racemes 2 or 3; pedunculate (upper), 8-

[4-jointed, 4-6 cm long. Peduncles 6-7 mm long; slightly hairy on margins. Joints longer
than pedicels, with ciliate margins, ca 4.5 mm long, triangular in section, base of joint
and pedicel producing a pore. Sessile spikelet triangular in section, elliptic, 5~5.5 mm
long, 1.5-2 mm wide. Lower glume elliptic-acute, not rugose, glabrous, chartaceous, bifid
(obscurely), 10-nerved, 2-keeled, with wings at apex; wings narrow. Upper glume acu-
minate (obscurely with awn to 2 mm long), glabrous, coriaceous, 4-nerved. Lower lemma
acute, glabrous, ca 6 mm long. Lower palea slightly shorter, hyaline. Upper lemma ca
> mm long, hyaline, bifid with linear lobes, bifid for 1/3 of length. Awn 12-16 mm long.
Upper palea lanceolate, ca 5 mm long. Anthers ca 2.5 mm long. Pedicelled spikelet
neuter, 2~7 mm long, elliptic, developed or reduced to 1 or 2 glumes. Lower glume
glabrous, not rugose below, coriaceous, 11-nerved. Upper glume subequal, acuminate,
glabrous, 7-nerved. Lower lemma acute, ca 5.5 mm long. Lower palea equal. Upper
lemma ca 5 mm long; awned. Palea ca 5.5 mm long. Anthers ca 1 mm long.

Specimens examined: Papua New Guinea. Morose District: Near Lae, Apr 1975, Henty NGF 49749
(A,BRI,CANB,L). Australia. Northern Territory, DARWIN AND GULF: George Creek near Robins Falls, S of
Adelaide River, Lazarides 252 & Adams (CANB,DNA,K,L); Robins Falls, Mar 1972, Passlow 270 (BRI).
Queensland. COOK District: Moa Peak, Moa Island, Aug 1985, Budworth 305 (BRI).

This is a fairly distinctive grass with affinities with the Malesian species J.
polystachyum Pres| and I. timorense Kunth and the African species J. afrum (J. Gmelin)
Dandy in that there is a distinct pore at the junction of the rachis joint and the pedicel.
This feature is not present in any other Australian species but is fairly common in
species of other regions.

89

ogoneae

*

ian Androp

Simon, Austral

.

Cu

otype of Ischaemum tropli

|

Ho

*

90 Austrobatleya 3(1): 1989

MNESITHEA Kunth

Mnesithea pilosa B. Simon, species nova affinis 4. formosae (R. Br.) Koning & Sosef
sed sine fimbria ciliata ad apicem articuli rhachidis, spiculis sessilis longioribus,
spiculis pedicellis rudimentalissimis differt. Typus: Bottom C Paddock, Swans
oo Queensland, 24 March 1983, P. Chaplain [AQ349053] (holo: BRI). Figs
5 &

Habit annual. Culms 25-30 cm tall, erect, branched; nodes hairy. Ligule ca 1 mm long.
Blades 3-6 cm long, 1-3 mm wide. Inflorescence single (rarely with 2 spikelet pairs at
a node). Racemes 1, pedunculate, 5-10-jointed, 1-4 cm. Peduncles glabrous, 2-3 mm
long. Joints and pedicels the same length (fused to halfway), glabrous, 3-5 mm long:
pedicel rigid and 3-nerved. Sessile spikelet dorsally compressed or terete, ovate, 2-3.5
mm long, 1.3-2 mm wide. Lower glume ovate, rounded on back, indurated, pilose,
rugose (slightly), 11-nerved (indistinctly), 2-keeled, without wings at apex. Upper glume
+ equal to lower glume, ovate, glabrous, membranous, not mucronate, 5-nerved. Lower
lemma elliptic-acute, glabrous. Lower palea absent. Upper lemma narrowly elliptic, as
long as upper glume, membranous. Upper palea elliptic, slightly shorter than its lemma.
Anthers | mm long. Pedicelled spikelet neuter, much shorter than sessile spikelet (reduced
to a glume fragment 0.5 mm long).

This grass, so far known only by the type, was originally thought to belong to the
genus Thaumastochloa. However the presence of a rudimentary pedicelled spikelet
necessitated it be moved to the genus Mnesithea, recently enlarged by the amalgamation
of a number of genera of the informal groups Rottboelliastrae of Stapf and Coelorachi-
dastrae of Clayton (Veldkamp, de Koning & Sosef 1986).

SCHIZACHYRIUM Nees

Schizachyrium mitchelliana B. Simon, species nova affinis S. pachyarthro C. Gardner
sed ligulis longioribus, inflorescentia ramis lateralibus multis differt. Typus: Mitch-
ell River Station, Admiralty Gulf, Western Australia, Nov/Dec 1973, 7. Kubicki
53 (holo: PERTH!). Fig. 7.

Ligule ca 3 mm long. Spatheoles involute towards tip, glabrous, 1.8-3 cm long, 3-4 mm
wide. Peduncles 3-5 mm long. Racemes 1-—1.5 cm long. Joints and pedicels + equal to
sessile spikelet; joints diffusely hairy, with hairs longer towards apex, narrowly cuneate
with parallel sides, ca 0.1 mm wide at base, ca 0.5 mm wide at tip. Sessile spikelet
dorsally compressed, narrowly ovate to narrowly ovate-oblong, ca 4 mm long, ca | mm
wide, 4 times as long as wide. Callus ca 0.1 mm long. Lower glume 2-keeled throughout,
narrowly winged, flattened, glabrous. Upper glume 1-nerved. Lower lemma as long as
to shorter than glumes. Upper lemma awn ca 2 mm long. Anthers ca 1.5 mm long.
Pedicelled spikelet ca 2 mm long (reduced to glumes). Awn {0-12 mm long.

This species is thus far known only by the type, which itself is rather fragmentary
in that most of the spikelets have broken up and only five inflorescences are present.
However, the general form of the inflorescence is so very different from other Australian
species of Schizachyrium and the locality of the Mitchell Plateau has and is revealing
many new taxa of plants. Further collecting 1n the area is required to collect better
material of this species.

THAUMASTOCHLOA CLE. Hubb.

Thaumastochloa heteromorpha B. Simon, species nova affinis 7. pubescenti (Benth.) C.E.
Hubb. sed spiculis heteromorphis (laevibus, areolatis vel semiareolatis) differt.
Typus: Cabbage Tree Creek on road from Laura to Cooktown, Lakefield National
Park, Cape York Peninsula, Queensland, 5 May 1987, Clarkson 6940 & B.K.
Simon (holo: BRI'!; 1s0o: K!,L!;MBA!,NSW!,PERTHI!,QRS!). Figs 8 & 9.

Culms 20-55 cm tall, nodes 6-9. Sheaths 0.8-1.5 cm long, pilose. Blades 1.5-8 cm
long, 1-2 mm wide, linear, flat to involute, pilose. Peduncle glabrous, 10-20 mm long
or 60-85 mm long, erect to recurved, to erect, straight. Spikes heteromorphous, with
spikelets alternating in 2 rows or + in | row, cylindrical to constricted, 1-2 cm long,
i-1.5 mm wide, or ca 0.5 mm wide, 5- or 6-jointed, or 2- or 3-jointed. Joints 3-4 mm
long; articulation straight or oblique. Spikelets 2.5-3 mm long. Lower glume triangular-

91

Simon, Australian Andropogoneae

|

Fig. 5. Holotype of Mnesithea pilosa.

92 Austrobaileya 3(1): 1989

oblong, convex (slightly), rugulose or smooth, 5-7-nerved (obscurely). Upper lemma

nerved.

Specimens examined: Q d. Cook District: 19 ml E of Mt Garnet, May 1967, D.E. Symon 4883
(AD,BRI,CANB): N et “hillazoe, gies 1938, Blake 13604 (BRI, CANB): track to Koolburra, Apr 1980, Clarkson
3146 (BRI,CANB,MBA): 40 ml NW of Mungana, Apr 1938, Blake 13738 (BRI); Fairview to Kimba road, Apr
1980, Clarkson 3176 (BRI,CANB,K,MBA,NSW,QRS): Jardine River, Brass 18880, in part (CANB).

=

mastochloa major S.T. Blake and T. ‘pubescens (Be th.) c rAUDE

taxon en one of these species) or a distinct species (de | S
me specimens have a greater representation of areolate than

1983). SO] 0 : if
nore others do not, suggesting some form of introgression is of erating. Som h
ea wi pagans spikelets pe ony Siig) tly narrower than spikes witl | areolate spikelets,
p wi! ck some spikelets are
‘ z have mt weakly areolate Sens
WESP he i widieation that this iakan shows evidence eet 0 : t
rehire and 7. major it is |
name until experim |

ma and th arte flatly careful
it 1s faily ea sy 10 1 mak mixed gathering. A slapper camer literature (Hi bbard
Thursday Island in | June 189
Aneieiseia and a numb

mub bard re ef ers to tb le an V er - elu me OD de1I ng

rig. 6. Two inflorescences of Mnesithea pilosa (holotype).

Simon, Australian Andropogoneae

Fig. 7. Holotype of Schizachyrium mitchelliana.

1989

: aera
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Nene

ol

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=

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Austrobaileya 3(1)

;
i
Ap S904
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;
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4 ui
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=

peters tenet naa

Pee ee

saeaparigiinon:

. Holotype of Thaumastochloa heteromorpha.

Simon, Australian Andropogoneae 95

Fig. 9. Inflorescence of spikelets of 7. heteromorpha: A. with smooth lower glumes (holotype); B. with areolate
lower glumes (holotype).

ria rigida B. Simon, species nova afhnis V. y filipiti (Benth.) C.E. ] lub. | sed r racemis
brevioribus eorum articulis crassioribus multo brevioribus differt. Typus:
River crossing on M Arukun road, Cape York Pennisi, Q

May 1982, J.R ~ Clarkson 4419 (holo: BRI!; iso: CANB!

Culms Senncly tufted, 130-150 cm tall, 3- or 4- noded, branched. Blades 30-50 cm long,
3-5 mm wide. Inflorescence 20-27 cm long, 8-10 cm wide, with branches in whorls of
3-5, 0.5-1 cm long. Racemes 3- or 4-jointed, 3-4.5 cm long. Joints 4-11 mm long, twice
as long as pedicels, both scaberulous upwards and filiform. Sessile spikelet lanceolate-
linear, acute, 6-9 mm long. Lower glume srtilagiious, scaberulous, muricate on nerves,
3- nerved. Upper glume 3-nerved, as long as lower glume, boat-shaped, muricate on keel
otherwise smooth. Awn to 2.5 mm long. Lower lemma ca 6 mm long, oblanceolate,
hyaline. Upper lemma stipe-like, ca 2 mm long, bifid. Awn 8- 10 mm long. Palea
lanceolate, ca 2.5 mm long. Grain ca 3.5 mm long. Pedicelled spikelet neuter, 4-) mm
long, linear-lanceolate. Lower glume 3-nerved. Upper glume a little shorter. Lower lemma
ca 3 mm long. Upper lemma ca 2 mm long.

Specimen examined: Queensland. Cook District: York Downs on Myall Creek, May 1981, Morton 1198
(BRL. MEL) (specimen originally selected as type and annotated as such but superseded by the one selected here).

Blake 1944), V. intermedia was separated from V. filipes
imilar to the one above. An examination of the type
imen could not eaalty be separated from V.
placed in synonymy with it. In more

and deserves recognition.

1989

Austrobaileya 3(1)

ida.

lg

a)

*

lverla r

Holotype of Vet

10

96
Fig.

Simon, Australian Andropogoneae 97

Nomenclatural Changes

As a result of some of my recent revisional work in some genera and study of
type material of Andropogoneae from European herbaria during a recent visit to Kew,
a few Australian species of this tribe require different names from those by which they
were formerly known. It seems a useful exercise to list them here as it may be some
time before the formal Flora of Australia account is published.

1. Dimeria chloridiformis (Gaudich.) Schumann & Lauterb., Fl. Schutzgeb. Sudsee 165
(1901).

Andropogon chloridiformis Gaudich. in Freyc., Voy. Uranie 412 (1826). Type: Guam,
M.C. Gaudichaud s.n. (holo: P!(photo BRI,K)).

Dimeria ciliata Merr., Philipp. J. Sci. 9: 262 (1914). Type: Palawan, Phillippines,
E.D. Merrill 9320 (holo: n.¥.; iso: K!(photo BRI)).

Dimeria chloridiformis has previously been kept separate from D. ciliata on the
basis of the pubescent foliage and rachis, and on having larger spikelets (Reeder 1948).
I do not consider the former character adequate for specific dilimitation whereas the
reported spikelet size difference was not apparent from examination of the types.

2. Eulalia aurea (Bory) Kunth, Revis. gramin. 1: 359 (1830).

Andropogon aureum Bory, Voy. iles Afrique 1: 367, t. 21 (1804); Pogonatherum
aureum (Bory) Roberty, Boissiera 9: 391 (1960). Type: Reunion, /.B.G.M. Bory
de Saint-Vincent s.n. (holo: P!(photo BRI,K)).

Saccharum fulvum R. Br., Prodr. 203 (1810); Erianthus fulvus (R. Br.) Kunth, Revis.
eramin. 1: 160 (1829); Pollinia fulva (R. Br.) Benth., Fl. austral. 7: 526 (1878);
Eulalia fulva (R. Br.) Kuntze, Revis. gen. pl. 2: 775 (1891). Type: Island c [Allen
Island, Carpentaria, Queensland], R. Brown [Iter Australiense 6186] (lecto (here
chosen): BM!(photo BRI,K); isolecto: K!(Qphoto BRI)).

Eulalia geniculata Stapf in Prain, Fl. trop. Afr. 9: 101 (1917). Type: Zaire, Homble
42 (syn: K!); Zimbabwe, Craster 18 (syn: K!); Rogers 4088 (syn: K!); Eyles 1137
(syn: K!).

The African species Eulalia geniculata and the Mascarene species EL. aurea were
formerly regarded as separate from each other and from the Australian FE. ful/va. They
were first placed together by Clayton and Renvoize (1982), who stated “there is consid-
erable variation in such characters as leaf-width, habit, raceme hairiness, and raceme
number, but the spikelet structure, particularly the shape of the lower glume, is constant.
A number of loosely defined and intergrading variants may be recognized, but it 1s an
exaggeration of their differences to treat them as separate species.” Following an exam-
ination of the types I concur with this view and the whole complex is best regarded as
one variable Old World species of the same nature as Hyparrhenia filipendula and
Themeda triandra.

3. Mnesithea Kunth

As a result of recent work on the generic limits of Rottboellia and its allies
(Veldkamp, de Koning & Sosef 1986), I have decided to follow this treatment and place
a number of Australian taxa in the genus Mnesithea. Although their solution is far from
ideal (as they say “clearly the last word has not been said”’), 1t represents an advance
on the understanding of the relationships of these taxa and is a compromise solution
for the time being. Only when further information from sources other than morphology
is available will stability be attained.

The nomenclatural details are as follows:

i) Mnesithea rottboellioides (R. Br.) Koning & Sosef, Blumea 31: 291 (1986).

Ischaemum rottboellioides R. Br., Prodr. 205 (1810); Andropogon rottboellioides (R.
Br.) Steudel, Syn. pl. glumac. 1: 382 (1855); Rottboellia rottboellioides (R. Br.)
Druce, Bot. Soc. Exch. Club Brit. Isles Rep. 1916: 644 (1917); Coelorachis
rottboellioides (R. Br.) A. Camus, Ann. Soc. Linn. Lyon 68: 197 (1921). Type:

98 Austrobaileya 3(1): 1989

Gulf of Carpentaria, North Coast, R. Brown [Iter Australiense 6156] (holo:
BM'(photo BRI,K); iso: K!(fragment BRI),MEL!(photo BRI),NSW!(photo BRI)).

Rottboellia ophiuroides var. commutata Hackel in DC., Monogr. phan. 6: 304 (1889).
Type: Etheridge R., Queensland, F. Mueller s.n. (holo: W!(fragment & photo BRI,
photo K)).

11) Mnesithea granularis (L.) Koning & Sosef, Blumea 31: 295 (1986).

Cenchrus granularis L., Mant. pl. altera 2: 575 (1771); Manisuris granularis (L.)
L.f., Nov. gram. gen.: 37 (1779); Hackelochloa granularis (L.) Kuntze, Revis. gen.
pl. 2: 776 (1891); Rottboellia granularis (L.) Roberty, Boissiera 9: 79 (1960). Type:
“India orientalis” (holo: LINN\(LINN 1217.12)).

iii) Mnesithea annua (Lazarides) Koning & Sosef, Blumea 31: 295 (1986).

Heteropholis annua Lazarides, Nuytsia 5: 288 (1985). Type: Guaging Station, Camp
Creek, Mitchell Plateau, Western Australia, K. Kenneally 8219 (holo: CANBI: iso:
BRI'!,DNA,K!(photo BRI),L,PERTH,US), fide Lazarides.

iv) Mnesithea formosa (R. Br.) Koning & Sosef, Blumea 31: 288 (1986).

Rottboellia formosa R. Br., Prodr. 206 (1810). Type: Island o [Mallison Island], R.
Brown {Iter Australiense 6157] (holo: BM!(photo BRI,K); iso: K!(photo
BRI),MEL!(photo BRI),NSW!(photo BRI)).

Rottboellia formosa var. pilosissima Domin, Biblioth. Bot. 85: 261 (1915). Type:
Cloncurry, Queensland, February 1910, K. Domin s.n. (holo: K!(photo BRI,
fragment NSW))).

Rottboellia formosa forma glabra Domin, Biblioth. Bot. 85: 261 (1915). Type: Castle
Hill, Townsville, Queensland, February 1910, K. Domin s.n. (?holo: K!(photo
BRI, fragment NSW!).

4. Polytrias diversiflora (Steudel) Nash, Torreya 5: 110 (1905).

Andropogon diversiflorus Steudel, Syn. pl. glumac. 1: 370 (Jul 1854). Type: Java,
Zollinger 539 (holo: n.v.; iso: P'{ photo BRI).

Andropogon amaurus Buse in Miq., Pl. jungh. 360 (Aug, 1854); Polytrias amaura
(Buse) Kuntze, Revis. gen. pl. 788 (1891). Type: Java, Miquel (holo: n.y.).

Lazarides (1980) takes up the name Polytrias amaura for this species, placing P.
diversiflora in synonymy, but the date of publication of Andropogon diversiflorus predates
that of A. amaurus by one month so the former’s specific epithet must be taken up
when the transfer to the genus Po/ytrias is made.

Acknowledgements

I am very grateful to the Australian Biological Resources Study scheme of the
Department of the Arts, Sport, The Environment, Tourism and Territories, for a grant
for 1986-1988 to undertake research on and to prepare an account of the Andropogoneae
for the Flora of Australia. With this financial support I was able to employ the diligent
technical assistance of Simone Stewart and Danny Healy, to both of whom I extend my
thanks. It also provided some funding to enable me to visit Kew in August 1986 during
which time many types of Australian Andropogoneae in Europe were examined. | also
thank Hans Dillewaard for the photographic work and for assistance in transferring text
between computers and Gordon Guymer for consultation of type material during his
term as Australian Botanical Liaison Officer at Kew (1986-1987). Thanks are extended
to herbarium curators who loaned specimens of Andropogoneae from their institutions.
Where specimens have been collected personally, support for field work has been given
by the Queensland Herbarium and, for recent collections in the Kimberley region of
Western Australia on the Kimberley 88 project mounted by the Linnean Society of
London and the Royal Geographical Society, by the sponsors of that project.

Simon, Australian Andropogoneae 99

References

BLAKE, S.T. (1944). Monographic studies in the Australian Andropogoneae, Part 1. Papers, Department of
Biology, University of Queensland 2(3). \-62.

BLAKE, S.T. (1969). Taxonomic and nomenclatural studies in the Gramineae No. 1. Proceedings of the Royal
Society of Queensland 80: 55-88.

BLAKE, S.T. (1974). Revision of the genera Cymbopogon and Schizachyrium (Gramineae) in Australia. Contri-
butions from the Queensland Herbarium, No. 17

CLAYTON, W.D. (1969). A revision of the genus Hyparrhenia. Kew Bulletin Additional Series I.

CLAYTON, W.D. & RENVOIZE, S.A. (1982). Gramineae 3. In Polhill, R.M., Flora of Tropical East Africa.
Rotterdam: A.A. Balkema.

CLAYTON, W.D. & RENVOIZE, S.A. (1986). Genera graminum. London: Her Majesty’s Stationary Office.

DALLWITZ, M.J. & PAINE, T.A. (1986). User’s guide to the DELTA system. A general system for processing
taxonomic descriptions, 3rd ed. CSIRO, Division of Entomology Report No. 13.

DE WET, J.M.J. (1987). Hybidization and Polyploidy in the Poaceae, pp 188-194. In Soderstrom, T.R., Halu,
K.W., Campbell, C.S., & Barkworth, M.E. (eds), Grass Systematics and Evolution. Washington D.C. &
London: Smithsonian Institution Press.

DE Bean J.M.J. & HARLAN, J.R. (1962). Species relationships in Dichanthium. IIL. D. sericeum and its allies.
Phyton 18: 11-14.

DE WET, J.M.J. & HARLAN, J.R. (1966). Morphology of the compilo-species Bothriochloa intermedia. American
Journal of Botany 53: 94-98.

DE WET, J.M.J. & HARLAN, J.R. (1968). Taxonomy of Dichanthium section Dichanthium (Gramineae). Boletin
de la Sociedad Argentina de Botanica \2: 206-227.

DE WET. I.M.J. & RICHARDSON, W.L. (1963). Morphological variation and species relationships in Dichan-
thiten. Phyton 20: 19-28.

DE WET, J.M.J. & SINGH, A.P. (1964). Species relationships in Dichanthium. V. The diploid species. Caryologia
17: 153-160.

FARUQI, S.A. (1969). Range of morphological variation within the Bothriochloa intermedia complex. Phyton
(Austria) 13: 285-303.

HUBBARD, C.E. (1936). Thaumastochloa C.E. Hubbard. In Hooker, W.J., cones Plantarum 34: t. 3313, 3314.

KONING, R. DE, SOSEF, M.S.M. & VELDKAMP, J.F. (1983). A revision of Heteropholis and Thaumastochloa
(Gramineae). Gardens’ Bulletin of Singapore 36: 137-162.

LAZARIDES, M. (1980). The tropical grasses of southeast Asia. Vaduz: J. Cramer.
REEDER, J.R. (1948). The Gramineae ~ Panicoideae of New Guinea. Journal of the Arnold Arboretum 29: 257-
392.

SIMON, B.K. (1982). New species of Gramineae from south-eastern Queensland. Austrobaileva 1: 455-467.
SOENARKO, S. (1977). The genus Cymbopogon Sprengel (Gramineae). Reinwardtia 9: 225-375.

TOTHILL, J.C. (1977). Flowering phenology of some native perennial tropical grasses from north-eastern Australia.
Australian Journal of Ecology 2: 199-205.

TOTHILL, J.C. (1981). Distribution, Adaptation and Evolution in Australian Andropogoneae (Poaceae). XIII
International Botanical Congress, Abstracts, 279. Sydney: Australian Academy of Science.

VELDKAMP., J.F.. KONING, R. DE & SOSEF, M.S.M. (1986). Generic delimitation of Rottboellia and related
genera (Gramineae). Blumtea 31: 281-307.

Accepted for publication 10 November 1988

Austrobaileya 3(1): 101-102 (1989) | 101

REVISION OF THE AUSTRALIAN VITACEAE,
4. CLEMATICISSUS PLANCHON

B.R. Jackes

Botany Department, James Cook University of North Queensland,
Townsville, Qid 4811

Summary

Clematicissus angustissima (F. Muell.) Planchon, endemic in the Geraldton area of Western Australia and the
only species in the genus, is redescribed and figured.

CLEMATICISSUS

Clematicissus Planchon in A. & C. DC, Monogr. phan. 5: 422 (1887). Type: C.
aneustissima (F. Muell.) Planchon

This monotypic genus is endemic in the Irwin District of Western Australia.

Clematicissus angustissima (F. Muell.) Planchon in A. & C. DC., Monogr. phan. 5: 422
(1887); Vitis angustissima F. Muell., Fragm. 1: 141 (1859). Type: Murchison
River, Oldfield (holo: MEL(MEL5348); iso: MEL(MEL5349),K).

F. Muell., Fragm. 1: 244 (1859), 9: 126 (1875); Benth., Fl. austral. lL: 450 (1863).

Deciduous vine or sprawling shrub to 2 m high, branches thin, striate, glabrous. Plants
deciduous, numerous tubers present. Mucilage cells and raphide sacs common throughout.
Tendrils leaf-opposed, usually bifid; branch or aborted branch subtended by a bract.
Leaves palmately compound (3-)5(-7)-foliolate, glabrous. Central leaflet from linear-
lanceolate to broadly lanceolate, (2.1-)4.5(-6.3) cm X (0.15-)0.4(-1.2) cm; apex acute;
base attenuated. Lateral leaflets similar but proportionally smaller. Margins entire to
deeply indented. Petiole slender, (0.8—)1.5(-2.8) cm long; petiolules often absent. Stipules
rounded, membraneous, 1.5-2 mm X 1i-1.5 mm. Inflorescence leaf-opposed, cymose,
multiflowered in congested heads which terminate one or both branches of the tendril;
peduncles and pedicels subtended by prominent membraneous bracts. Flowers bisexual,
pentamerous. Calyx shortly 5-lobed, 0.5-1 mm long, membraneous. Corolla with 5 free
lobes weakly cohering in bud; lobes reflexed after anthesis, soon caducous, greenish white
to mustard yellow, 1.5-2 mm long. Stamens inserted on the receptacle at the base of
the disc, opposite the petals; filaments erect, flattened, tapering towards the apex, 0.75~
1 mm long; anthers ovate, introrse, opening by longitudinal slits, ca 0.75 mm long.
Pollen grains tricolporate. Disc adnate to and entirely surrounding the base of the ovary,
broad; edge often spreading, pressure grooves formed by the filaments. Ovary 2-locular
with 2 anatropous, basally attached ovules per locule; style conical, stigma capitate.
Berry fleshy, globular to depressed-globular, purplish black to black at maturity, 7-10
mm X 4-8 mm, with | or 2(-4) pyriform seeds. Shape of seeds in transverse section
varies according to the number maturing in the fruit. Seeds 5.5-7 mm X 3.5-4 mm;
perichalaza extends to the dorsal surface where it forms a rounded depression. Endotesta
irregularly lignified, endosperm ruminate, in transverse section U-shaped. Cotyledons
similar, broadly lanceolate, persisting for up to twelve months whilst tubers are forming.
Figs 1-5.

Representative specimens: Western Australia. IRWIN District: Red Bluff ca 5 km S of Kalbarri, May 1968,
Wilson 6517 (AD,K,PERTH); Northampton, Sep 1926, Gardner 1964 (PERTH); Greenough River, Yuna East
Reserve, 70 km NE of Geraldton, Jan 1982, Jackes (CBG,L,NSW,PERTH); 37 km N of Geraldton, 28°34’S,
l'4°37E, Hnatiuk 760377(PERTH): 1.1 miles [1.8 km] N of Mingenew, Mar 1968, Carr 352 (K,PERTH).

Specimens from 35 localities examined.
Distribution and habitat: This species is found in the Geraldton area of Western Australia

extending north to the Murchison River and south to the Irwin River. Here it is found
in woodlands on a variety of soil types but chiefly on sand or loam.

102 Austrobaileya 3(1): 1989

Fig. 1. Clematocissus angustissima: A. young leaf and inflorescence X 0.5. B. leaf X 0.5. C. seed, ventral surface
showing position of raphe x 5. D. seed, dorsal surface showing position and shape of perichalaza X 5. E. median
T.S. of mature seed X 6 ( e —lignified endotesta).

Phenology: Flowering mainly late summer and autumn; fruits mature autumn to late
spring.

Notes and observations: Superficially this summer deciduous species resembles the
partially or totally winter deciduous eastern species Cissus opaca F. Muell. The obvious
distinguishing features are the dense congested inflorescence, the different shape of the
endotesta and the pattern of formation of the tubers. A good description and illustration
of the tubers are given by Pate and Dixon (1982). Leaf shape and size are highly variable.

Reference

PATE, J.S. & DIXON, K.W. (1982). Tuberous, Cormous and Bulbous Plants. Nedlands: University of Western
Australia Press.

Accepted for publication 10 April 1989

Austrobatleya 3(1): 103-107 (1989) | 103

BRYOPHYTES IN A SUB-TROPICAL MANGROVE
COMMUNITY

J. Windolf

53 Pandanus Avenue, Coolum Beach, Queensland 4573

Summary

A descriptive and quantitative analysis of the bryophyte species in sub-tropical mangroves in southern Queensiand,
their host/substrata relationship and their occurrence relative to the adjacent non-mangrove environment is presented
here. The study is based on 337 collections.

Introduction

Bryophyte ecology has become an important discipline not only to the specialist
working within his particular field but also within general ecology. The majority of work
has been carried out in temperate or polar regions where a considerable amount of
progress has been made, resulting in at least a reasonable understanding of the position
of bryophyte life in relation to the overall environment. Unfortunately this cannot be
said of the tropics and sub-tropics, particularly as regards marginal or highly specific
habitats such as those of the savannah, lowland rainforests or mangroves (Richards
1984).

Geography, climate and physical aspects

The areas studied comprise the tidal or mangrove-supporting sections of the
Noosa, Maroochy and Mooloola River systems on the southern Queensland coast between
approximately 26°20’/S and 26°45’S, 80-120 km north of Brisbane. They contain typical
sub-tropical mangrove communities (Lear & Turner 1977) extending up to 20 km inland.
Those of the Noosa and Mooloola Rivers are principally composed of a narrow fringing
zone and saline marshes, while the Maroochy River, with its richer nutrients and hence
substrate, supports a more varied mangrove flora including extensive areas of closed
Bruguiera forest. Upstream, the landward margin of the mangroves merges with freshwater
swamps, rainforest or sugar cane plantations while near the river mouths considerable
urban development has taken place in recent years. In spite of this, remarkably little of
the original mangrove environment has been disturbed up to the present time. The
climate of the region is of the sub-tropical, humid, east-coast type, characterized by hot
summers, mild winters and a clearly defined wet-season but with sufficient rainfall in
all seasons to allow continuous growth of vegetation (Windolf 1985b).

Collecting and recording

Collecting was carried out along transects aligned at right angles to the species/
structural zonation. Sufficient transects were employed so as to include all obvious
combinations of zoning in various sections of the three separate river systems. Individual
host trees (or sub-strata) were numbered and all species found thereon were recorded
using this number. In addition such details as density of the surrounding vegetation,
height above high-tide level, proximity to non-mangrove vegetation and pertinant
information on micro-habitat were also recorded.

Host trees and sub-strata

Because of the inherent saline influence in the mangrove environment, soil, except
in the case of upturned tree bases, is virtually eliminated as a sub-stratum and 99% of
specimens were found growing either as epiphytes on living trees or on decaying timber
above the highest tide level. |

Sub-tropical mangrove environments support far fewer tree species than do those
of the true tropics. In the area under consideration six mangrove species have been

[04 Austrobaileya 3(1): 1989

recorded compared with the eastern Australian coast at 12°S with 28 species and 30°S
with 2 species. In addition several other species occur with varying degrees of frequency,
not only in the mangroves, but also in adjacent non-saline environments. Of these only
one, Casuarina glauca, was found to support epiphytic bryophytes within the mangrove
environment proper.

The phorophytes/sub-strata which supported bryophytes and the number of each
from which specimens were collected are listed in Table 1.

Table 1
l. Mangroves
Avicenniaceae
Ayvicennia marina (Forsskal) Vierh. 15
var. australasica (Walp.) Mold.
Rhizophoraceae
Bruguiera gymnorhiza (L.) Savigny 48
Rhizophora stylosa Griffith |
Euphorbiaceae
Excoecaria agallocha L. 36
Zs, Other
Casuarinaceae
Casuarina glauca Sieber ex Sprengel 51
Decaying timber (species not defined) {1
Soil 4
Total 166

No bryophytes were found on two mangrove species, Aegiceras corniculatum (L.)
Blanco and Cerlops tagal (Perrottet) C.B. Robinson var. australis C. White. In the
transitional landward zone there is a rapid changeover between mangrove species and
those more commonly associated with the adjoining non-mangrove forest. Specimens
were collected only from those areas where mangroves formed the major plant type.

Notes on bryophyte species
Hepaticae
(SEOCALYCACEAE

Lophocolea semiteres (Lehm. & Lindenb.) Mitt. Cosmopolitan. Very common locally in
a wide variety of habitats. Those occurring on mangrove plants were invariably small
and weak and in many cases appeared to be struggling to survive.

FRULLANIACEAE

Frullania ericoides (Nees) Mont. Pan-tropical. Locally common in wet sclerophyll forest,
woodland and vine scrub. The specimens observed in the mangrove forest were robust
and prolifically fertile.

Frultania ferdinandi-muelleri Steph. Endemic to the central-east Australian coastal region
(26°S-30°S). Locally common, usually on the fringes of rainforest or in lush wet sclerophyll
forest. This species had a definite preference for Bruguiera gymnorhiza and Casuarina
glauca in closed forest or well shaded sites. Specimens of F. ferdinandi-muelleri were
generally large, robust and often fertile.

Frullania rostrata (J.D. Hook. & Tayl.) J.D. Hook. & Tayl. East Australian coast, New
Zealand and the south-west Pacific region. Locally common in light scrub and woodland.
It had very limited occurrence in the mangroves, being confined to stunted Bruguiera
gymnorrhiza in low open woodland in one particular spot.

Windolf, Bryophytes in Mangroves 105

Table 2. Bryophyte species/host relationships

Mangroves Other
Avicennia Brugtiera Rhizophora Excoecaria Casuarina Decaying Soil Total
marina gyninorrhiza — stylosa agallocha glauca timber specimens

Hepaticae
Lophocoieca senuteres 9 4 l 14
Frullania ericoides 3 1S 16 7 4]
F. ferdinandi-muelleri {3 2 9 24
F. rostrata 6 6
F. subtropica 12 9 [3 15 i 50
Acrolejeunea aulacophora 6 13 i 12 16 3 51
Cheifolejeunea intertexta l 12 l 16 i 3]
Cololejeunea lanciloba 6 18 3 27
Lejeunea cucullata | I
L. flava

ssp. ortentalis 4 l 2 1 8
Leptolejeunea maculata 8 12 4 3 i 28
Schiffneriolejeunea tumida

var. haskarliana | |
Metzgeria decepiens i I
Total Hepatic specimens 27 89 2 81 65 18 1 283
Total Hepatic species 4 12 2 8 7 8 I 13
Musci
Fissidens humutis I 4 3 8
Macromitrium aurescens 3 7 16 5 3
Sematophyliunt sp. j 8 6 45
Total Musci specimens 3 2 7 28 11 3 54
Total Musci species l 2 | 3 2 l 3
Total specimens 30 91 2 8&8 93 29 4 337
Total species 5 i4 2 9 10 id 2 16

Frullania subtropica Steph. Endemic to the central-east Australian coast (20°S-—30°S).
Locally common, often in quite dry habitats. This species preferred open sites in the
mangroves and often covered large areas of the trunks of Casuarina glauca and the near
horizontal limbs of Avicennia marina. The plants were robust and often fertile. In terms
of volume, F. subtropica is probably the dominant bryophyte in the mangroves.

LEJEUNEACEAE

Acrolejeunea aulacophora (Mont.) Steph. Widespread throughout the Pacific and Indian
Ocean regions. Very common locally in a wide variety of habitats and it was well
diversified throughout most parts of the mangrove environment.

Cheilolejeunea intertexta (Lindenb.) Steph. Pan-tropical and sub-tropical. Uncommon in
the surrounding areas. Had a distinct preference for Excoecaria agallocha and Bruguiera
gymnorrhiza, particularly in stunted mangrove scrub.

Cololejeunea lanciloba Steph. Tropical south-west Pacific, south-east Asian regions and
Japan. Relatively common in nearby rainforest and scrub. This species was generally
well developed, prolifically fertile and obviously relishes this particular habitat. kxcoecaria
agallocha in dense closed forest was the preferred host/habitat.

106 Austrobaileya 3(1): 1989

Lejeunea cucullata (Reinw., Blume & Nees) Nees. Widespread in tropical areas of the
Indo-Pacific region. As only a single specimen was collected, it suggests that the mangroves
are not a preferred habitat, but the species 1s also rare in the surrounding vegetation.

Lejeunea fiava (Sw.) Nees subsp. orientalis Schust. Widespread, the species being cos-
mopolitan. Extremely common locally in rainforest, wet sclerophyll forest and damp
woodland. In the mangroves it was confined to the lush closed forest and the plants
collected were not particularly well developed.

Leptolejeunea maculata (Mitt.) Schiffn. Widespread in tropical regions. The species is
reasonably common locally and the specimens from the mangroves were robust and well
established in a variety of micro-habitats.

Schiffneriolejeunea tumida (Nees) Grad. var. haskarliana (Gott.) Grad. & Terken. South-
west Pacific and south-east Asian regions. Uncommon locally and as only a single
specimen was collected from the mangroves, it indicates its rarity in that habitat as well.

METZGERIACEAE

Metzgeria decipiens (Massal.) Schifin. & Gott. Australia, New Zealand and the Pacific
region. Locally common, but mostly in upland rainforest and wet sclerophyll forest. The
single specimen found in the mangroves was small and poorly developed.

Muscli
FISSIDENTACEAE

Fissidens humilis Dix. & Watts. Eastern Australia and New Zealand. Reasonably common
locally on damp banks and tree bases, particularly Melaleuca quinquinervia in freshwater
swamps. In the mangroves it favoured relatively open positions with broken sunlight,
only a few centimetres above the high tide level, occurring on bark and on soil retained
between the roots of upturned trees.

ORTHOTRICHACEAE

Macromitrium aurescens Hampe. Endemic to north-east Australia. Local distribution is
unknown apart from the mangroves but it is probably not common. Its preferred habitat
in the mangroves was on Excoecaria agallocha in open woodland and in open forest on
Casuarina glauca, often in very exposed situations above the main canopy. This tallies
with Vitt and Ramsay’s (1985) comment that this species of Macromitrium appears to
be somewhat xerophytic having an affinity with rough barked host species.

SEMA TOPHYLLACEAE

Sematophyllum sp. Sematophyllum 1s a widespread, very complex genus usually associated
with damp habitats although some species are found 1n seasonally drier areas. The genus
is very common in surrounding districts where a number of species occur. Specimens
have not yet been identified to species level.

Conclusions

Analysis of the data revealed some significant ecological peculiarities as regards
the bryophyte flora on the mangroves of this area. These can be summarised as follows:

A. Hepatics outnumbered the Musci both in terms of species (Hepatics 84%, Musci 16%)
and individual plants.

B. Hepatic specimens were dominated by the genera Frullania (43%), Acrolejeunea (18%),
Cheilolejeunea (11%), Leptolejeunea (10%) and Cololejeunea (10%), others being
of only spasmodic occurrence. Macromitrium (57%) and Sematophyllum (28%)
account for the majority of mosses.

C. Virtually all specimens were epiphytic (99%) because of the peculiar nature of the
mangrove environment.

D. Species prevalence varied somewhat from that of adjacent non-mangrove environments.

E. Some marked bryophyte/phorophyte relationships existed.

Windolf, Bryophytes in Mangroves 107

Although the area studied is essentially sub-tropical in form, it exhibits many
positive similarities to those observed by Thaitong in the tropical mangrove forests of
Thailand (Thaitong 1984). This is particularly so with regard to Hepatic genera. Of the
12 recorded by Thaitong and the 9 described here, 7 are common to both areas, Whether
or not these factors are constant in mangrove environments will only be revealed by
comparing detailed studies carried out in different areas.

Acknowledgements

Acknowledgement is due to Sinski Hattori, Barbara Thiers, Helen Ramsay and
George Scott for their help in identifying specimens and providing general information;
George Scott for reading the text and to Phillip Sharpe for his encouragement and
assistance on numerous collecting trips.

References
LEAR, R. & TURNER, T. (1977). Mangroves of Australia. Brisbane: University of Queensland Press.

RICHARDS, P.W. (1984). The Ecology of Tropical Forest Bryophytes. In Schuster, R.M. (ed.), New Manual of
Bryology. Vol. 2. 1233-1270. Nichinan, Miyazaki: Hattori Botanical Laboratory.

THAITONG, U. (1984). Bryophytes of the Mangrove Forest. Journal of the Hattori Botanical Laboratory 56:
85-87.

VITT, D.H. & RAMSAY, H.P. (1985). The Afacromitrium Complex in Australasia (Orthotrichaceae: Bryopsida).
Part 1. Journal of the Hattori Botanical Laboratory 59: 325-451.

WINDOLF, J. (1985b). Survey of the Hepaticae and Anthocerotae of the Sunshine Coast Region, Queensland.
Journal of the Hattori Botanical Laboratory 58: 171-176.

Accepted for publication 11 August 1988

Austrobaileya 3(1): 109-133 (1989) 109

NOTES ON ASCLEPIADACEAE, 1
P.1. Forster

Botany Department, University of Queensland, St Lucia, Qld 4067

Summary

Cynanchum liebiana (F. Muell.) P. Forster comb. noy. and C. christineae P. Forster sp. noy. are described.
Sarcostemma esculentum (L.f.) Holm is newly recorded for Australia with Marsdenta brockmaniana W. Fitzg.
placed in synonymy. The status of 7ylophora erecta F. Muell. ex Benth., 7. macrophylla Benth. and Tf. stelligera
Schitr. is reviewed with 7. macrophylla and T. stelligera being relegated to synonymy. Tylophora benthamii Tsiang
is shown to be the earliest available name for the taxon known variously as 7. floribunda Benth. or T. crebriflora
S.T. Blake. Isotype material of Sarcolobus ciliolatus Warb. and S. obovatus Rintz at the Queensland Herbarium
is documented and S. obovatus recorded for the Solomon Islands and northern Papua New Guinea. A generic
account for Finlaysonia Wallich in Australia and New Guinea is given, with F. obovata Wallich being recorded
for the region. Gyrmnema sylvestre (Retz.) R. Br. ex Schultes is lectotypified. The species does not occur in Australia.
Gymnema pleiadenium F. Muell. is an earlier name for the taxen known as Gymnema dunnii (Maiden & Betche)
P. Forster. G. pleiadenium is distinguished from G. geminatum R. Br. and G. sylvestre. Brachystelma glabriflorum
(F, Muell.) Schltr. is the earliest name for the species of Brachystelma that occurs in Australia and New Guinea.

Contents
CVHAMCHVNIES wos x als Goce in OM ee Ege be tn Se te cee AE pS te a sah 110
Sarcéstenntia BK Br. .. os <p es Ve ee Ye Se ee BS ee Oe we Seog he RE ae 114
AVIGBWOTHIRG. BE. -. 02 2% eon Bw. whois Fue he woe ALS ode poe aS Ee ge re 115
Sarcolobus GL. se <a. uses. Velie we QE oe oe nee J ed 2 bende ae cle ee Ee 121
Finlaysonia. Wallich. ..... 0... 2 to ke te be a ke HAE Hb 123
Gymnena Ry Bri foo. cee aa ot tee) oo ve web da BY ae ce te 125
Brachystebiia SHAS! =. 2.00 nace Wiese Ge Ado dh poo oll, pHa ee Bake 2 25 130

There are relatively few collections of certain Australian and New Guinean
Asclepiadaceae available for taxonomic study. Furthermore, many specimens in herbaria
are of limited use in the preparation of accurate descriptions, due to the poor quality
of the specimens compounded by the distortion to the intricate floral structures brought
about by pressing and drying. Consequently comprehensive treatments of the larger
genera such as Marsdenia R. Br., Cynanchum L., Gymnema R. Br. and Tylophora R.
Br. cannot be completed until a number of satisfactory specimens in spirit (obtained
from both field collections and cultivated material) can be examined.

However, due to some recent excellent collections of material, several new taxa
have been recognised. Some of these are known only from a few specimens or apparently
have a restricted geographical range and they are described here to draw attention to
their conservation status.

In addition to the descriptions of new species, notes on nomenclature are provided
for a number of taxa, and some new records for Australia are discussed.

Materials and Methods

Holdings at BRI, BRIU, CBG, DNA, JCT, MEL, PERTH, QRS, partial holdings
at CANB and selected types at K and LD were examined. In specimens cited below,
material preserved in spirit is indicated by an *. Material at NSW was unavailable for
loan due to staff shortages (K. Hill, pers. comm. 1988). Collections were made or obtained

110 Austrobaileya 3(1): 1989

in Queensland and the Northern Territory, with material in spirit (70% alcohol, plus
glycerol) used in most instances for preparing descriptions.

The small structures at the base of the lamina in many taxa of the Asclepiadaceae
and related Apocynaceae have been variously referred to as ‘finger glands’ or ‘glands’ in
Australian literature (Williams 1984; Stanley & Ross 1986). These are correctly termed
extrafloral nectaries (Arekal & Ramakrishna 1980; Mohan & Inamdar 1986) and should
not be confused with the floral nectaries present in most species in the family (Christ
& Schnepf 1988).

CYNANCHUM L.

Examination of the type specimen of 7y/ophora liebiana F. Muell. revealed that
the species should be placed in Cynanchum and that it is allied to C. brachystelmoides
P. Forster and C. christineae P. Forster. Von Mueller’s (1891) placement of this taxon
in Tylophora is not altogether surprising, as it possesses a staminal corona which is
superficially similar to that of several taxa of Tylophora in that the lobes are rounded
and do not overtop the staminal column. In this respect, the taxon is similar to
Cynanchum hydrophilum Tsiang & Zhang, C. riparium Tsiang & Zhang and C. gracilipes
Tsiang & Zhang from China (Tsiang & Li 1976). The form of the pollinia (not globose
as in 7 ylophora), corolla and bud shape, and the position of the staminal corona all
clearly indicate that 7. /iebiana is a species of Cynanchum.

Cynanchum liebiana (F. Muell.) P. Forster, comb. nov.

Tylophora liebiana F. Muell., J. & Proc. Roy. Soc. New South Wales 24: 78 (1891).
Type: Port Darwin, 1890, 14. Holtze 1010 (holo: MEL).

Herbaceous perennial. Roots thinly fibrous-fusiform, to 60 mm long, 2-3 mm diameter.
Stems cylindrical, upright, to 60 cm long, 0.4-0.5 cm diameter, up to 30 nodes on
flowering plants, glabrous; internode length variable to 7 cm. Leaves narrow-linear,
glabrous, 40-50 mm long, c. 1 mm wide. Flowers borne on top 2 or 3 nodes. Cymes
umbelliform, appearing at nodes between the pair of leaves, 1-5-flowered; peduncles
glabrous, 5-8 mm long, c. | mm diameter. Flower pedicels 11-12 mm long, c. 0.5 mm
diameter, filiform, pendulous during anthesis, glabrous. Calyx without basal glands; sepals
broadly triangular, acute, glabrous, c. 2.5 mm long, c. | mm wide. Petals 5, distinct,
twisted in bud; linear-lanceolate, 10-11 mm long, c. 1.5 mm wide, abaxial surface
glabrous, purple-brown or cream, adaxial surface glabrous or with white cilia, purple-
brown or cream. Corona of 5 separate lobes adnate to base of staminal column, c. 3.75
mm diameter; lobes broadly ovate to globose, tip rounded, c. 0.5 mm long, c. 1 mm
wide, purple-brown or cream. Staminal column c. 1.5 mm long, c. 1.25 mm diameter,
corona adnate to bottom two-thirds. Anther connectives incurved, apical appendage
ovate, c. 0.5 mm long, c. 0.5 mm wide. Slit between anther wings c. 0.75 mm long.
Pollinaria 0.52 mm long, 0.36 mm wide; pollinia solitary in each anther cell, pendulous,
oblong-ovoid, 0.28 mm long, 0.13 mm wide; caudicles curved, translucent, 0.1 mm long,
0.04 mm wide; corpusculum oblong, brown, 0.2 mm long, 0.13 mm wide. Style-head
conical, obtuse, cream. Follicles fusiform-globose, glabrous, c. 6 cm long, c. | cm diameter:
fruiting pedicel 15-16 mm long, c. | mm diameter. Seed 15 mm long, 10 mm wide,
ovoid, light-brown; coma absent. Fig. 1.

Specimens examined: Northern Territory. DARWIN AND GULF District: 15 km S of Darwin, Swamp W of Rural
Caravan Park, 12°55’S, 130°58’E, Dec 1984, Jones 1738 (DNA); Cnr Bonson Terrace & Elrundie Avenue,
Palmerston, Dec 1987, Tingey [AQ408479] (BRI*); ditto, Feb 1988, Tingey [AQ408482] (BRI); Elrundie Avenue,
Palmerston, 12°30’S, [31°O0’E, Feb 1987, Tingey s.n. (DNA*).

Distribution and habitat: The species has been recorded only from the area surrounding
Darwin, from sandy embankments on disturbed ground adjacent to black soil paperbark
swamp. This is the ARNM Region of Barlow (1985).

Phenology: Flowering occurs from November to December with the flowers opening in
the morning and closing by midday (R. Tingey, pers. comm. 1987). Fruiting occurs from
February to March.

Notes: Table 1 provides a comparison of the major differences between C. liebiana, C.
brachystelmoides and C. christineae.

Forster, Ascleptadaceae, | 111

| Se \\\ =
LD pai

IRSA

Cn

te 1:
i Si fe
~ ETE
ff

Fig. 1. Cynanchum liebiana: A. habit < 0.5. B. cyme with several buds x 3. C. ventral view of flower x 3. D.
lateral view of flower with petals partly unfurled X 3. E. apical view of staminal corona and column X 5, F,
lateral view of staminal corona and column x 5. G. front view of anther x 20. H. lateral view of anther and
associated coronal lobe X 20. L. pollinarium Xx 40. All from spirit material of Tingey s.n. (DNA). Del. A.
Thongpukdee.

112 Austrobaileya 3(1); 1989

Table 1. Comparison of major distinguishing characters of Cynanchum liebiana, C. christineae and
C. brachystelmoides (C. brachystelmoides data from Forster & Thongpukdee 1988).

Character C. liebiana C. brachystelmoides C. christineae

Sepal size c. 2.5 * 1mm 1-11.25 X 0.1-0.2 mm c. 1.5 X 1.0 mm

(length X width)

Petal shape linear-lanceolate ovate-oblong linear-lanceolate

Petal size 10-lII Xe 15mm 1,5-1.7 X¥ ¢. 0.4mm cc. 5.0 X 1.5 mm

(length X width)

Corona lobe shape broadly ovate- broadly-ovate, tip cylindrical-oblong,
globose, tip rounded ovate-obtuse tip rounded

Pollinium size c 0.28 0.13 mm ¢.0.1 X¥ 0.025 mm c,. 0.21 X 0.13 mm

(length X width)

Corpusculum length c. 0.2 mm c. O.1 mm c. 0.2 mm

Caudicle length c. 0.1 mm c. 0.015 mm c. 0.14 mm

The lack of a coma on the seed of C. liebiana and C. christineae is an unusual
feature as a coma is normally present on seeds of Asclepiadaceae. Rintz (1980) records
Sarcolobus carinatus Wall., S. globosus Wall. and S. oblongus Rintz as having seed
without a coma. Unfortunately the follicles and seed of C. brachystelmoides are still
unknown. C. /iebiana is a particularly beautiful plant and efforts should be made to
propagate material for cultivation.

Conservation status: At present this species is only definitely known from the area of
the cited collections. These are from areas of disturbed ground or near to human
habitation and are under immediate threat from housing development. An appropriate
conservation coding is IE (Leigh ef a/. 1981).

Cynanchum christineae P. Forster species nova affinis C. brachystelmoidi P. Forster sed
sepalis ca 2.5 mm longis et 1 mm latis, petalis linearibus-lanceolatis, lobis coronae
cylindricis-obloideis, polliniis ca 0.21 mm longis et 0.13 mm latis; corpusculis ca
0.2 mm longis et caudiculis ca 0.14 mm longis differt. Typus: Palmerston, 12°29’S,
i30°58’E, December 1987, C. Cox [AQ408484] (holo: BRI).

Herbaceous perennial. Roots thinly fibrous-fusiform, to 80 mm long and | mm wide.
Stems cylindrical, upright, to 30 mm high, 1-2 mm diameter, up to 19 nodes on flowering
plants, finely pubescent with uniseriate hairs; internode length variable to 25 mm. Leaves
linear-lanceolate, to 80 mm long, 1-8 mm wide, main rib single, secondary veins 3 or
4, sparsely pubescent with uniseriate hairs. Flowers borne on top 2-10 nodes. Inflorescence
an umbelliform cyme appearing at nodes between the pairs of leaves, 1-6-flowered:
peduncles finely pubescent, 2-11 mm long, c. 0.5 mm diameter. Flower pedicels 3-7
mm long, c. 0.5 mm diameter, finely pubescent with uniseriate hairs, filiform, pendulous
during anthesis. Calyx without basal glands; sepals linear-lanceolate, c. 1.5 mm long, c.
1.0 mm wide, generally glabrous but with a few sparse cilia on the edges. Corolla deeply
)-parted, c. 5 mm long, 4 mm diameter; petals twisted in bud, linear-lanceolate, generally
glabrous but with a few sparse cilia on the top edges, c. 5 mm long, 1.5 mm wide,
cream. Corona a continuous ring around the base of the staminal column, double lobed,
c. 2 mm long, 2.5 mm diameter, cream; main (inner) lobe obloid-cylindrical with tip
rounded, curving inwards over the stamens, c. | mm long and 0.5 mm diameter:
secondary (outer) lobe a small protuberance at base of the main lobe. Staminal column
c. 2 mm long, 1.5 mm diameter with corona adnate to bottom third. Anther connectives
incurved; apical appendages acute, c. | mm long, | mm wide; slit between anther wings
0.5 mm long. Pollinarium 0.44 mm long, 0.6 mm wide; pollinia pendulous, oblong,
golden, 0.21 mm long, 0.13 mm wide; corpusculum 0.20 mm long, 0.11 mm wide,
oblong, tan; caudicles 0.14 mm long, 0.05 mm wide. Style-head conical, cream, c. 1 mm
wide and extending c. | mm above the anthers. Follicles fusiform, glabrous, c. 4.5 cm
long and c. 5 mm diameter. Seed 9 mm long, 3 mm wide, ovate, tan; coma absent. Fig.
2.

Forster, Asclepiadaceae, ! 113

Specimens examined: Northern Territory. DARWIN AND GULF District: Leander Swamp Bombing Range, Jan
1969, Byrnes NB1284 (DNA); Palmerston, Dec 1987, Tingey [AQ408486] (BRI*); ditto, Sep 1988, Tingey (BRD.

Distribution and habitat: This species occurs in the ARNM Region of Barlow (1985)
and is recorded from cleared black soil/sandy loam in association with Xanthostemon
paradoxus, Terminalia sp., Eucalyptus miniata, Pleurocarpaea denticulata, Hypoxis
marginata and Brunontella sp.

Phenology: Flowering occurs from December to January.

a!

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atta
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*
+
it

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Fig. 2. Cynanchum christineae. A. habit X 0.5. B. apical view of flower x 3. C. view of single flowered cyme X

3. D. apical view of staminal corona and column X 19. E. lateral view of staminal corona x 19. F. pollinarium
x 35, All from spirit material of Tingey AQ408486. Del. K. Harold.

appa:

AMMEN SIGE DON EON CDP EN

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COCO OR ee re Lr cCLCceCecLons

i14 Austrobatleya 3(1): 1989

Conservation status: As with C. /iebiana, further populations may be found in suitable
habitats in northern Australia. The status of the population at the Leander Bombing
Range requires investigation. A conservation status of 1E is provisionally given.

Etymology: Named for Mrs Christine (Chris) Cox who together with Mrs Rita Tingey
and Mrs Joyce Stobo, has made many useful collections of Asclepiadaceae from around
the Darwin-—Palmerston area.

SARCOSTEMMA R. Br.

Over the last few years a number of specimens in Australian herbaria have been
identified as Sarcostemma esculentum (L.f.) Holm. While most of these specimens have
been previously undetermined, those from Western Australia have usually been referred
to as Marsdenia brockmaniana W. Fitzg. Fitzgerald (1918) provided a relatively detailed
description of his new Marsdenia, which clearly shows that it is conspecific with
Sarcostemma esculentum.

This species is most commonly placed in the small genus Oxystelma R. Br. The
relegation of Oxystelma to subgeneric status under Sarcostemma by Holm (1950) has
not been followed by subsequent authors (Bullock 1956; Huber 1983; Ali 1983). Both
Huber and Ali take very narrow views of the genera they provided accounts of, with
many of the key diagnostic characters they used representing intergrades of character
states. Huber (l.c.) stated concerning Sarcostemma and Oxystelma, “the shallowly lobed
corolla, the pubescent outer corona attached to the corolla and the lanceolate tips of the
inner corona segments clearly distinguish [Oxystelma from Sarcostemma sens. strict.)”.

However S. insigne (N.E. Br.) Descoings from Madagascar has a shallowly lobed
corolla (Descoings 1961), but does not have a “pubescent outer corona” (in reality merely
an area with hairs present) or lanceolate tipped corona lobes. Although the presence of
hairs has been a much used character in distinguishing both genera and species in the
Asclepiadaceae, often the presence/absence of hairs may be quite variable. This has been
found in such genera as Ceropegia L. (Field & Collenette 1984; Bruyns 1985), Stapelia
L. (Leach 1985), Brachystelma Sims (Forster 1986; Forster 1988), Caralluma R. Br.
(Bruyns 1986, 1987) or Marsdenia R. Br. (Forster unpubl. data). Consequently I consider
attributes of the indumentum overrated as diagnostic characters in the Asclepiadaceae.

The degree and shape of lobing of the staminal corona in the Asclepiadaceae is
also variable (e.g. in Stapelia (Leach 1985), Brachystelma (Dyer 1983), Ceropegia (Dyer
1983; Bruyns 1985) and Caralluma (Bruyns 1986, 1987) and has also been greatly
overrated in both specific and generic classifications of the family. When the variation
of the staminal corona lobes of species in large genera like Marsdenia, Ceropegia,
Brachystelma, Tylophora R. Br. and Stapelia L. is taken into account, the linear-tipped
corona lobes in Sarcostemma esculentum are not all that distinctive compared with the
globose lobes of other species of the genus. Hence it is considered that the corolla
indumentum and linear-tipped corona lobes of S. esculentum are useful only in defining
the species. Holm’s (1950) placement of the species (in the subgenus Oxystelma (R. Br.)
Holm) 1s therefore supported here.

Sarcostemma esculentum (L.f.) Holm, Ann. Missouri Bot. Gard. 37: 482 (1950); Oxystelma

esculentum (L.f.) R. Br. ex Schultes, Syst. Veg. 6: 89 (1820); Periploca esculenta
Lf, Suppl. pl. 168 (1781). Type: Koenig, Linnean Herbarium 307/7 (holo:
LINN(microfiche!)).
Sprengel, Syst. Veg. 1: 850 (1825); Wight, Contr. bot. India 54 (1834); Hook.,
Companion Bot. Mag. 2: 52-53, t. 22 (1836); Decne in DC., Prod. 8: 542 (1844):
Wallich, Numer. List 8219-8220 (1828-1849); Griffith, Not. pl. asiat. 4: 61 (1854);
Griffith, Ic. pl. asiat. t. 399, 400 (1854); Miq., Fl. Ned. Ind. 2: 483 (1856): Dalz.
& Gibson, Bombay fl. 150 (1861); Benth. in Benth. & J.D. Hook., Gen. pl. 2:
749 (1876); J.D. Hook., Fl. Brit. India 4: 17 (1885); K. Schum. in Engl. & Prantl.
Nat. Pilanzenfam. 4(2): 229 (1897); Tsiang, Sunyatsenia 4: 102 (1939): Bakhuizen
van den Brink, Blumea 6: 371 (1950); Backer & Bakhuizen van den Brink, FI.
Java 3: 250 (1965). H. Huber, Fl. Ceylon Handb. 1: 38 (1973); Rev. Handb. f1.
Ceylon 4: 83-84 (1983); Ali, Fl. Pakistan 150: 20-23 (1983).

Forster, Asclepiadaceae, | 1i5

Oxystelma wallichii Wight, Contr. bot. India 54 (1834). Type: Adripas Gangis atque
Irawaddi, Wallich (syn: K-W(microfiche!)); Pathacottah, in prov. Tanjorenst?,
Wight (syn: K(microfiche!)).

Decne in DC., Prod. 8: 542 (1844); Wallich, Numer. List 8220 (1828-1849).

Marsdenia brockmaniana W. Fitzg., J. & Proc. Roy. Soc. Western Australia 3: 197
(1918). synon. nov. Type: Devils Pass, Napier Range, Western Australia, May
1905, WV. Fitzgerald 626 (holo: PERTH).

Slender vine, perennial, glabrous, with white latex. Stems much branched, cylindrical,
to 2 mm diameter; grey-green to green; internode length variable to 12 cm long. Leaves
linear-lanceolate, glabrous, lacking extrafloral nectaries, pale green, seasonally deciduous;
lamina 3-9 cm long, 3-9 mm wide; tips acute; petiole 0.5-1.3 cm long, c. 1 mm diameter.
Flowers borne on top 2-6 nodes. Cymes appearing at the nodes between the pair of
leaves, 1-4-flowered; peduncles 4~5 cm long, c. 1 mm diameter, pale green. Pedicels 1-
2 cm long, c. | mm diameter, pale green. Flowers c. 7 mm long, 15-25 mm diameter;
cream with irregular pale pink stripes or pale rose. Sepals 5, distinct, acute, glabrous, 2-
3.5 mm long, c. | mm wide, generally with 1-3 glands at base on the adaxial side.
Corolla shallowly campanulate to rotate, glabrous outside, lobes about as long as tube,
imbricate in bud: corolline corona of dense hairs present at base adjacent to staminal
column, c. | mm high, 4 mm diameter; petals lanceolate, 5-10 mm long, c. 7 mm wide.
Staminal corona of 5 separate lobes adnate to staminal column below anthers, c. 6 mm
long, 4 mm diameter; lobes ovate-lanceolate, tips incurved over anthers, c. 5 mm long
and 1.5 mm wide at base, cream. Staminal column c. 4 mm long, 2 mm diameter.
Anther appendages acute, c. 1 mm long, | mm wide. Slit between anther wings c. 2—-2.5
mm long. Style-head depressed, globular, pentagonal, c. | mm long, | mm wide. Ovaries
free, glabrous, c. 2.5 mm long, 1.5 mm wide. Pollinaria 1.45 mm long, 0.4 mm wide;
pollinia pendulous, attached to base of corpusculum, oblong-ovoid, 1.1 mm long, 0.28-
0.3 mm wide: corpusculum 0.38-0.4 mm long, 0.23-0.24 mm wide; caudicles 0.15 mm
long, 0.06 mm wide. Follicles fusiform, 4-8 cm long, c. 1 cm diameter, glabrous; seed
broadly ovate, tan, 3.5-4 mm long; coma white, attached to micropylar end of seed, c.
20 mm long.

Specimens examined. Sri Lanka. NORTHERN PROVINCE, MANNAR District: c. 8 miles [13 km] NE of Manta
along the road to Pooneryni, near mile marker 14/3, Dec 1974, Davidse & Sumithraanachchi 9167 (BRI; PDA
n.y.). Australia. Western Australia. Bank of Ord River, 5 km W of Kununurra, 15°46'S, 128°41"E, Mar 1978,
Paijmans 2556 (CANB,PERTH); 25.3 km WSW of Mt Blythe on Charnley River, 16°23’S, 125°13’E, Feb 1989,
Kenneally 10927 & Hyland (BRITPERTH); Near Uramurdah Ck, 16°39’S, 120°2S’E, Apr 1978, Craven 5310
(CANB); Ord River, 17°24’S8, 128°52’E, Sep 1949, Langfield 105 (PERTH); Barker River Gorge, Oct 1921, Gardner
447a (PERTH); Kalumburu, Dec 1973, Crawford 10 (PERTH). Northern Territory. DARWIN AND GULF DISTRICT:
“of the South Alligator’, undated, Anon. (P, photo at BRI); Mainorn, Jan 1963, Barlow 623 (BRI,JCT); Peron
Is, 13°$1’S, 130°O1’E, Oct 1974, Henshall 830 (ADW,CANB,DNA,MEL,NT). :

Distribution and habitat: In Australia S. esculentum occurs in the KIMB and ARNM
Regions of Barlow (1985) (Map 2) where it has been recorded from alluvial mulga
woodland or grasslands fringing swamps.

Phenology: In Australia flowering occurs from October to March and fruiting two to
four months after flowering.

Notes: The species is a popular subject for experimental purposes having been used in
studies of embryology (Devi & Lakshminarayana 1977), chemistry (Trivedi ef ai. 1988),
leaf venation (Mohan & Inamdar 1984), plasmodesmata in trichomes (Inamdar ef al.
1973) and floral anatomy (Rao & Ganguli 1963; Puri & Shiam 1966).

Conservation status: Not endangered, vulnerable or rare in Australia at present. 5S.
esculentum is widely distributed in Egypt, Pakistan, India, Iraq, Sri Lanka, China,
Thailand, Burma and Indonesia (Tsiang 1939; Backer & Bakhuizen van den Brink 1965;
Ali 1983; Huber 1983).

TYLOPHORA Rk. Br.

Tylophora erecta F. Muell. ex Benth. and T. macrophylla Benth. were published
simultaneously by Bentham (1869) with two syntypes cited for each respectively. Bentham
distinguished his two taxa in the accompanying species key:

116 Austrobaileya 3(1): 1989

“Leaves narrow-lanceolate.............. 0.0.0.2... l. Z. erecta
Leaves ovate-lanceolate ...........0. 0.0.0.0... 2. T. macrophylla”

In his two descriptions, the only other difference seems to be corolla size, with T.
macrophylla having only slightly larger flowers.

Wide variation in leaf size occurs in material identified as T. erecta and T
macrophylla in herbaria. Leaf lamina length and width in relation to stem node number
was examined in material of Forster & Bolton 3707, a large multiple collection from
four different plants within the same population. Lamina length/width ratio was found
to be relatively constant (Fig. 3A). Leaf lamina length and lamina width were found to
increase until node 4 below the stem apex after which both were relatively constant (Fig.
3B,C). Considerable variation occurred in these dimensions. Hence the size of a particular
leaf on a 7. erecta individual (and consequently on an herbarium specimen) may be
dependent on both the conditions under which growth occurred and also the number of
nodes available for examination. For small specimens with few nodes the larger leaves
towards the base of the stem tend to be absent. Therefore the leaf shape difference given
by Bentham (1869) to distinguish species, which is dependent on the lamina length/
width ratio is not considered a valid distinguishing feature.

Flower size was found to be variable within populations (e.g. Forster & Bolton
3705A-D; Forster & Bolton 3707A-D) with the corolla diameter varying from 9-12 mm
and petals 4-6 mm long. Therefore the difference in scale given by Bentham (viz 3 lines
diameter for T. erecta versus nearly 4 lines diameter for 7. macrophylla) is not sufficient
for the recognition of two species.

Tylophora erecta F. Muell. ex Benth., Fl. austral. 4: 334 (1869); Vincetoxicum erectum

(F. Muell. ex Benth.) O. Kuntze, Revis. gen. pl. 2: 424 (1891). Type: Burdekin
River, undated, F. von Mueller (syn: K(photo!); isosyn: MEL(MEL 113602)):
Sellheim & Bowen Rivers, undated, Bowman (syn: K, both on same sheet(photo!);
rein oe 113604 in part, excl. Nebo River); MEL(Sellheim River, MEL
113603)).
F. Muell., Fragm. 9: 71 (1875); Fragm. 11: 79 (1878); Syst. census Austral. pl. 94
(1882); Second syst. census Austral. pl. 158 (1889); Bailey, Syn. Queensl. fl. 316
(1883); Catal. pl. Queensl. 30 (1890); Bailey, Queensl. fl. 3: 1005 (1900); Compr.
cat. Queensl. pl. 326 (1913); Domin, Biblioth. Bot. 89 (6): 1085-1086 (1920):
Williams, Native pl. Queensland 3: 310-311 (1988).

T'ylophora macrophylla Benth., Fl. austral. 4: 334 (1869); Vincetoxicum benthamiana
O. Kuntze, Revis. gen. pl. 2: 424 (1891) synon. nov. non V. macrophylla Sieber
& Zucc, (1846). Type: Port Essington, Feb 1839, Armstrong (syn: K(photo!));
Adam’s Bay, undated, Hulls (syn: MEL(MEL 113619)).

Tylophora stelligera Schitr., Bot. Jahrb. Syst. 40, Beibl. 92: 5 (1908). synon. nov.
Type: Islands of Torres Strait, Hartmann (holo: Bt; iso: MEL(MEL 113610)).

Woody erect perennial, with dense indumentum of uniseriate yellow hairs; with yellow
latex. Stems cylindrical, c. 1 m high, 1-4 mm diameter, yellowish green to grey-green:
up to 30 nodes on flowering plants; internode length variable to 8 cm. Leaves linear-
lanceolate to ovate-elliptical; lamina up to 14 cm long and 6 cm wide, upper surface
yellowish green to grey-green, paler beneath; petiole 5-8 mm long and 1 mm wide: 4
extrafloral nectaries at lamina base. Flowers borne on upper 2-10 nodes. Cymes appearing
at the nodes between the pair of leaves, |-many-flowered; peduncle 10-60 mm long, 1-
1.5 mm _ diameter, pale green. Pedicels 6-10 mm long, c. 1 mm diameter, pale green.
Flower 3-4 mm long, 9-12 mm diameter; tube up to | mm deep; dark brown, tan, red,
reddish brown or green. Sepals 5, distinct, lanceolate 3-4 mm long, [-1.5 mm wide,
generally with 3 or 4 glands at the base on the adaxial side. Petals 5, distinct, ovate-
lanceolate, externally pubescent with white cilia, internally glabrous or with sparse white
cilia, 4-6 mm long, 2.5-4 mm wide. Corona of 5 separate lobes adnate to staminal
column below anthers, 0.5-1 mm long, 2.5-3 mm diameter; lobes rounded, c. 0.75 mm
long, 0.5 mm wide, dark brown, tan, red, reddish brown or green. Staminal column c.
1.5 mm long, | mm diameter. Anther appendages obtuse, c. 0.25 mm long, 0.5 mm
wide. Slit between anther wings c. 0.25 mm long. Style-head depressed globose, with a
bifid style, 0.5-1.0 mm longer than staminal column, 1-1.25 mm diameter, pale green.

Forster, Asclepiadaceae, | 117

non © a OFF DO N @

Lamina length/width ratio

Lamina length (mm)
~ » © & OA OO
© 0 o oc Oo @ oO

©

25

20

15

10

Lamina width (mm)

1 2 3 4 §5§ 6 FJ 8B Y 10 44 412 143 14
Node number below apex

Fig. 3. Tylophora erecta: A. lamina length/width ratio versus node number below apex. B. lamina length versus

node number below apex. C. lamina width versus node number below apex. All from Forster 3707. Del. L.
Jessup.

118 Austrobaileya 3(1): 1989

Pollinaria c. 0.15 mm long, 0.20 mm wide; pollinia globose, held erect, 0.13-0.15 mm
long, 0.11 mm wide; golden; corpusculum oblong, 0.06 mm long, 0.03 mm wide, tan;
caudicle 0.04 mm long, 0.01 mm wide. Ovaries free, glabrous, c. | mm long and 1.25
mm wide. Follicles fusiform, 5-11 cm long, 12-22 mm wide; seed oblong, tan, c. 6 mm
long and 4 mm wide; coma white, attached to germinating end of seed, c. 20 mm long.
Figs 4 & §.

Selected specimens: Northern Territory. Old Marrakai Track, 47 mile, Stuart Hwy, £2°54’S, 131°10’E, Dec 1987,
Cox [AQ408474] (BRI*); Marlowe’s Lagoon, Palmerston, Dec 1987, Tingey [AQ408476] (BRI*); Adelaide River
Camp, 13°15’S, [31°10Q’E, Aug 1946, Blake 16688 (BRI); Bank of Daly River, [3°53’S, 130°48’E, Jul 1946, Blake
16481 (BRI); near Bull Ck, 13°53’S, 131°16@’E, Jul 1946, Blake 16286 (BRI); near Beatrice Hill, 12°38’S, 131°16’E,
Sep 1946, Blake 16980 (BRI). Queensland. Cook District: 1.3 km N of Duckholes Ck crossing, Peninsula Road,
14°24’S, 143° 22’E, Apr 1988, Forster 4252 & Liddie (BRI*,CANB,K); 4.5 km N of old homestead site on Cape
York Rd, Lockerbie, Sep 1985, Williams 85199 (BRI); Just E of Myall Ck on the road from Weipa to Sudley,

1 2°39’S, 142° 16’E, Aug 1983, Clarkson 4886 (BRI,QRS); Laradeeniya Ck, Sep 1963, Stephens [AQ431122] (BRI):
3km W of Peninsula Development Rd on track to Kooburra Stn, 15°18’S, 143° SE, Apr 1980, Clarkson 3151
(BRI); 1i km W of Peninsula Development Rd on track to New Dixie Stn, 15°O1S. 143°30°E, Apr L980, Clarkson
3137 (BRI); Archer Bend N.P., 13°39’S, 141°20’E, Aug 1981, Aforton AM1308 (BRI); Long Scrub, Bamaga, 1962,
Webb & Tracey 6921 (BRLCANB). BURKE District: Chudieigh Park, Oct 1961, Copeman [AQ216955] (BRI).
NorTH KENNEDY DistTrRIcT: 16 km past “Lolworth” HS on Pentland Rd, 20°17’S, 145°00’E, Mar 1988, Forster
3705A-D & Bolton (BRI*,CANB,K,MO), 3.7 km by road NE of Ravenswood, 20°0S’S, 146°55’E, Mar 1988,
Forster 3707A-D & Bolton (BRI*¥,CANB,K,MO); Scrubby Ck, S of Rishton Scrub, 20°11’S, 146°34’E, Mar 1988,
Forster 3776 (BRI*,CANB,K); 2.5 miles [4 km] NW of Collinsville, 20°30’S, 147°50’E, Nov 1968, Ozanne
[AQ216959] (BRI). SoUTH KENNEDY District: Collinsville Coal Company Lease area, Jun 1984, Thompson
C161 (BRI. LEICHHARDT District: Southern end of Lake Elphinstone, c. 6 km from road to Glenden, 21°33’S,
148°13’E, Jul 1986, Champion 173 (BRI).

Distribution and habitat: The species occurs in the ARNM, CARP, CYRK and BURD
Regions of Barlow (1985) (Map 1). Collections have been recorded from reddish lateritic
soils or pale grey podsols in open eucalypt forest in association with overstorey taxa
such as Eucalyptus confertiflora and Erythrophieum chlorostachys on Cape York Peninsula
or with Eucalyptus quadricostata near Pentland or Eucalyptus crebra near Ravenswood.

Phenology: Flowering occurs from November to April, fruiting from March to June.

Notes: The species has been implicated in cases of scouring in calves (Ozanne AQ216959)
and pounded roots were reputedly used as an remedy for toothache by aborigines
(Stephens AQ431122).

Conservation status: 7. erecta 1s widespread in north tropical Australia and is not
endangered at present.

When Bentham (1869) described 7ylophora floribunda for a taxon that occurs on
the east coast of Australia, he was obviously unaware of T. floribunda Miq. from Asia.
Subsequently Blake (1948) ‘provided a new name, 7. crebriflora for the Australian species
and this has universally been used since. However Tsiang (1936) had already recognised
the illegitimacy of Bentham’s name and substituted 7. benthamii for Bentham’s species.
The synonymy is thus as follows.

Tylophora benthamii Tsiang, Sunyatsenia 3: 231 (1936); 7ylophora floribunda Benth.,

Fl. austral. 4: 335 (1869), non Mig., Ann. Mus. Bot. 2: 128 (1866); Vincetoxicum
polyanthum ©. Kuntze, Revis. gen. pl. 2: 424 (1891); Tylophora crebriflora S.T.
Blake, Proc. Roy. Soc. Queensl. 59: 168 (1948) nom. superfl. non Vincetoxicum
polyanthum K. Schum., Bot. Jahrb. Syst. 17: 136 (1893); non Tylophora polyantha
Volkens, Bot. Jahrb. Syst. 31: 473 (1902). Type: Mount Warning, Tweed River,
undated, C. Moore (holo: MELC(MEL 113591)).
Williams & Harden, Rainforest Climbing Pl. 37 (1980); Jacobs & Pickard, PI.
New South Wales 70 (1981); Willams, Students fl. N. E. New South Wales 5:
729-730 (1984): Stanley & Ross, Fi. S. E. Queensland 2: 313 (1986); Jones &
Gray, Climbing pl. Austral. 351, 355 (1988); Williams, Native pl. Queensland 3:
310-311 (1988).

Woody perennial liane, with dense to sparse indumentum of uniseriate hairs; with yellow
latex. Stems erect, cylindrical, twining; grey green becoming corky with age, to 2 cm
diameter; internode length variable to 13 cm long. Leaves petiolate; lamina broadly
ovate to elliptic with base cordate, rounded or truncate and tip mucronate or abruptly
acuminate, 4-12 cm long, 2.5-8 cm wide, 5-7 main veins prominent below, deep glossy
sreen above, pale green to bluish-green below, sparsely pubescent to glabrous above,

Forster, Asciepiadaceae, | 119

= axe,! ‘47
& +F ‘Ls ee a
= + gPF a =
* 7 = *45 = +
= a7 Fe, Sion “i
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« * = * tout ta 5 sa et 47
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Fig. 4. Tylophora erecta: A. habit X 1. B. apical view of part of calyx x 5. C. lateral view of flower x 4.5. D.
apical view of flower < 4.5. E. apical view of staminal corona and staminal column and representative hairs on
corolla X 21. F. lateral view of staminal corona and staminal column X 21. G. pollinarium X 37. All from spirit
material of Forster 3705A & Bolton. Del. K. Harold.

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RRP er rere re rccorrcecereer rrr r rs

[20 Austrobaileya 3(1): 1989

tll
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5. Tylophora erecta: A. compound cyme X |. B. apical view of part of calyx x 7. C. lateral view of flower
3. (Note: entire surface pubescent except for a few broad interior corolla margins.) D. apical view of flower
5. E. apical view of staminal column and staminal corona X 21. F. lateral view of staminal corona and
staminai column X 21. G. pollinarium X 37. All from spirit material of Forster 4252 & Liddle. Del. K. Harold.

Forster, Asclepiadaceae, | [21

sparsely to densely pubescent below; petiole 10-33 mm long, c. | mm diameter; 2-9
extrafloral nectaries at lamina base. Flowers borne on upper 2-5 nodes. Cymes appearing
at the nodes between the pairs of leaves; |-many-flowered, up to 8 cm wide; bracts on
cyme, linear-lanceolate, c. 4 mm long, 0.5 mm wide; peduncle to 15 mm long, c. 1 mm
diameter, pale green; pubescent. Pedicels 7-15 mm long, 0.5—1 mm diameter, pubescent,
light green. Flowers rotate, purple, brown or red, 3-7 mm long, 7-11 mm diameter:
tube up to 2 mm deep. Sepals 5, distinct, linear-lanceolate, 2.5-4 mm long, 1-1.5 mm
wide, sparsely pubescent externally; generally with 1-2 glands at the base on the adaxial
side. Petals 5, distinct, lanceolate, 3-5 mm long, 2~3 mm wide, finely pubescent externally
and internally. Corona of 5 separate lobes adnate to stamina! column below anthers, 1-
2 mm long, 1.5-4 mm diameter; lobes flattened, tongue shaped, 1.5-1.75 mm long,
0.75-1 mm wide; same colour as corolla. Staminal column 1-2 mm long, c. 1.5 mm
diameter. Anther appendages ovate-obtuse, c. 0.25-0.5 mm long, c. 0.25-0.5 mm wide.
Slit between anther wings 0.25-0.4 mm long. Style-head flattened pentagonal, with a
bifid style, c. 1.5 mm wide, pale green, 0.5-1.0 mm longer than staminal column.
Pollinaria 0.15-0.16 mm long, 0.3-0.4 mm wide; pollinia globose, held horizontally to
erect, 0.13-0.15 mm long, 0.12-0.15 mm wide, golden; corpusculum oblong, tan, c. 0.10
mm long, 0.05 mm wide; caudicles unwinged, 0.05-0.07 mm long, 0.01 mm wide.
Ovaries free, few sparse hairs, c. 2 mm long, 1.5 mm wide. Follicles fusiform, 9-12 cm
long, 1-2 cm wide; seed oblong, tan, 8-9 mm long, 4 mm wide; coma 20-25 mm long,
attached to germinating end of seed. Fig. 6.

Selected specimens: Northern Territory. DARWIN AND GULF District: Murganella Area, 11°26’S, 132°58’E, Feb
1984, Jones 1387 (BRI,DNA*); 6 km 8S of Brogden Pt, Murganella, 11°34’S, 133°04’E, Mar 1987, Russell-Smiith
1985 & Lucas (BRI,DNA*); Little Nourlangie Rock, 12°52’S, 132°48’E, May 1978, Dunlop 4826 (DNA); Lightning
Dreaming, Arnhemiand, 12°55’S, 133°02’E, Feb 1984, Dunlop 6592 & Wightman (DNA); East Alligator River,
10 km 8 of Cannon Hill Ranger Stn, 12°27'S, 132°58’E, May 1980, Craven 6010 (CANB,DNA); Katherine Gorge
N.P., 14°19’S, 132°28’E, Jun 1983, King 237 (DNA); Liverpool River mouth, Tudor Pt, 11°59’S, 134°15’E, Jun
1987, Russell-Smith 2564 & Lucas (DNA); Cobourg Peninsula, 11°09’S, 132°09’E, Jul 1982, Wightman 137 &
Dunlop (DNA); Goomadeer: Gundjumbul outlier, 12°04’S, 133°44’E, Oct 1987, Russell-Smuth 3834 & Lucas
(DNA). Queensland. Cook District: Tolga, Nov 1985, Williams 85310 (BRI); Bamaga, c. 27 km SW of Cape
York, Oct 1965, Smith 12664 (BRD; cult. ex S.F.R. 144, 16°17/S, 145°0S’E, Nov 1978, Gray 1120 (QRS*);
Wongabel, 17°20’S, 145°28’E, Sep 1957, Smith 10171 (BRI). NortTH KENNEDY District: Wilkin Hill, Hinchinbrook
Is, L8°16’S, 146°14VE, May 1972, Webb & Tracey 12006 (BRI,CANB); Tully, Dec 1949, Clemens [AQ217019]
(BRI). WIDE Bay District: Mt Allan, Booloumba Ck S.F.R. near Kenilworth, Mar 1987, Sharpe 4659 (BRI*);
Cold Ck, S.F. 301, 25°53’S, 152°24’E, Nov 1988, Forster 4839 (BRI*). MoRETON District: W Slopes of Mt
Roberts, McPherson Range, Mar 1945, Blake 15483 (BRI); c. | km S of Coolum Beach P.O., Apr 1978, Sharpe
2486 (BRI). New South Wales. Fingal Head, May 1967, Blake 22789 (BRI).

Distribution and habitat: Near the coast from Sawtell in New South Wales to the top
of Cape York Peninsula in Queensland and into the north of the Northern Territory
which equates to the MCPH, CYRK, BURD, ARNM and DWSN Regions of Barlow
(1985) (Map 2). It occurs in a wide variety of rainforest types from littoral rainforest at
low altitudes to higher altitude rainforests on the Atherton Tableland (e.g. Tolga and
Wongabel).

Phenology: Flowering and fruiting throughout the year.

Conservation status: While the species is widespread in northern Australia, it is markedly
disjunct and very rarely encountered in southeastern Queensland and northeastern New
South Wales. It is not endangered, threatened or vulnerable at this stage.

SARCOLOBUS R. Br.

In his revision of Sarcolobus R. Br., Rintz (1980) lists the type citation for S.
ciliolatus as being “Moluccas, Aroe & Kei Is., Warburg 21317” with an isotype being
present at A. Material labelled S. ciliolatus Warb. (a synonym of S. retusus K. Schum.)
exists at BRI with the label formation “Plants of Key Arch. 1889 Warburg 21317”.
There have been numerous examples of specimens with different label information
forming part of the same collection in material collected during the 19th century,
especially with duplicated collections at different herbaria (Merrill 1915). If the collections
have identical collection numbers (as opposed to the species numbers used by some
collectors, which generally do not run into 5 digits) and are of the same species, and
there 1s no information available on the location of duplicates, it is reasonable to assume
that they represent the same collection. The BRI material of Warburg 21317 is referable
to S. refusus and 1s therefore considered to represent an isotype of S. ciliolatus.

Fig. 6.

pollinarium (lacking { pollintum) X 37. All from Gray 1534. Del. K. Harold.

Austrobaileya 3(1): 1989

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ees
=
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eee Veet
= at ¢e* += + Fe ei

. part of cyme X !{. B. apical view of part of calyx showing glands X 3. C, latera
view of flower * 4.5. (Note: entire surface pubescent except for a few broad interior corolla margins.) D. apica
view of flower X 4.5. E. apical view of staminal corona and staminal column, showing some hairs on corolla x
15. F. lateral view of staminal corona and staminal column showing hairs at base of staminal corona X 15. G.

Forster, Asclepiadaceae, | 123

Isotype material of S. oblongus Rintz also exists at BRI in addition to the holotype
(A) and isotypes (BM,L) listed by Rintz (l.c.: 77). Rintz cited material only from the
southern coast of Papua New Guinea, from the Fly River and Purari River deltas. S.
oblongus also occurs in the Solomon Islands and the northern coast of Papua New
Guinea.

Specimens examined: Papua New Guinea. WESTERN District: Daru Is, Mar 1936, Brass 6278 (BRI; A v.yv.), Sep
1972, Streimann & Lelean NGF18424 (BRI; CANB,L,LAE i.y.). MADANG District: Madang, Croft & Galore
LAE71097 (BRI; A,CANB,K,L,LAE n.y.). GULF District: Uramu Is, 7°30’S, 144°30’E, Jul 1955, Gray & Floyd
NGF8032 (BRI; LAE n.y.). Solomon Islands. Meringe Lagoon, Isabel {Santa Isabel Island], Nov 1932, Brass 3156
(BRI; A w.¥.).

FINLAYSONIA Wall. (subfamily Periplocoideae)

The monotypic genus Fin/aysonia was first described by Wallich (1831) who also
provided an excellent coloured plate and detailed description of F. obovata Wallich.

F. obovata, though until now considered non Australian, does occur in Australia
and represents a third genus in the subfamily Periplocoideae. Finlaysonia may be
distinguished from Cryptostegia R. Br. and Gymnanthera R. Br. by the following key.

|. Corolla without tube; lobes pubescent ...................... Finlaysonia
Corolla with tube; lobes glabrous

2. Corona lobes broad, cusped or toothed; corolla tube cylindrical .. Gymnanthera
Corona lobes pointed or forked; corolla tube funnel-shaped ...... Cryptostegia

Finlaysonia Wallich, Pl. asiat. rar. 2: 48, t. 162 (1831). Type: F. obovata Wallich

Wight, Contr. bot. India 65 (1834); Decne in DC., Prod. 8: 494 (1844): Miq., FI.
Ned. Ind. 2: 464-465 (1856); Griffith, Notul. pl. asiat. 4: 70 (1854); Benth. in
Benth. & J.D. Hook., Gen. pl. 743 (1876); J.D. Hook., Fl. Brit. India 4: 7-8
(1885); K. Schum. in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 211 (1897); Merr.,
Interpr. Herb. amboin. 434 (1917), Sp. blancoan. 314 (1918), Enum. Philipp. fl.
pl. 339 (1923); Ridley, Fl. Malay Penins. 2: 373 (1923); Backer & Bakhuizen van
den Brink, Fl. Java 2: 248 (1965).

Woody liane with white latex. Stems twining, glabrous; with papery bark. Leaves slightly
fleshy, glabrous, opposite, margins entire, lacking extrafloral nectaries at lamina base.
Inflorescence appearing at nodes between the pair of leaves, cymes with I-many flowers.
Sepals 5, distinct, without glands at base. Corolla rotate; petals 5, distinct, lobes
overlapping to right in bud. Corolline corona of 5 filiform-subulate lobes inserted adnate
to the stamens. Stamens inserted in corolla throat, filaments free, not alternating with
corona lobes; anthers with inflexed membranous tip. Pollinaria comprising a granular
mass with 2 pollen masses in each anther cell; translators erect, dilated. Follicles widely
divergent, ovoid with a narrow base and hooked acuminate top, ribbed. Seed smooth:
coma short, reflexed.

Finlaysonia obovata Wallich, Pl. asiat. rar. 2: 48, t. 162(1831), Numer. List 4466 (1831).
Type: Kaluen River, 2 December 1827 (syn: K-W(4466A, photo!)); Sundriban 23
sep 1809, ex Hb. Ham. (syn: K-W(4466B, photo!)).

Tabernaemontana cirrhosa Blanco, Fl. Filip. 115 (1837). Type: not designated.
Blanco, Fl. Filip. Ed. 2, 83 (1845), Ed. 3. I: 152 (1877); A. DC., Prod. 8: 377
(1844): Merr., Sp. blancoan. 314 (1918).

[Finlaysonia maritima auct. non Backer ex K. Heyne: Heyne, Nutt. pl. Ned.-Ind.
ed. 2, 2: 1293 (1927).]

Stems to 8 mm diameter. Leaves petiolate; lamina obovate-oblong, oblanceolate or
elliptic, 40-125 mm long, 16-60 mm wide; base cuneate to acute, tip obtuse or apiculate:
dark glossy green above, paler beneath, coriaceous, conspicuously reticulate-veined
beneath; petioles 7~20 mm long, c. | mm diameter, top grooved, with an occasional
uniseriate hair. Inflorescence with sparse to dense uniseriate hairs, up to 10 cm wide.
Peduncle with occasional uniseriate hairs, to 2 mm diameter; bracts ovate, with scattered

124 Austrobaileya 3(1); 1989

uniseriate hairs on edge, c. | mm long, 1.5 mm wide. Flower to 15 mm diameter, c. 5
mm long. Sepals ovate, uniseriate hairs on edge, c. 1.5 mm long, 2 mm wide. Petals c.
7 mm long, 5 mm wide, yellow-brown to green, lower surface glabrous, upper surface
with lower 4 mm covered in dense uniseriate multicellular hairs. Corona c. 2.5 mm
long, 4 mm diameter, consisting of 5 separate lobes/filaments fused to base of corolla
lobes: each lobe rounded at base, c. 1.5 mm diameter, linear-subulate above for c. 3.5
mm in length and 0.25 mm diameter. Stamens fused to base of corolla filament, not
connate, enclosing stigma; individual stamens c. 1.5 mm long, 0.75 mm wide; slit between
anther wings c. 0.75 mm long. Style-head on stalked style, c. | mm long, | mm diameter.
Pollen mass c. 0.5 mm long, 0.25 mm wide. Follicles ribbed, 5-8 cm long. Seed ovate,
tan, c. 18 mm long, 10 mm wide; coma around the entire seed edge, c. 10 mm long.
Fig. 7.

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wt
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Fig. 7. Finlaysonia obovata: A. single fascicle from cyme X 1. B. apical view of flower x 1. C. apical view of
flower showing detail of staminal column, corona lobes and hairs on corolla (several corona lobes lacking upper
part) X 10. D. lateral view of flower showing detail of staminal column, corona lobes and hairs on corolla (several
corona lobes lacking upper part) X 10. All from spirit material of Craven 6559. Del. K. Harold.

Forster, Asclepiadaceae, | 125

Specimens examined: Singapore. Kranji Nature Reserve, 14 1/2 m [23.2 km] Woodlands Road, Aug 1965,
Ngadiman HMB376 (BRI; SENG n.v.), Papua New Guinea. GULF District: Kinomeri Village, Uramu Is, 7°30’S,
144°30’E, Jun 1955, Gray & Floyd NGF8009 (BRI; LAE #.y.); Paia Village near mouth of Omati River, Jan
1955, Womersiey & Simmonds NGF5060 (BRI; LAE n.v.); Tributary of Era River, 7°25’S, 144°46’E, Apr 1974,
Croft & Vinas LAE61302 (CANB; L,LAE a.v.). WESTERN Division: Daru Is, Feb 1936, Brass 6226 (BRI; A n.y.):
Upper Wassi Kussa River (left branch), Jan 1936, Brass 8623 (BRI; A n.v.); Sabi, lower Wassi Kussa River,
9°05'S, 142°O0"E, Jun 1973, Henty NGF49645 (BRI; A,CANB,L,LAE v.y.). Australia. Northern Territory. East
Alligator River, 19.5 km NW of Cannon Hill Ranger Stn, 12°14’S, 132°49.5’E, Jun 1980, Craven 6434 (CANB):
ditto, Mar 1981, Craven 6559 (CANB*).

Distribution and habitat: The species is widely distributed in India, South-east Asia,
Philippine Islands, New Guinea and known from a single locality in the Northern
Territory of Australia in the ARNM region of Barlow (1985) (Map 3). It grows as a
twiner among mangroves or in fringing forest, often inundated by water at high tide. In

Australia it is recorded from cracking clay on a semi-saline floodplain with mixed grasses
and herbs.

Phenology: Flowering in March and probably earlier. Wallich (1831) records August to
October in the northern hemisphere. Fruiting in June. Wallich (l.c.) records November
in the northern hemisphere.

Notes: There are two sheets in the Wallich herbarium at Kew (K-W) that are relevant
to typification of Finlaysonia obovata. The protologue lists Gurua obovata Herb. Hamilton
as a synonym and states the species as “Crescit ad litora aestuariorum Gangis, Irawaddi
et fluminum Martabaniae, florens Augusto-Octobre, fructifera Novembre.”. From Wal-
lich’s contemporaneous Numerical List, the following label has been attached to sheet
4466A in K-W:

4466 Finlaysonia obovata Wall
A Rupe fl. Martabonia 1827
B Gurua obovata HHam. [= herb. Hamilton] e Sunderban”’.

Both sheets A and B are therefore regarded as syntypes of Wallich’s name. As both
Sheets represent the same taxon, a lectotype is not selected here.

Material of F. obovata from Java was erroneously included under the name F.
maritima Backer ex Heyne by Heyne (1927). That name was based on Secamone
maritima Blume which applies to a species of Toxocarpus and is not referable to
Finlaysonia (Bakhuizen van den Brink 1950).

Conservation status: Apparently of restricted occurrence in Australia. Surveys of the
mangrove communities of northern Australia are required to ascertain the extent of this
species’ occurrence in Australia.

GYMNEMA R. Br.

Bentham (1869) did not accept Gymnema geminatum R. Br. for Australia, placing
the name in synonymy under “Gymnema sylvestre R. Br.” with the comment “The
species 1s also in Ceylon and the Indian Peninsula, and has probably a much wider
range in tropical Asia. R. Brown indicated the close resemblance of the Australian to
the Indian plant; and, after a careful analysis of the flowers of both, I can detect no
character to separate them unless the scales of the corolla may be slightly more decurrent
in the former than in the latter.” This name was not validly published until 1820 by
Schultes so G. geminatum therefore has priority over G. sy/vestre. Bailey (1900) followed
Bentham in accepting “G. sy/vestre R. Br.” and added another species, G. pleiadenium
F. Muell., described by von Mueller in (1878). G. pleiadenium and G. sylvestre sensu
Benth. were distinguished by Bailey with the following key.

“Leaves ovate to ovate-lanceolate. Umbels mostly in pairs .... 1. G. sylvestre
Leaves ovate, shortly acuminate, with many superficial glands at the
base. Umbels mostly solitary ets A) ee) ee 2. G. pleiadenium”

Examination of the type sheet of G. pleiadenium F. Muell. has found it to be
synonymous with the species described as G. dunnii (Maiden & Betche) P. Forster
(Forster 1987) and is hence the prior name. Examination of the type sheet of G. sy/vestre
(Retz.) R. Br. ex Schultes (Fig. 8) and other material from India has shown that the
Australian plants are specifically distinct. Vegetative material of G. sylvestre is almost

long, 0.06
2mm wide
ibrous, 4s

t, Aug 1969, .
ane road, Kai

126 Austrobaileya 3(1): 1989

indistinguishable from that of G. pleiadenium; however a number of floral and vegetative
differences may be used to distinguish both these species and G. geminatum (Table 2).

There are no comprehensive published descriptions of G. sylvestre and none of
the specimens cited here have mature fruits, but Hooker (1885) states that the follicles
are slender and terete and the immature follicle on Thomson AQ217135 (BRI) is similar
to those of G. pleiadenium. G. pleiadenium has fusiform follicles whereas G. geminatum
has fusiform to ovoid follicles. G. sylvestre (Retz.) R. Br. ex Schultes appears closely
related to both G. pleiadenium and G. geminatum but differs in a number of significant
ways as outlined above.

Table 2. Comparison of major distinguishing characters of Gymnema sylyestre, G. pleiadenium and
G. geminatum.

rn a a SS ees 0 0 eee

Character G. sylvestre G. pleiadenium G. geminatum
Sve Te gg en ee ee eS SS ee
secondary venation obscure obscure prominent

Corolla tube length c. 1.5 mm c. 2-3.5 mm c. 3.0 mm

Staminal corona of absent present absent

small protuberances

Corolline corona | present absent present
in small lobes extending
beyond corolla tube

Corpusculum length c. 0.12 mm 0.10-0.12 mm 0.14-0.15 mm
Pollinia length c. 0.26 mm 0.14-0.20 mm c. 0.12 mm
Follicles fusiform fusiform fusiform-ovoid

Gymnema sylvestre (Retz.) R. Br. ex Schultes, Syst. Veg. 6: 57 (1820); Periploca sylvestris
Retz., Observ. bot. 2: 15 (1791). Type: Koenig 6733 (lecto (here designated): LD).
Sprengel, Syst. Veg. 1: 844 (1825); Wight, Contr. bot. India 44 (1834); Wight,
Icon. pl. Ind. orient. 2: t. 349 (1840); Decne in DC., Prod. 8: 621 (1844); Thwaites,
Enum. pl. zeyl. 197 (1860); J.D. Hook., Fl. Brit. India. 4: 29 (1883); Trimen,
Handb. fl. Ceylon 3: 153 (1895); Tsiang & Li, Acta Phytotax. Sin. 12: 12-13
(1976); H. Huber, Rev. Handb. fl. Ceylon 4: 100-102 (1983).

Asclepias geminata Roxb., Fl. ind. ed. 1832 2: 45 (1832). Type: not designated.

Woody vine, perennial; with white latex. Stems twining; internode length variable to 7
cm. Leaves petiolate; lamina ovate, oblong-ovate to oblong-elliptic, up to 9 cm long and
5.5 cm wide, base rounded, truncate or shallowly cordate, apex acuminate or shortly
acuminate, pinnately veined, secondary venation not prominent, upper surface green to
grey-green, paler beneath, densely pubescent with uniseriate hairs, petiole 0.4-1.2 cm
long, c. 1 mm diameter; extrafloral nectaries 2-4 at lamina base. Flowers borne on upper
2-10 nodes. Cymes umbelliform, appearing at the nodes between the pair of leaves,
usually paired, 1-many-flowered; peduncle 2-4 mm long, c. 1 mm diameter, finely
pubescent, pale green. Pedicels 2-4 mm long, c. 0.25 mm diameter, finely pubescent.
Flowers cream, campanulate, c. 3 mm long, 3 mm diameter; tube up to 1.5 mm long.
Sepals 5, distinct, elliptic to oblong, externally finely pubescent, c. 1 mm long, | mm
wide, generally with 1-2 small glands at base on the adaxial side. Petals 5, distinct, acute
to oblong, externally glabrous, internally with few sparse hairs, semi-reflexed at anthesis,
1.52 mm long, c. | mm wide. Corolline corona of 5 thickened lobes c. 1 mm long,
running longitudinally down corolla tube and alternating with corolla lobes, densely
pubescent on edges; free ends of lobes erect, subulate, just exceeding corolla throat, 0.25-
0.5 mm long. Staminal column J-1.5 mm long, 0.75-1 mm diameter. Anther appendages
acute, 0.25-0.5 mm long, 0.25-0.5 mm wide. Slit between anther wings c. 0.75 mm long.
Style-head conical, c. 0.75 mm longer than staminal column, c. 0.75 mm diameter.
Pollinarium 0.15 mm long, 0.26 mm wide; pollinia oblong, held erect, 0.15 mm long,

Forster, Asclepiadaceae, |

0.07 mm wide, golden; cor oblong, 0.12 wide, tan; caudicles
attached to base of corpusculum, 0.06 mm long, 0.02 varies free, glabrous,
c. 0.75 wide. Follicles fusiform, glabrous, 4.5 cm long, lameter.

eed not seen.

cim ined: India. Hill beyond Bageshpur, Hassan District, Aug 1969
Shivadi Ghort, May 1969, Saldanha 13395 (BRI; US n.v.); Kadamane road, Ramamoorthy
aisor & Carnatic, Thomson [AQ217135] (BRI; K n.v.).

*
*

Saldanha 14378 (BRI; US n.¥.);
FP1892 (BRI; US

‘all

und herbarium stamp is 31 mm diameter.

128 Austrobaileya 3(1): 1989

Distribution: G. sy/vestre occurs in the Indian subcontinent, parts of China and southeast
Asia.

Notes: Tsiang & Li (1976) list a long series of synonyms for G. sylvestre, however it has
not been possible to see the types of all of these and hence they are not given above.

Huber (1983) gave the type of G. sylvestre as being “Koenig s.n. (LD not seen)’.
The Koenig herbarium consists of numbered specimens. There are two sheets numbered
6733, one of which is referable to the original (albeit brief) description of G. sy/vestre
as Periploca sylvestris (Retzius 1781) (Fig. 8) and hence is chosen lectotype of that name.
The other sheet is from a species of Cynanchum and may represent the type material
of Periploca tunicata Retz.

Gymnema pleiadenium F. Muell., Fragm. 11: 77-78 (1878). Type: East Australia, 1848,
W. Hill (holo: MEL(MEL 1515481)).
Bailey, Queensl. fl. 3: 104 (1900).

Gymnema dunnit (Maiden & Betche) P. Forster, Austrobaileya 2: 401 (1987),
Marsdenia rostrata var. dunnii Maiden & Betche, Proc. Linn. Soc. New South
Wales 31: 736 (1906), synon. nov. Type: New South Wales, Acacia Creek,
McPherson Range, December 1905, J.L. Dunn (holo: NSW; iso: BRI(AQ333081)).

The species is described in detail in Forster (1987).

Additional specimens examined: Western Australia. Devil’s Pass, Napier Range, May 1905, Fitzgerald 595
(BRI,PERTH); Hann & Barnett Rivers, Jun 1905, Fitzgerald 1122 (PERTH); Champagny Is, Bonaparte Archipelago,
May 1972, Wilson 10946 (PERTH); Under cliff of Mt Trafalgar, St George Basin, 15°17’S, 125°04’E, Jan 1989,
Kenneally 10918 & Hyland (BRI,LPERTH). Queensland. Cook District: Forty Mile Scrub, 18°07’S, 144°49/E,
Feb 1972, Hyland 5870 (QRS); ditto, Apr 1977, Gray 450 (QRS); ditto, (cult. Atherton) Nov 1981, Barbagallo
14 (QR*) (cult. Tolga) Apr 1988, Sankowsky [AQ408489] (BRI*); Jervoise Holding, 18°54’S, 144°43’E, May 1979,
Hyland 9921 (QRS). LEICHHARDT DistRicr: Gracemere, Dec 1869, O’Shanesy 1204 (MEL); Glenmoral Gap, 13
km WSW of Theodore, 24°58’S, 149°57’E, Jan 1987, Forster 2858 (BRI); Expedition Range, Rolleston Road,
24°37'S, 149°02’E, Mar 1988, Forster 3606 (BRI). WIDE Bay District: Mt Glastonbury, 26°14’S, 152°29’E, Jan
BRS Forster 2873 & Sharpe (BRI); St Agnes Ck, Walla Stn, 25°09’S, 151°56’S, Nov 1987, Forster 3267 & Zillman
RI).

Distribution: In addition to that cited in Forster (1987), material has been collected or
noted for three further pastoral districts in Queensland and the Kimberley region in
Western Australia. G. pleiadenium occurs in the MCPH, CYRK, BURD, DWSWN and
KIMB regions of Barlow (1985) (Map 3).

Notes: F.M. Bailey in a letter to von Mueller attached to the type of G. pleiadenium
mentions that Hill “thought he got it from the Pine River’. This would be the Pine
River east of Brisbane, as Hill’s collecting activities during this period were apparently
restricted to south-east Queensland.

The populations from northern Australia (illustrated by Williams (1988: 198)) and
Western Australia show some slight differences to those from southern Queensland and
northern New South Wales described by Forster (1987). These are the somewhat larger
leaves, leaf base nearly always cordate, seed pods 6-7 cm long, pedicels up to 7 mm
long, slit between anther wings ca 0.75 mm long and pollinia 0.17-0.2 mm long (Fig.
9). The generally larger leaves of the northern populations are to some extent a result
of phenotypic expresssion, as material from the Forty Mile Scrub (Gray 450) which was
subsequently cultivated at Atherton (Barbagallo 14) produced smaller leaves similar to
material from the southern part of the range in Australia. By comparison material from
the same locality cultivated at Tolga (Sankowsky AQ408489) did not differ from the
field collected material.

Gymnema geminatum R. Br., Prod. 462 (1810). Type: Islands of the Gulf of Carpentaria,
December 1802, R. Brown (lecto (here designated): BM; isolecto: MEL(MEL
113347), BRI(AQ333083)).

Sprengel, Syst. Veg. 1: 844 (1825); Decne in DC., Prod. 8: 623 (1844).

[Gymnema sylvestre auct. non (Retz.) R. Br. ex Schultes: Benth., Fl. austral. 4: 342
(1869); Bailey, Queensl. fl. 3: 103-104 (1900).]

Woody vine, perennial, with white latex. Stems twining, to 2 cm thick, with corky grey-
brown bark; internode length variable to 6 cm. Leaves petiolate; lamina ovate to ovate-

me MP mm Hy Mw MWh BB PW Ma Pw a aw yeh a nny a yt My Bay ty a gt ey a I a

129

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Forster, Asclepradaceae, |

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ALAR Sd PUPS eth bE RRO TEEER PEPio no a atettht TL aes ee

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==

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populations, and longer

dary venation X 0.5. C. fruit X
, Forster 3958. Del. L. Jessup.

*

ing prominent secon
C-D

*
+

howl

. B, Forster 3955

¥

larger more cordate leaves of northern
ir $

ing

B. leaf pa

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+

habit show
feminatuim

Pld}?

: pleiaden
fruits. X 0.5. B-D. Gywinema

*

.9. A. Gypninema

Fig
0

130 Austrobaileya 3(1): 1989

elliptic, up to 90 mm long and 50 mm wide, base rounded, apex acuminate, pinnately
veined, secondary venation prominent, upper surface green to grey-green, paler to yellow-
green beneath, finely pubescent with uniseriate hairs; petiole 7-26 mm long, | mm wide;
extrafloral nectaries 3 or 4 at lamina base. Flowers borne at upper 2-10 nodes. Cymes
umbelliform, appearing at the nodes between the pair of leaves, 1-many-flowered, often
2 per node; peduncle 3-8 mm long, c. | mm wide, finely pubescent, pale green. Pedicels
2-5 mm long, c. 0.5 mm diameter, finely pubescent. Flowers cream, campanulate, c. 4-
5 mm long, 2.5 mm diameter, tube up to 2 mm long. Sepals 5, distinct, ovate to elliptic,
lobed from 1/2 to 3/4 length, externally finely pubescent, generally with 2-4 small glands
at base on the adaxial side. Petals elliptic to oblong, externally glabrous, internally shortly
pubescent, semi-reflexed at anthesis, c. 1.5 mm long, | mm wide. Corolline corona of 5
thickened ridges running down corolla tube and alternating with corolla lobes, densely
pubescent on edges; free ends of lobes erect, linear-subulate, arching over staminal
column, 0.4-0.75 mm long. Staminal column c. 2 mm long, | mm diameter. Anther
appendages acute, c. 0.55 mm long, 0.5 mm wide. Slit between anther wings c. 0.75 mm
long. Style-head conical, c. 1 mm longer than staminal column, c. | mm diameter,
cream. Pollinarium 0.2 mm long, 0.4 mm wide; pollinia oblong, held erect, 0.12 mm
long, 0.07 mm wide, golden; corpusculum oblong, 0.14-0.15 mm long, 0.05-0.06 mm
wide, dark brown; caudicles attached to base of corpusculum, 0.15-0.16 mm long, 0.07
mm wide at point of attachment to pollinium, 0.02 mm wide at point of attachment to
corpusculum. Ovaries free, glabrous, c. 0.75 mm long, 0.75 mm wide. Follicles fusiform
to narrowly ovoid, shortly pubescent, 2.5-5 cm long, 4-13 mm wide. Seed oblong, tan,
5-7 mm long, 2-3 mm wide; coma white, attached to micropylar end of seed, 10-20
mm long. Fig. 9.

Selected specimens. Western Australia. Kununurra District, {5°18’S, 128°45’E, 1978, Gowland 613 (DNA); Goose
Hill, E. Kimberley, Sep 1906, Fitzgerald 1602 (BRI,LNSW,PERTH); Boongaree Is, Prince Frederick Harbour,
15°55’S, 125°10’E, Jul 1973, Wilson 11373 (PERTH); Cape Leveque, 16°25/S, 122°57’E, Apr 1988, Dunlop 7828
(BRI,DNA,NSW,PERTH). Northern Territory. near Prawn farm, off Channel Is Road, 12°29%S, 130°58’E, Dec
1987, Cox [AQ408477] (BRI*); East Point, Darwin, Jan 1988, Stobo [AQ408490] (BRI*); Port Darwin, 1883,
Holtze 300 (MEL); Humpty Doo, Dec 1961, McKee 8322 (BRI), Liverpool River Mouth, Tudor ?t, L1°S9’S,
134°15’E, Jun 1987, Russell-Smith 2558 (DNA); Groote Eylandt, Umbakumba, 4 mile Jungle, 13°52’S, 130°47E,
Jul 1987, Russell-Smith 2868 & Lucas (DNA); Murganella area, 11°26’S, 132°58’E, Feb 1984, Jones 1389 (DNA);
Brogden Pt, Murganella area, 11°31’S, 133°04’E, Feb 1984, Jones 1371 (DNA*). Queensland. Cook DISTRICT:
Mapoon, May 1901, Bailey [AQ216534] (BRI); Z North Fauna Survey Site 17, Weipa, 12°39’S, 141°54’E, 1982,
Morton [AQ378134] (BRI); Few miles SE of Chillagoe, 17°10'S, 144°35’E, Jan 1972, Hyland 5828 (QRS); 17.5
km from Almaden on Chillagoe Rd, 17°16’S, 144°34’E, Mar 1988, Forster 3955 (BRI,CANB), 1.5 km past
Mungana trucking yards, 17°06’S, 144°23’E, Mar 1988, Forster 3958 (BRI,CANB). BURKE DISTRICT: Karumba, '
Aug 1943, Blake 15135 (BRI).

Distribution and habitat: Apparently restricted to Australia in the KIMB, ARNM, CARP
and CYRK regions of Barlow (1985) (Map 4). G. geminatum occurs in deciduous
microphyll vine thickets on a variety of substrate types.

Phenology: Flowering throughout the year with fruiting occurring two to three months
after flowering.

Conservation status: The species is widespread and not vulnerable, endangered or rare
at present.

BRACHYSTELMA Sims

An account of Brachystelma Sims in Australia and New Guinea has been given
by Forster (1988). As noted by P.G. Wilson (in litt.), the prior name to be used for the
single species recognised, is B. glabriflorum (F. Muell.) Schltr. and not B. microstemma
Schltr. which is a nom. nov. based on Microstemma tuberosum R. Br.

Acknowledgements

Illustrations were provided by K. Harold, L.G. Jessup, and A. Thongpukdee. Plant
material was provided by C. Cox, B.P.M. Hyland (QRS), K.F. Kenneally (PERTH), G.
Sankowsky, P.R. Sharpe, J. Stobo and R. Tingey. Collections were made on trips with
L.H. Bird, M.P. Bolton, P.D. Bostock, D.J. Liddle, R.A. Harvey, G. Kenning, M.C.
Tucker, P.R. Sharpe and E.E. Zillman. The Queensland Department of Forestry issued
various permits to collect on land administered by that department. G.P. Guymer (BRI)
and K.L. Wilson (NSW) while Australian Botanical Liasion Officers at Kew checked for

be NADAS ALN mm pm et mm ae IA a ah NANPA mmm? 9 yt NN 4 mp IAW IIA IANA ” Ll ” ” om jm yaa ” sao ig yo he ANANSI NL MMM my minnie ay Pian) yy enya Pymyly ame ae ay my," nnn wl ea ae VSI mmm i lh lt wl HAWN Ne Hy WMH mH my mmm
EIEN IE HWE My Ha ahh Physi mt ng mt hy me hn mmm? monger 1 HH Why ey yh yh i ‘. a tanh oy ae Po eat am one ol rn ‘ cel ea ltetuachsaaht wis i

131

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@ Gymnema

gem cere
mse 4 <i
oy Te tte

*

1 | |

bo |

Od
wt
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co =
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=
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=
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esculentum
geminanun.

-4. Distribution in Australia mapped at 1:250 000

Forster, Asclepiadaceae, |

Maps 1

132 Austrobaileya 3(1): 1989

type material of several species and arranged for photographs to be taken. The latter
provided detailed notes on the type sheets of F inlaysonia obovata. P.G. Wilson (PERTH)

brought to my attention the priority problem in Brachystelma. The Queensland Her-
barium provided working space and processed loan material. The Directors of BRI,
CANB, CBG, DNA, JCT, LD, MEL, PERTH and QRS provided access to material
either at their institutions or on loan. The Australian Biological Resources Study provided
partial financial support in 1988. E.M. Ross (BRI) provided constructive comments on
the manuscript. All are gratefully acknowledged.

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of unpublished East Indian Plants. 2: 48-50. London: Treuttel & Wutz, Treuttel Jun., & Richter.

WILLIAMS, J.B. (1984). Asclepiadaceae. In N. Beadle (ed.). Student’s Flora of North Eastern New South Wales.
Part V. Armidale: University of New England.

WILLIAMS, K.A.W. (1988). Native Plants of Queensland. 3: 198. Ipswich: K.A.W. Williams.
Accepted for publication 10 March 1989

Austrobaileya 3(1): 135-139 (1989) 135

NOTES ON TRACHYMENE RUDGE (APIACEAE)
IN QUEENSLAND, 1

A.FE. Holland

Queensland Herbarium, Meiers Road, Indooroopilly, Qld 4068

Summary

Trachymene clivicola Boyland & Holland is described and 7. procumbens var. hookert (Domin) B.L. Burtt is raised
to specific rank as 7. hookeri based on Didiscus procumbens var. hookeri Domin. T. cussonit (Montr.) B.L. Burtt
is recorded for Australia and described.

Trachymene clivicola Boyland & Holland, species nova, T. g/aucifoliae (F. Muell.) Benth.
affinis a qua mericarpis alatis (ala 1-2 mm lata) et habitu caespitoso effuso
imprimis differt. Typus: Queensland. GREGORY NORTH DISTRICT: Kurran
homestead, ca 160 km WNW of Jundah, 24°09’S, 141°34’E, 5 June 1974, D.E.
Boyland 7060 (holo: BRI; iso: BRI).

T. clivicola Boyland in J. Jessop, ed., Fl. Central Australia 269-270 (1981), nom.
inval.

Semi-erect, tufted, spreading herb. Stems several, much branched, to 40 cm high, glabrous.
Leaves mostly cauline, alternate or opposite, petiolate; petioles to 4 cm long, glabrous
or with a few hairs at base: blades broadly ovate in outline, deeply tripartite, up to 3
cm long, 4 cm wide, glabrous or with scattered hairs below; lobes cuneate at base,
obovate or obtrullate, simple or further shallowly lobed or trisected, ultimate segments
2-5 mm wide. Inflorescences numerous; peduncles up to 90 mm long, 0.5—-1 mm wide,
glabrous; umbels 40-60-flowered, 10-15 mm wide in flower and 15-20 mm wide in
fruit: bracts 12-15, narrow, subulate, 5-8 mm long, 0.2-0.5 mm wide at base, glabrous;
pedicels slender, 4-6 mm long. Flowers blue or white; petals ovate or elliptic, 1-1.5 mm
long, I1-1.2 mm wide; styles 1.5-2 mm long. Fruit a single mericarp;, mericarps obliquely
rounded, laterally flattened, 5-8 mm long, 4-5 mm wide including wing, glabrous,
minutely papillose; wing !-2 mm wide, glabrous, smooth. Fig. 2A-B.

Specimens examined (both BRI): Queensland. GREGORY Nort District: Elderslie, W of Winton, on upper
parts of Mt Booka Booka, Oct 1935, Blake 10057; Warlus 6, site 546, Mesa, 8km N of “Warra’’, Jun 1978, Purdie
| 250.

Distribution and habitat: Far western Queensland in sparse forbland on mesa slopes in
shallow lithosol soils.

Affinities: This species is most closely related to 7. g/aucifolia (F. Muell.) Benth. but
differs as follows:

Erect, sparsely branched herbs 30-100 cm high; mericarps wingless
CIT CISIAVEA) woth ok EN Seah oo nn ee Fie ate le we oes tol lk case ae A T. glaucifolia
Semi-erect, spreading, much branched herbs, to 40 cm high; mericarps with

wings 1-2 mm wide (Gn) ........ .. 0-02 2 ee ee. T. clivicola

Note: The name J. clivicola had not been validly published when used by Boyland and
Stanley in Flora of Central Australia (Boyland & Stanley, 1981). I have taken the
opportunity to formally name the species here.

Trachymene hookeri (Domin) Holland, comb. nov.

Didiscus procumbens var. hookeri Domin, Sitzungsber. Konigl. Bohm. Ges. Wiss.
Prag. 10: 55 (1908); Trachymene procumbens var. hookeri (Domin) Bailey, Compr.
cat. Queensl. pl. 228 (1913). Type: Cook District: Lizard Island, August 1848,
J. MacGillivray s.n. (lecto (here designated): K(2 sheets!, photo BRI')).

Erect perennial herb to 2 m high with a fleshy primary root. Stems one to several,
mostly glabrous, often with sparse spreading hairs at base. Lower leaves alternate, petioles

136 Austrobaileya 3(1): 1989

3-10 cm long, glabrous or sparsely hairy; blades deeply palmatisect, 3- or 5-lobed, 4-10
cm long; lobes deeply pinnatisect, often bipinnatisect, ultimate segments linear or
lanceolate, acute or acuminate, 1-4 mm wide, glabrous or with scattered hairs below:
upper leaves opposite, sessile or shortly petiolate, simple or deeply trisected; lobes linear,
acute, 1-2 mm wide, glabrous. Inflorescences few or many; peduncles 4-18 cm long,
0.5~1 mm wide, glandular hairy at base, otherwise glabrous; umbels 20-45-flowered, 8-
12 mm wide in flower, 10-16 mm wide in fruit; bracts 13-18, narrow, subulate, 5-10
mm long, 0.2-0.5 mm wide at base, glabrous or with spreading hairs; pedicels slender,
3-6 mm long. Flowers white; petals ovate, obtuse, 1-1.5 mm long, 0.5-1 mm wide;
styles 1-1.5 mm long. Fruit of paired mericarps; mericarps obliquely rounded, laterally
flattened, 2-3 mm long, 1.5-2 mm wide, wingless, glabrous, papillose or smooth. Fig.
1.

Specimens examined (all BRI): Queensiand. Cook District: on turnoff to Ebagoola, Sep 1983, Scarth-Johnson
1364; Kennedy Rd., 15 miles [24 km] N of Musgrave, Aug 1965, Gittins 992; 25 km NW of Starke Homestead,
Jul 1980, Clarkson 3278; Telegraph Line, between Saltwater Ck and Morehead River, Aug 1929, Brass 19979:
Isabella falls, North of Cooktown, Oct 1986, Champion 174; 41.6 km WNW of Cooktown, without date, Pedley
2634, [4 miles [22.4 km] NW of Cooktown, Jun 1968, Ped/ey 2627; Emu Swamp, 8.5 km west of the North
Kennedy River on the Road to Kimba Station, May 1982, Clarkson 4285*; 11 km from the Kennedy River
Crossing on the Fairview-Kimba Road on the track to King River outstation, Apr 1980, Clarkson 3216*. NORTH
KENNEDY District: Hinchinbrook I., Ramsay Bay, Aug 1975, Sharpe 1772. (Specimens marked * represent a
more robust form with thick stems woody at base).

\¢
fx cs
ik, &

Fig. 1. Trachymene hookeri: A. habit X 0.5. B. fruit * 6. C. flower X 6.

Holland, Trachymene, | 137

Distribution and habitat: North-eastern Australia from Ebagoola, just south of Coen, to
Hinchinbrook I. It occurs in sandy soil in Eucalyptus woodlands.

Affinities: Trachymene hookeri was previously included in 7. procumbens (F. Muell.)
Benth. but is specifically distinct from that species. The two species can be distinguished
as follows.

Prostrate, multistemmed herbs with soft, glabrous stems; leaf blades

20-50 mm fone; involucral bracts 10-14, 2-5 mm long (Bn, Dd,Le,

Mo,Wb) . .. L. procumbens
Erect herbs with one to several glabrous or - sparsely hairy stems: leaf

blades 40-100 mm ba involucral bracts 13-18, 6-10 mm long

(Co,Nk) . ay T. hookeri

Note: As Domin failed to indicate the location of his type and possibly saw sheets of
Macpgillivray’s Voyage of Rattlesnake Bot. N. 384 in more than one herbarium,
MacGillivray’s material (two sheets) at K is selected lectotype of Domin’s Didiscus
procumbens var. hookeri.

Trachymene cussonii (Montr.) B.L. Burtt, J. Bot. 79: 45 (1941); Hydrocotyle cussoni
Montr., Mem. Acad. Lyons, 10: 211 (1860); Didiscus cussoni (Montr.) Guillaumin
& Beauvisage, Ann. Soc. Bot. Lyons 38: 94 (1913, repr. 1914). Type: De Pile Art
(pers de la Nouvelle Caledonie) (n.¥.).

Trachymene homei Seem., J. Bot. 2: 75 (1864); Didiscus homei (Seem.) Domin,
Sitzungsber. Konig]. Bohm. Ges. Wiss. Prag. 10: 56 (1908). Type: Isle of Pines,
New Caledonia, Sir E. Home (holo: BM n.v., photo BRIE).

Didiscus austrocaledonicus Brongn. & Gris, Ann. Sci. Nat. ser. 5, 3: 235 (1865),
Bull. Soc. Bor. France 12: 272; Trachymene austrocaledonica (Brongn. & Gris)
F. Muell., Phytogr. New Hebr. 13 (1873), J. Bot. 11: 340 (Nov 1873). Syntypes:
New Caledonia: “prope Balade (Vieillard no. 610), et insula Pinorum (Pancher)”

(n.¥.).

Tussock-forming herbaceous perennial to 30 cm high. Stems ascending, robust, glabrous.
Leaves numerous, mostly crowded in a basal rosette; basal leaves long petiolate; petioles
2-12 cm long, glabrous, dilated at base; blades deeply trisected, 2~5 cm long, glabrous,
lobes further deeply bi- or trisected, often further divided, ultimate segments linear,
oblong, acute or obtuse, 2-4 mm wide; cauline leaves similar, alternate or opposite,
sessile or shortly petiolate. Inflorescences numerous; peduncles 2-11 cm long, 1-2.5 mm
wide, glabrous; umbels with 40-100 flowers, 8-13 mm wide in flower, 15~25 mm wide
in fruit; bracts 12-18, narrowly triangular, acuminate, 3~5 mm long, 0.5-1 mm wide at
base, glabrous. Flowers white; petals ovate or elliptic, 1-1.5 mm long, 0.5-1 mm wide;
styles 0.2-0.4 mm long. Fruits of paired mericarps; mericarps obliquely rounded, somewhat
flattened, 4-5 mm long, 3-4 mm wide, wingless, glabrous, entirely smooth. Fig. 2C—D.

Specimens examined (all BRI): Queensland. Port Curtis District: Wreck Island, Apr 1986, Osmund [AQ440690];
One Tree Island, Apr 1986, Beinssen [AQ440693]; Hoskyn Island, Apr 1986, Beinssen [AQ440692]; Fairfax Island,
Apr 1986, Upton [AQ440691]; Lady Elliot Island, May 1969, Firth {AQ8059].

Distribution and habitat: This species 1s a native of New Caledonia and Vanuatu. In
Australia it is at present known only from beaches on the Islands of the Capricornia
and Bunker Groups in the Capricornia Section of the Great Barrier Reef.

Affinities: This species is distinguished from all other Queensland species by its tussock-
forming habit and large (4-5 mm long), completely smooth mericarps.

Acknowledgements

I extend my thanks to Mrs I. Champion and Dr M. Chaloupka and associates
for their collections, the Director of K for the loan of type material and Dr Hy Eichler,
Mr R. Henderson and Miss S. Reynolds for their helpful comments. Mr W. Smith
produced the line drawings.

138 Austrobaileya 3(1): 1989

i ">
psstias
PN ee

ip 35
ve
ba

Fig. 2. Trachymene clivicola: A. habit X 0.5. B. fruit 6. Trachymene cussonii: C. fruit X 6. D. branch with
mature umbel X 0.5.

Holland, Trachymene, | 139

Reference

BOYLAND, D.E. & STANLEY, T.D. (1981} Umbelliferae. In J. Jessop (ed.), Flora of Central Australia 266-271.
Sydney: A.H. & A.W. Reed Pty. Ltd.

Accepted for publication 26 April 1989

Austrobaileya 3(1): 141-148 (1989) 14]

NEW SPECIES OF BULBOPHYLLUM
SECTION OXYSEPALUM (ORCHIDACEAE) IN AUSTRALIA

B. Gray
CSIRO Division of Plant Industry, PO Box 780, Atherton, Qid 4883
and D.L. Jones

Australian National Botanic Gardens
GPO Box 1777, Canberra, ACT 2601

Summary

Bulbophyllum grandimesense, B. lewisense and B. windsorense, all members of section Oxysepalum from north-
eastern Queensland, are described as new and their relationships with previously known species of the section are
discussed. A key to Australian species in section Oxysepalum is included.

Introduction

As outlined by Dockrill (1969), the section Oxysepalum Schltr. in the genus
Bulbophyllum Thouars in Australia contains two species, viz B. gadgarrense Rupp and
B. wadsworthii Dockr. Field studies by the senior author have revealed three further
species which are here described as new. Two species are closely related to B. wadsworthil,
the other to B. gadgarrense.

‘Taxonomy

Bulbophyllum grandimesense B. Gray & D. Jones species nova affinis B. wadsworthii
Dockr. sed foliis angustioribus et petiolatis, floribus majoribus concoloribus
eburneis ad albis differt. Typus: Queensland. Cook District: T.R. 146, Parish
of Monkhouse, Tableland L.A., 15°43’S, 145°17’E, 800 m, 3 May 1983, B. Gray
3045 (holo: QRS; iso: BRI).

Plant epiphytic, consisting of a few semi-erect to pendulous rarely branched stems to
about 8 cm long; roots wiry, ca 0.5 mm diameter, mainly from basal nodes, some stem-
embracing adventitious roots arising from upper nodes; rhizomes ca | mm diameter,
covered by pale brown to dark brown scarious bracts, 8-25 mm_ between pseudobulbs;
pseudobulbs 4-6 mm X 1.5-2.5 mm, more or less ovoid to cylindrical, green, sulcate
on underside, decurrent. Leaf lamina 14-50 mm X 3-5 mm, linear to linear-oblong,
more or less round in cross-section with a narrow groove on the upper surface, on
petioles 4-5 mm long. Pedicels ca 1.5 mm long, bracteate; flowers ca 8 mm long, solitary
at nodes, white throughout, sepals spreading to about 7 mm apart at tips; sepals subsimular,
8-9 mm X 2-2.5 mm, narrowly triangular, constricted in distal third, apices long-
acuminate to filiform: petals ca 2.5 mm X ca 1.5 mm, more or less ovate, narrowing at
base, slightly incurved; labellum ca 2.3 mm X ca 1.3 mm, fleshy, distinctly 3-lobed,
erect in proximal half, then curving in a semi-circle, surface smooth, lateral lobes ca lL.
mm long, erect, mid-lobe obtuse; column ca 1.2 mm long; stelidia ca 0.6 mm long, very
short, broadly triangular, blunt to rounded; column foot ca 2.2 mm long, curved; anther
cap ca 0.5 mm X ca 0.5 mm, with a prominently deflexed rostrum ca 0.13 mm long.
Capsule not seen. Fig. 1.

Specimens examined (ail QRS): Queensland. Cook District: T.R. 146, Parish of Monkhouse, Tableland L.A.
15°43’S. 145°17’E, 800 m, May 1983, Gray 3045; T.R. 146, Parish of Monkhouse, Big Tableland, 15°42’S, 145°16’E,
500 m, Aug 1984, Gray 3524.

Distribution and habitat: At present this species is known only from the Big Tableland
on the Trevethan Range north of the Bloomfield River. It grows in highland rainforest
between 500-800 m altitude. Plants favour the small, upper branches of trees in the
rainforest canopy.

Flowering Period: May—June.

142 Austrobaileya 3(1}; 1989

Notes: Bulbophyllum grandimesense has some affinities with B. wadsworthii but it can
be readily distinguished by its much narrower, petiolate leaves and the larger, uniformly
coloured cream to white flowers. The flowers of B. wadsworthii are also cream to white
but have a brown to orange-brown labellum. B. grandimesense is the only member of
the section Oxysepalum known to grow on the Big Tableland.

Conservation Status: Bulbophyllum grandimesense is a poorly collected species and from
the collections available appears to be restricted to a fairly limited, highland habitat. Its
conservation status is 2RC according to Leigh, Briggs and Hartley (1981).

Etymology: The specific epithet is derived from the Latin grandis, big, and the Spanish
mesa, tableland, a reference to the distribution of this species on the Big Tableland.

amm

Fig. 1. Bulbophyllum grandimesense. A. habit. B. flower from front. C. flower from side. D. column and labellum
from side. E. column from front. F. column and labellum from front. G. pollinia from front. H. pollinia from
side. 1. anther from front. J. anther from side. All from Gray 3045.

Gray & Jones, Bulbophyllum 143

Bulbophyllum lewisense B. Gray & D. Jones species nova affinis B. gadgarrensi Rupp
sed plantis exilioribus, pseudobulbis conspicuis nudis, floribus albis differt. Typus:
Queensland. Cook District: S.F.R. 143, South Mary L.A. 16°30/S, 145° 15’E,
1000 m, 13 October 1979, B. Gray 1539 (holo: QRS; iso: BRI,NSW).

Plant epiphytic, consisting of a few semi-pendulous or spreading, rarely branched stems
to about 15 cm long; roots wiry, ca 0.5 mm diameter, mainly from basal nodes, some
adventitious stem-embracing roots arising from upper nodes; rhizomes ca | mm diameter,
covered by pale brown, scarious bracts, 5-10 mm between pseudobulbs; pseudobulbs 4-
5 mm X 3-3.5 mm, more or less ovoid, green, sulcate on underside, decurrent. Leaf
lamina 15-22 mm X 3-5 mm, narrowly oblong to narrowly elliptic, thick and fleshy,
dark green, slightly curved, deeply grooved on upper surface, more or less cordate in
cross section, shortly petiolate. Pedicels ca 3.5 mm long, bracteate, flowers ca 5 mm
long, white throughout, solitary at nodes, sepals spreading to about 4 mm apart at the
tips: sepals subsimilar, 5-6 mm X 1.5-2 mm, narrowly triangular, apices cymbiform,
acuminate: petals 2-2.5 mm X ca 0.8 mm, oblong, slightly spreading; labellum ca 2 mm
x ca 1 mm, fleshy, distinctly three-lobed, erect in proximal half then curving in a seml-
circle, surface smooth, lateral lobes ca 1 mm long, erect, mid-lobe obtuse; column ca
0.8 mm long; stelidia ca 0.4 mm long, narrowly triangular, bluntly pointed; column foot
ca 2 mm long, curved; anther cap ca 0.4 mm X 0.5 mm, with a prominent deflexed
rostrum ca 0.1 mm long. Capsule not seen. Fig. 2.

Specimens examined. (all QRS). Cook District: S.F.R. 143, Parish of Riflemead, North Mary L.A., 16°31'S,
(45°16’E, 1100 m, Aug 1986, Collins s.n., Zarda Roots Creek, Jan 1936, Flecker, Flecker Herbarium 1303; S.F.R.
144, Parish of Whypalla, Bower Bird L.A., 16°L5’S, 145°OV’E, 1240 m, Aug 1984, Gray 3535; S.F.R. 143, South
Mary L.A., 16°30’S, 145°1S’E, 100 m, Oct 1979, Gray 1539: S.F.R. 144, Cockatoo L.A., 16°17’S, 145°05’E, 960
m, Jui 1982, Gray 2671; S.F.R.143, Parish of Riflemead, 16°32'S, 145°1VE, 1100 m, Oct 1984, Gray 3574.

Distribution and habitat: B. /ewisense occurs on Mt Lewis, Carbine Tableland and
Windsor Tableland. It grows on trees in rainforest at medium to high elevations (900-
1200 metres) and seems to favour slender trunks or smal! branches in the upper canopy.

Flowering Period: September and October.

Notes: B. Jewisense has as its closest congener B. gadgarrense but it can be readily
distinguished by the smaller, more open plants, naked, conspicuous pseudobulbs and
white flowers. The two species grow together in some areas but are easily distinguished.
The new species always forms small, sparse clumps and the roots arise mainly from the
base of the stems. This contrasts with B. gadgarrense in which many of the roots arise
from aerial nodes along the stems.

Conservation Status: Sparsely distributed but not rare.

Etymology: The specific epithet refers to the type locality, Mt Lewis.

Bulbophyllum windsorense B. Gray & D., Jones species nova affinis B. wadsworthii Dockr.
sed foliis brevioribus valde canaliculatis, floribus majoribus concoloribus eburneis
ad viridi-eburneis differt. Typus: Queensland. Cook District: S.F.R. 144, Parish
of Whypalla, Bower Bird L.A., 16°14’S, 143°00’E, 1220 m, 15 June 1981, B. Gray
4503 (holo: QRS; iso: BRI,NSW).

Plant epiphytic, consisting of a few to several, upright to pendulous, rarely branched
stems to 30 cm long; roots wiry, ca 0.5-0.8 mm diameter, arising from basal nodes or
base of branches: rhizomes 1-1.5 mm diameter, covered by brown scarious bracts, 8—-
20 mm between pseudobulbs; pseudobulbs 5-8 mm X 4-5 mm, obovate to almost
cylindrical, decumbent, truncate, only partly obscured by bracts. Leaves 12-30 mm X
4-6 mm, narrowly elliptic to oblong, sessile, acute, thick and fleshy, more or less round
in cross section, broadly and deeply grooved on the upper surface, often compressed
laterally closing the groove. Peduncles ca 4-6 mm long including the ovary, bracteate;
flowers solitary or in pairs from nodes and internodes, cream or greenish cream,
10-13 mm long, sepals spreading 7-16 mm apart at the apices; sepals subsimilar, 9-12
mm X 3.5-4.5 mm, narrowly triangular, constricted in the distal half, apices filiform,
lateral sepals connate for about 2 mm at the base; petals 2-3 mm xX 1.2~1.5 mm, obovate;
labellum 2.2-2.5 mm X 1.6-2 mm when flattened, fleshy, erect in proximal half then
curved in a semicircle, surface smooth, lateral lobes ca 1 mm long, erect, midlobe
recurved, 1-1.2 mm long, more or less obtuse; column I-1.2 mm long; stelidia very

144 Austrobatleya 3(1): 1989

Fig. 2. Bulbophyllum lewisense: A. habit. B. flower from side. C. flower from front. D. column and labellum from

side. E. column from front. F. anther from side. G. anther from front. H. pollinia from front. All from Gray
1539.

Gray & Jones, Bulbophyllum i45

short, oblong; column foot 1.5-1.8 mm long, curved almost in a semicircle with the
column: anther ca 0.35 mm X 0.35 mm, with a deflexed rostrum. Capsule not seen. Fig.
3.

Distribution and habitat: Known only from highland rainforests of the Mt Windsor
Tableland. Common at elevations above 1000 m and growing in small clumps on the
upper branches of trees in the rainforest canopy.

Flowering Period: May—August.

Notes: B. windsorense is probably most closely related to B. wadsworthii, but it can be
readily distinguished by the larger, uniformly coloured flowers and the shorter, deeply
srooved leaves.

Conservation status: Its conservation status is 3RC according to Leigh, Briggs & Hartley
(1981),

Etymology: The specific epithet refers to the type locality, Windsor Tableland.

Discussion

The five Australian species of Bulbophyllum in section Oxysepalum are confined
to north-eastern Queensland where they grow in highland rainforests above 500 m
altitude. All species grow on trees with B. wadsworthii and B. gadgarrense sometimes
also found on boulders. Field observations indicate that the newly described species are
relatively uncommon to rare and they tend to occupy different ecological niches. Further
exploration of isolated highland regions in north-eastern Queensland could result in the
discovery of new taxa in this group. Members of this section can be identified from
non-flowering material and an illustration of cross-sections and outlines of leaves is
included (Fig. 4), as these provide valuable diagnostic characters.

Key to Australian species of Bulbophyllum
section Oxysepalum

[.. eaves PetOlate a Ge yy eee wh del eas be Ge So ate tee om ee eS Me a he ee 2
aes SESS 88. nena cll ot Tel ae hates fy ov Dl oY eg be See ne on ete at ges 4

2. Leaf lamina narrowly grooved (more or less round in cross-section);

petioles 4-6 mm long; pseudobulbs 4-7 mm long, 2-3 mm broad,

conspicuous; flowers white or creamy white; sepals long acumin-
Ate 0 cc ee ce ee ee se ee ee sete vevs.aee..... 3B. grandimesense
Leaf lamina broadly grooved or channelled in cross-section ............ 3

3. Leaf lamina very thick and fleshy, nearly oblong, 15-25 mm long, 3-4

mm broad: petioles 1-3 mm long; pseudobulbs 2-3 mm long, I-1.2

mm broad, almost concealed by stem bracts; flowers yellow or orange;
sepals narrowly triangular, obtuse .................... B. gadgarrense
Leaf lamina thin, narrowly elliptic 20-45 mm long, 3-6 mm broad;
pseudobulbs 4-5 mm long, 3-3.5 mm broad, conspicuous; flowers
white or creamy white; sepals broadly triangular, acute or shortly
BORTHINATES eri Gee atv lee wee eA gens! Ca od. aie et athe th omy cts B. lewisense

4. Leaf lamina narrowly grooved (nearly round in cross-section), 20-80 mm
long; flowers white, creamy white or pale green; labellum orange or
orange brown; sepals long acuminate eee ee Pe B. wadsworthii

Leaves broadly grooved in cross-section .. .. 2. 6. ee ee ee ee ee ee 5

5. Leaf lamina short, thick and fleshy, nearly oblong, 15-30 mm long, 4-6

mm broad; flowers pale yellow or greenish yellow; sepals long
acuminate .... 2... ee ee ee ee ee ee ee te ee ea es ee... B. windsorense

Leaf lamina thin, narrowly elliptic 20-45 mm long, 3-6 mm broad; flowers
white or creamy white; sepals acute or shortly acuminate ...... B. lewisense

146 Austrobaileya 3(1): 1989

Fig. 3. Bulbophyllum windsorense. A. habit. B. flower from front. C. flower from side. D. column and labellum
from side. E. column from front. F. labellum from front. G. anther from side. H. anther from front. 1. pollinia
from front. All from Gray 4503.

147

Gray & Jones, Bulbophyllum

Fig. 4. Leaf, pseudobulb and bract habit of Bulbophyllum sect. Oxysepalum. 1. B, grandimesense. 2. B. windorense.
3B. wadsworthii. 4. B. lewisense. 5. B. gadgarrense.

148 Austrobaileya 3(1): 1989

Acknowledgements

We wish to thank Lyn Craven, CSIRO, for the Latin translations, Reg Lockyer
and Ron Collins for supplying live plants for study, and Barbara Jones for typing the
manuscript. Jim Armstrong and Mark Clements, both of the Australian National Botanic
Gardens, read the draft manuscript. Bob Johnson, Director, Queensland Herbarium:
provided access to material at BRI.

References
DOCKRILL (1969). Australian Indigenous Orchids, Vol. [. Sydney: Society for Growing Australian Plants.

LEIGH, J., BRIGGS, J. & HARTLEY, W. (1981). Rare or Threatened Australian Plants. Australian National
Parks and Wildlife Service Special Publication No. 7. Canberra: Australian National Parks and Wildlife
Service.

Accepted for publication 17 April 1989

Austrobaileya 3(1): 149-158 (1989) 149

REVISION OF THE AUSTRALIAN VITACEAE,
5. TETRASTIGMA (MIQ.) PLANCHON

B.R. Jackes

Botany Department, James Cook University of North Queensland,
Townsville, Qld 4811

Summary

Tetrastigma nitens (F. Muell.) Planchon, previously the only known species of Tetrastigma recorded for Australia,
is redescribed. Three new species T. crenatum, T. petraeum and T. thorsborneorum are described and illustrated.

A description of T. pisicarpum is included as specimens from a vine growing in vine forest at the tip of Cape York
possibly belong to this species. A description of the genus is also given, together with a key to the species.

This paper completes the revision of the Australian species of the Vitaceae. The
genera described are Ampelocissus Planchon (Jackes 1984), Cayratia Juss. (Jackes 1987),
Cissus L. (Jackes 1988a) and the Western Australian endemic C/lematicissus (F. Muell.)
rire (Jackes 1989). A key to the genera in Australia has been given by Jackes
(1988b).

TETRASTIGMA

Tetrastigma (Miq.) Planchon in A. & C. DC., Monogr. phan. 5: 423 (1887); Vitis sect.
Tetrastigma Miq., Ann. mus. lugd.-bat. 1: 72 (1863). Type: Vitis lanceolaria Roxb.
(= Tetrastigma lanceolarium (Roxb.) Planchon.)

Climbing evergreen vines, dioecious. Stems striate, initially rounded but often becoming
flattened with age; underground root system extensive. Tendrils leaf-opposed, unbranched
or branched, branches subtended by a bract. Leaves alternate, trifoliolate or pedately 5-
{1-foliolate, margins indented rarely entire; indentations usually small. Stipules 2, cad-
ucous. Inflorescence in Australian species axillary, shortly pedunculate, cymose with
terminal multiflowered umbels: peduncles and pedicels subtended by bracts. Flowers
unisexual, tetramerous. Calyx cup-shaped; margin entire or shallowly lobed. Corolla with
4 free lobes: lobes cohering in bud, cucullate at apex, usually reflexed after anthesis,
soon caducous. Stamens inserted on the receptacle at base of disc opposite petals;
filaments erect: anthers in male flowers ovate-triangular, dorsifixed, introrse, opening by
longitudinal slits; anthers in female flowers on staminodia, much reduced. Pollen grains
tricolporate. Disc obscure, adnate to and entirely surrounding the base of the ovary in
female flowers, almost completely covering the much reduced non-functional ovary in
male flowers. Fertile ovary 2-locular with 2 anatropous, basally attached ovules per
locule, conical; style short or absent; stigma large, 4-lobed, often ciliolate. Berry fleshy,
globose to elliptical, with 1-4 ovoid to globose seeds. Shape of seeds in median transverse
section varies according to the number maturing in the fruit; perichalaza extends for
two-thirds to almost the total length of the dorsal surface. Endotesta irregularly lignified,
endosperm ruminate, appearing convoluted in median transverse section. Mucilage ceils
and raphide sacs often containing mucilage appear to be present in all organs. Cotyledons
lanceolate, three-veined at base, margins entire.

This genus is readily distinguished from the other genera of the Vitaceae by the
dioecious flowers and the large 4-lobed stigma present in the pistillate flowers, and the
capitate, entire stigma in the staminate flowers. In Australia, non-fertile specimens of
Tetrastigma are usually readily recognised in the field by the firm, palmate or pedately
compound, distinctly petiolate leaves which are glabrous or apparently glabrous but
minutely puberulent when examined under magnification.

There appears to be little intraspecific variation with respect to leaf characters.
This is unusual when compared to the other genera of Vitaceae occurring in Australia.
Main characters used for distinguishing between the species are. leaflet number, size of
leaflets, and presence or absence of hairs. All measurements and observations in this
paper refer to dried mature organs except where otherwise indicated.

150 Austrobaileya 3(1): 1989

Zetrastigma occurs chiefly in the Asian tropics and subtropics. It extends from
India north to China, south to Australia and eastward to Fiji; more than half the 120
species which have been described occur in Malesia. Eighteen have been described from
Papua New Guinea but many of these are inadequately known. Australian species fall
into three groups: 7. nitens (F. Muell.) Planchon and T. petraeum sp. nov., which form
a closely related pair with different habitat preferences, the former being found chiefly
in or adjacent to rainforest, whilst the latter occurs in drier habitats: T. crenatum Sp.
noy., which is closely related to the Papua New Guinean species 7. lauterbachianum
Gilg.; and 7. thorsborneorum sp. nov., which is closely related to T. pisicarpum (Miq.}
Planchon, which occurs in southern Papua New Guinea. Vegetative material and
specimens with immature staminate inflorescences collected in the Lockerbie scrub area
at the tip of Cape York possibly belong to this species; however its status cannot be
resolved until mature flowering and fruiting material is collected.

Terms used to describe rainforest habitats are those of Webb (1978),

Key to Species

b, dGaves SelOHOlale ss 1-8 4-6 kk oe cog Lee ee nw vaneec My eles cee ee nt Oe. 2
PEEVES POC ACiees cot: dey ia aha S86. ta covet Deas prey her lal om Ss ba eis 4

2. Leaves pubescent chiefly on lower surface of the midrib; margins crenate-
serrulate, peg-like lenticels present on older stems.......... 1. T. crenatum
Leaves glabrous, margins entire to irregularly serrate, lenticels not peg-like .. 3

3. Central leaflet more than 9 cm long, length/breadth ratio 2.08 +
Central leaflet rarely exceeding 8 cm long, length/breadth ratio 1.45 +
EI Pert chen oo cnt nce, cheb d hte s MR oh Sey ot Sail y ccc here aden alls 3. T. petraeum

2. I. nitens

4. Seed with 2 lateral furrows which converge with central furrow on ventral
surface at base; staminate inflorescence 3-5 cm long; leaflets usually

DIO hag ecnt a ott 4 hag URI BAGS tte tiie twee ml, lee 4. T. thorsborneorum
seed with 2 lateral furrows which converge with central furrow on ventral
surface towards centre of seed; staminate inflorescence usually more

than 6 cm long; leaflets usually 7-11 ................ 5. I. pisicarpum

1. Tetrastigma crenatum B.R. Jackes species nova affinis speciebus novoguineensibus T
lauterbachiano Gilg et T. papilloso (Blume) Planchon autem differt foliis trifo-
liolatis marginibus crenato-serrulatis, pilis uniseriatis multicellularibus in costa
infra et papillis parvis in lamina consociata praeditis, floribus in caulibus vetioribus
portatis et lenticellis prominentibus in caulibus vetis ornatis. Typus: Queensland.
Cook District: Cardstone via Tully, 17°45’S, 145°37’E, 3 April 1984, EM. Jackes,
" rainforest vine. Flowers minute, greenish on old wood. Red loam”. (holo: BRI:
iso: CANB,K,L,NSW).

Slender vine climbing over rainforest trees; stems lenticellate striate, with uniseriate
multicellular hairs present on young stems; lenticels rounded, pustular with margins
forming hollow peg-like protuberances to 3 mm long on older stems. Tendrils bifid.
Leaves trifoliolate; leaflets glabrous above, lower with sparse uniseriate multicellular hairs
confined to midrib and base of main laterals; papillae associated chiefly with the upper
midrib and veins on the lower surface; margins crenate-serrulate: central leaflet broadly
lanceolate, apex acuminate, base shortly angustate, (13.2~)14.8(-18) cm x (6.8—)7.4(-8.5)
cm; laterals lanceolate, slightly oblique at the base, (10.7-)11.9(-14) cm x (3.2~)5.9(-
6.4) cm. Petiole often sparsely pubescent, 6-10 cm long; central petiolule striate with 2
shallow grooves on upper surface, 1.7-2.3 cm long, lateral petiolules similar, 0.6-1 cm
long. Stipules triangular, borne at right angles to the stem, 4-5 mm X 3-4 mm: margins
papillate. Inflorescence borne on older wood; bracts prominent, several often clustered:
uniseriate hairs present on branches. Staminate inflorescence not seen. Pistillate inflo-
rescence with congested terminal umbels 6-7 cm long; pedicels pubescent, c. 1 mm long;
calyx c. 0.25 mm long, with 4 small pubescent lobes; corolla c. 1-1.25 mm long, greenish,

Jackes, 7etrastigma 151

pubescent, with lobes cucullate and extended to form a pouch near the apex, not strongly
reflexed: ovary conical, c. 0.5 mm long; style short, stigmatic lobes narrow. Fruit not
seen. Fig. 1A-B; Fig. 2A-D.

Representative specimens: Queensland. Cook District: Ryker’s Track, Cape Tribulation, Oct 1982, Jackes
(BRIL.CBG,NSW), Mossman Gorge, Jul 1976, Jackes (BRI,CANB,QRS); Babinda, 17°20’S, 145°55’E, May 1978,
Jazo (QRS).

Specimens from 5 localities examined.

Distribution and habitat: This species (Map 1) has been found only between Cape
Tribulation to the north and the Tully area to the south, in complex mesophyll vine
forest between sea level and 200 m altitude.

Phenology: Flowers have been collected in April and May; male flowers and fruits have
not yet been collected.

Notes and observations: This species is related to 7. /auterbachianum, from which it
may be distinguished by the smaller pistillate inflorescence and the leaves which do not
have uniseriate hairs on the abaxial surface, only small papillae being associated with
the midrib, and to 7. papillosum which has glabrous leaves. I could not distinguish
between specimens of 7. /auterbachianum and T. gilgianum, and hence regard these as
synonymous. Flowering is apparently irregular. It took five years of regular visits to two
populations to obtain flowering material! Plants grown from cuttings took two years to
flower in Townsville. Adventitious roots on the cuttings emerged through the lenticular
protuberances.

Fig. 1. Tetrastigma crenatum: A. \eaf x 0.5. B. pistillate inflorescence borne on old wood X I.

Austrobaileya 3(1): 1989

Be

ee

Fig. 2. Tetrastigma crenatum: A. portion of stem showing pustular lenticels. B. buds of female flowers. C. open
female flower. D. ovary with expanded 4-lobed stigma and staminodes. All scale bars equal 0.5 mm.

The specific epithet was chosen because this is the only

| ustralian species
with crenate-serrulate leaf margins.

Planchon in C. DC., Monogr. phan. 5: 427 (1887):
ragm. 2: 73 (1861), 6: 178 (1868). pe: Clarence

E EL 540130),P); Hastings River, Beckler (syn:
540142)); Macleay Beckler (syn: MEL(MI P).

obust vine; mature stems lenticellate, usually flattened, thickened at the nodes, glabrous
lenticels elongated, often with raised margins. Tendrils simple or bifid. Leaves trifoliolate:
leaflets thinly coriaceous, glabrous; margins undulate or slightly revolute with 4-6 shallow
teeth present on both sides of the lamina, often prominent on juvenile leaves, rarely

Vitis nitens
3eckler (syn:

a

*

Jackes, Tetrastigma [53

entire; central leaflet broadly lanceolate, apex acuminate, base shortly angustate, (9-
)11.9(-15) cm X (4.5-)6.3(—7) cm; lateral leaflets lanceolate, slightly oblique at base, (7.5-
)9.3(-12.2) cm X (3.6-)4.8(-7) cm. Petiole shallowly grooved, often swollen at junction
with stem, (1.5-)3.9(—5.8) cm long; central petiolule striate, shallowly grooved above,
1-3.2 cm long; lateral petiolules 0.2-1.8 cm long. Stipules triangular, c. 3 mm X 3 mm,
glabrous. Inflorescence with prominent bracts, and sometimes small uniseriate papillae.
Staminate inflorescence with 1-3 peduncles per axil, 4-6 cm long; pedicels 3-4 mm long;
calyx spreading, c. 0.5 mm long; corolla c. 2 mm X 1 mm; flaments c. 1 mm long;
anthers triangular-ovate, c. 0.5 mm long; ovary much reduced and almost covered by
the disc. Pistillate inflorescence with | peduncle per axil rarely more, 2-3 cm long;
pedicels 1-2 mm long; calyx spreading, c. 0.5 mm long; corolla creamy-green, c. 3 mm
x 1.5 mm: lobes strongly reflexed after anthesis; staminodes present; ovary conical;
stigma sessile, lobes ciliolate, spreading. Berry ovoid to ellipsoid, purplish-black to black
at maturity, 1.2-1.5 mm x 1-1.3 mm; seeds ovoid, [-1.1 cm X 0.6-0.9 cm; | or 2(-4)
= fruit; endosperm in transverse section broken by the convoluted endotesta. Fig. 3A-

Representative specimens: Queensland. Cook District: 0.5 miles {[c. 1 km] NW of Stuckies Gap, Bloomfield
River area, 15°50’S, 143°19’E, May 1969, Webb & Tracey 8936 (BRI); Little Mulgrave Creek crossing near
Gordonvale, May 1979, Jackes (A,BRI,CANB,CBG,JCT,K,L,MEL,NSW,QRS). NORTH KENNEDY DISTRICT: Herv-
ey’s Range W of Townsville, 19°20’S, 146°29’E, Jan 1978, Jackes (L,LPERTH). SOUTH KENNEDY DISTRICT:
Dalrymple Heights and vicinity, Jun-Nov 1947, Clemens (BRI,K). Port Curtis District: Rockhampton,
1863-1865, Dietrich (BM,PR). MORETON DistRICT: Blackall Range, May 1977, Byrnes 3524 (BRD; Mt. Greville,
Jun 1963, Fyerist 7253 (BRI,.K). New South Wales. NortH Coast: Sandiland Ranges, Nov 1904, Boorman
(NSW); Mt. Elaine, c. 12 miles [c. 20 km] SE of Grafton, Nov 1956, Constable (NSW); Manning River, 5 miles
[c. 8.2 km]) NW of Burdook, Dec 1953, Johnson (K,NSW); Cabbage Tree Island, offshore island east of Port
Stephens, Jan 1969, Mfartensz (CANB). NORTHERN TABLELANDS: Platypus Ck., New England National Park, Dec
1960, Williams (NE). CENTRAL Coast: Martinvale, Nov 1965, Blaxell (NSW).

Specimens from 91 localities examined.

Distribution and habitat: This species (Map 2) is found in an area extending from south
of Cooktown to around Sydney, chiefly in vine forest and along moist gullies. It occurs
on a variety of soil types.

Phenology: Flowering occurs mainly in winter but flowers have been collected until
midsummer. Fruits have been collected throughout the year.

Notes and observations: This widespread species shows affinities with 7. /anceolarium,

which occurs in Papua New Guinea, but may be distinguished from it by the broader
leaves and larger fruits. It is closely related to 7. petraeum which has smaller broader
leaves and prefers drier inland habitats.

3. Tetrastigma petraeum B.R. Jackes species nova affinis 7. nitenti (F. Muell.) Planchon
a quo foliis brevioribus latioribusque et fructibus parvioribus distinguitur, et
addite areas siccas [exempli gratia dumetum cum plantis volubilibus (‘vine thicket’)
et silvam apertam] praeopta. Typus: Queensland. NORTH KENNEDY DISTRICT:
Walleroo Hill, 3 miles [c. 4.8 km] SW of Cashmere crossing, Herbert River,
18°08’S, 145°20’E, 30 March 1980, E.M. Jackes, “vine thicket on basaltic outcrop”
(holo: BRI; iso: CANB,L,NSW).

A robust scrambling vine; stems striate to rounded, thickened at nodes, glabrous. Tendrils
simple or shortly bifid. Leaves trifoliolate; leaflets coriaceous, glossy above but may be
yellowish near marginal teeth; both surfaces glabrous; margins with 4-6 small mucronate
teeth usually present, rarely entire; central leaflet broadly elliptical, apex acuminate to
shortly caudate, base often truncate, (6-)6.5(-9) cm X (3.5-)4.5(-6.3) cm; lateral leaflets
broadly elliptical, slightly oblique at base, (5.3-)5.6(-7.5) cm X (3.5-)4(-5.5) cm. Petiole
striate, 1.8-4 cm long; central petiolule striate, 1.5—2 cm long; lateral petiolules 0.3-0.4
cm long. Stipules triangular, carried at right angles to the stem, revolute, c. 3 mm X 2.5
mm, glabrous. Inflorescence bracts inconspicuous. Staminate inflorescence not seen.
Pistillate inflorescence 2-3 cm long, glabrous; pedicels c. 3 mm long; calyx truncate c.
0.5 mm long; corolla creamy-green, 2-3 mm X 1.5 mm; lobes strongly reflexed after
anthesis; staminodes present, aborted anther reduced and awl-shaped; ovary conical, 1.5-
2 mm long: stigma sessile, lobes ciliolate, spreading. Berry ovoid, black at maturity, 0.9-
1.2 cm X 0.7-1 cm: seeds ovoid, 6-7 mm X 3.5 mm; usually 2 per fruit. Endosperm in
transverse section much dissected by the convoluted endotesta. Fig. 3E-H, Fig. 5A-B.

154 Austrobaileya 3(1): 1989

Representative specimens: Queensland. Cook District: Granite Creek, Mareeba, 17°05’S, 145°21’E, Jul 1985,
Jackes (A,BRI,CANB,K,L,MEL,NSW,QRS); Near Lancelot Mine, Silver Valley, Ravenshoe, Jul 1978, Lockyer
(CBG). NorTH KENNEDY District: 3 km W of “Gregory Springs” Station, [19°40’S, 144°22’E, Oct 1984, Jackes
(JCT,MEL); “Lochwall” Station, Charters Towers, Dec 1977, Adarsh [AQ441436] (BRI).

Specimens from 7 localities examined.

Distribution and habitat: This species is found in open forest and vine thickets (Map 2)
where it is often growing over rocks. Soils are variable.

Fig. 3. Tetrastigma nitens: A. leaf with staminate inflorescence X 0.5. B. seed, ventral surface showing the position
of the raphe < 5. C. seed, dorsal surface showing the position of the perichalaza x 5. D. median transverse
section of mature seed X 5. Tetrastigma petracunt. E. leaf with pistillate inflorescence < 0.5. F. seed, ventral
surface showing the position of the raphe X 5. G. seed, dorsal surface showing the position of the perichalaza X
5. H. median transverse section of mature seed. (e — lignified endotesta: s - endosperm.)

Jackes, Tetrastigma 155

Phenology: Flowers have been collected in March and fruits throughout the year.

Notes and observations: This species is easily distinguished from the closely related 7.
nitens by the broadly elliptical almost ovoid leaves, which are also shorter than in that
species, the smaller less pulpy berries and its habitat. Male flowers have not been seen.

Etymology: The specific epithet refers to the species’ habit of growing over rocks exposed
to the sun.

4. Tetrastigma thorsborneorum B.R. Jackes species nova affine 7. pisicarpo (Miq.)
Planchon a quo fructibus grandioribus globulo-ovoideis caesionigris ubi maturis,
seminibus ovoideis, sulcis lateralibus duobus convergentibus in sulcum centralem
a raphe in pagina ventrali prope basem seminis factum ornatis, foliolis plerumque
paucioribus et inflorescentia staminata robustioribus distinguitur, et a speciebus
australianis ceteris foliis pedatis differt. Typus: Queensland. NORTH KENNEDY
DisTRIicT: Gustav Creek, Nelly Bay, Magnetic Island, 19°09’S, 146°51’E, 15 March
1985, B.R. Jackes ‘“‘closed forest, vine, flowers white’? (holo: BRI; iso:
A,CANB,K,L,MEL,NSW).

A robust vine; stems glabrous, lenticellate, rounded initially, becoming flattened with
age: lenticels prominent. Tendrils bifid. Leaves pedate, with 5-7({-11) leaflets; leaflets
firm, dark glossy green; both surfaces glabrous; margins irregularly toothed with 5 or 6
teeth on either side of the lamina; central leaflet elliptical, apex acuminate, base shortly
attenuate, (12-)13(-18) cm X (4.3-)6(-8.5) cm; lateral leaflets elliptical, proportionally
smaller, oblique at the base, particularly the lowest leaflets. Petiole rounded, (5.5—)6.6(-
11) cm long; lIenticels prominent even before leaf is fully expanded; central petiolule
rounded, lenticels often obvious, (1-)2.7(—4.2) cm long; lateral petiolules (0.5-—)1.8(-—3.3)
cm long. Stipules triangular, c. 2.5 mm X 2 mm. Inflorescence bracts inconspicuous;
scattered papillae present on pedicels. Staminate inflorescence robust, 3-5 cm long; calyx
spreading, c. 0.5 mm long, papillose; corolla whitish, c. 2 mm X 1.5 mm, cucullate at
the apex, lobes strongly reflexed; filaments c. 1.5 mm long, anthers c. 0.5 mm long.
Pistillate inflorescence 3.5-5.7 cm long; pedicels 3 mm long; calyx as for male flower;
corolla 2.5 mm X 1.5 mm; aborted anthers triangular; ovary broadly conical, c. 1.5 mm
long; stigma sessile, lobes spreading, ciliolate. Berry globular to ovoid, bluish black to
black at maturity, 0.9-1.2 cm X 0.7-1 cm; seeds ovoid, c. 7 mm X 6 mm; 1 per fruit;
furrows on the ventral surface converge at base of seed; endosperm in transverse section
much dissected by the convoluted endotesta. Fig. 4A—E, Fig. 5C-D.

Representative specimens: Queensland. Cook District: Tozer Gap, Jun 1983, Morton (JCT); Chester River,
13°40’S, 143°25’E, Jul 1977, Hyland 9455 (QRS); Wallaby Creek, S of Cooktown, Jun 1982, Jackes (BRI,CANB).
NortH KENNEDY District: Murray Upper, 18°00’S, 145°00’E, Jun 1977, Cooke (DNA,QRS); Cardwell, Sep 1978,
A. & M. Thorsborne (BRI). SouTH KENNEDY District: Airlie Beach, Nov 1982, Jackes (JCT).

Specimens from 14 localities examined.

Distribution and habitat: This species occurs (Map 1) along the eastern Queensland
coastline from Tozer Gap in Cape York Peninsula as far south as Airlie Beach (20°16’S,
148°44’E) between sea level and 300 m, and is commonly found along streams in vine
forest. Soils are variable.

Phenology: Flowers have been collected in autumn and fruits in winter and early spring.

Notes and observations: This species is easily distinguished by its firm, pedate leaves
with leaflets with irregularly toothed margins, and numerous conspicuous lenticels on
the petiole even before the lamina is fully expanded. The number of leaflets per leaf is
variable even on the one plant, but the number most frequently encountered is 5-7.
Fruits are reported to be eaten by the fig bird (J. Wieneke, pers. comm.). It 1s closely
related to 7. pisicarpum which occurs in Papua New Guinea; however, the latter has
thinner usually smaller leaves, less prominent lenticels on the petioles, the staminate
inflorescence is longer and on the specimens examined appears to be more fragile and
easily broken when dry. The furrows on the ventral surface of the seed converge about
midway along the seed rather than at its base.

Etymology: This species is named for Margaret and Arthur Thorsborne of Cardwell, who
collected the first fertile material of it, in recognition of their contribution to biology in
the Rockingham Bay area.

i989

3
=

i

Austrobaileya 3(1)

156

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w
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cotyledons and first leaf x 0.5.

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Jlanchon in A.
us. lugd.-bat. 1: 79

(1863).

stems rounded, lenticellate; bark flaky; lenticels dotted.
res ped h (7-)9(-11) leaflets; leaflets thin, drying somewhat like

entral leaflet elliptical, apex shortly acute, base acute, 6-12 ¢ 3-7 cm: lateral le
elliptical, proportionally smaller, oblique at the base. Petiole striate, 10-10.5 c
lenticels inconspicuous until lamina fully expanded; central petiolule 3-5 c
ong. sta
pedicels thin and narrow, in the dry state a
Berry obovoid to elliptical, blackish at maturity, 0.8-0.9 X 0.6-0.7 c

ovoid, c. 5-6 mm X 6-7 - 1 per fruit; lateral furrows converge on the ventral surface
with the central furrow at about the |

onesia. 1869, Zollinger (K); de Vogel 5059 (Ks).

long;
long; lateral

inate inflorescence with flattened branches exceeding 6
opearing to be easily

ener

inea. New Guinea, Forbes

ST RINSE eee

TAS

= BE oe

Os
pes

Fig. 5. Tetrastigma petraeum: A. unopened bud of female flower. B. open female flower. Jetrastigma thorsbor-
neorum: C. open male flower and bud. D. side view of open male flower showing reduced ovary surrounded by

the disc. All scale bars equal 0.5 mm.

158 Austrobatleya 3(1): 1989

Vegetative material and material with immature staminate inflorescences collected
from the Lockerbie Scrub area at the top of Cape York Peninsula appears to belong to
this species. Confirmation of this identification at this stage is not possible because no
mature flowering or fruiting material has been seen. All specimens examined have usually
had between 7 and 11 leaflets per leaf and the leaflets are thin so that even tertiary
veins are easily seen on fresh material; lenticels are relatively inconspicuous on all but
old petioles. The collection by Gray 4295 bears young staminate inflorescences. Although
not fully mature, the buds are considerably smaller than buds of T. thorsborneorum
which are somewhat younger, and the top of the ovary and stigma are projecting well
above the disc. |
[Bamaga Rd, 8 km S of Cape York,10°45’/S, 142°31’E, Feb 1986, Gray 4295 (JCT,QRS); Between Lockerbie and
Somerset, 10°46’S, 142°30’E, May 1981, Hyland 21138V (JCT,QRS); Carnegie Range, Apr 1984, Jackes (JCT):

(BRT I3taioy Jul 1986, Jackes (JCT); South of Bamaga, 27 km SW of Cape York, Oct 1965, Smith 12603
1412)j.

Acknowledgements

_ [am indebted to Mr L. Pedley for kindly preparing the Latin diagnoses. The
assistance of the Directors of Herbaria who loaned specimens and/or allowed me access
to their collections is much appreciated. To my husband and those who went out of
their way to try to obtain fertile material of several species, a sincere ‘thank you’ — your
job is still unfinished! Mrs H. Winsor prepared the the Scanning Electron Micrographs.

References
JACKES, B.R. (1984). Revision of the Australian Vitaceae, 1. Aympelocissus Planchon. Austrobaileya 2(1): 81-86.
JACKES, B.R. (1987). Revision of the Australian Vitaceae, 2. Cayratia Juss. Austrobaileya 2(4): 365-379.
JACKES, B.R. (1988a). Revision of the Australian Vitaceae, 3. Cissus L. Austrobaileya 2(5): 481-505.

JACKES, B.R. (1988b). A Key to the Genera of the Australian Vitaceae. Proceedings Ecological Society of Australia
Lor 2y il.

JACKES, B.R. (1989). Revision of the Australian Vitaceae, 4. Clematicissus Planchon. Austrebaileya 3(1): 101-
102.

WEBB, L.J. (1978). A general classification of Australian Rainforests. Australian Plants 9: 349-363.
Accepted for publication 28 April 1989

(

Maps 1-2. 1. @? Tetrastigma pisicarpum;, @ T. thorsborneorum, * T. crenatum. 2. @ Tetrastigma nitens, * T.
petraeum.

Austrobaileya 3(1): 159-161 (1989) 159

NOTES

Eriocaulon longifolium Nees ex Kunth (Eriocaulaceae), a new record for Australia

Bentham (1878) stated that Eriocaulon australe R. Br. is “The largest of the
Australian species’. Certainly F. australe has the largest plants of the Australian species
of Eriocaulon and with leaves to 80 cm long and peduncles to 100 cm long it 1s readily
distinguished from other Australian species, the majority of which have plants which
are mostly less than 25 cm high. In the absence of any recent treatment of Australian
Eriocaulon, Bentham’s account contains the only Australia-wide key for identification
of Australian material of this genus. This situation appears to have led to the identification
of all material from tall plants with long leaves as E. australe, based solely on the size
of the plant.

Examination of some unidentified material at NI and DNA by the author in
preparation for a revision of the genus in Australia led to the identification of some
robust specimens as E. /ongifolium Nees ex Kunth using the accounts of Eriocaulon by
van Royen (1959) and Ruhland (1903). The superficial similarity of this material to that
of E. australe was apparent. Examination of further material from BRI and NSW
confirmed that two species were indeed represented by specimens identified as FE. australe
in Australia. Australian specimens thought to belong to E. /ongifolium were found to be
conspecific with material under that name from Malesia and also matched material cited
by van Royen (1959). The type does not appear to have been examined by van Royen
nor has it yet been seen by the author; however, based on the descriptions and illustration,
the species is readily identified. E. /ongifolium has thus been collected in Australia as
early as 1937 (Flecker s.n. [AQ200288] BRI).

The important distinguishing features of the two species are summarised in the
key below. Distribution of each of the two taxa in Australia is shown in Map 1. 2&.
longifolium is also widely distributed through Malesia, India and Madagascar while &.
australe is also recorded from New Guinea.

1. Leaves pubescent at base; heads 7-10.5 mm _ wide; involucral bracts

pubescent at base; receptacle pubescent with long hyaline hairs; flowers
TPEPVIELOUIS: Selene tee a. zp tae to ce i atl lame oo fle acetal oS “eS E. australe

Leaves mostly glabrous, rarely slightly pubescent at base; heads 4-6 mm
wide; involucral bracts glabrous; receptacle glabrous; flowers
dimerous.. .. 8% en carer ee ere anemia E. longifolium

Index to collectors of material examined
(1 = E. australe, 2 = E. longifolium)

Aston, H.1. 2668, 2; Bell, C. 380, 1; Blake, S.T. 2142, i; 2424, 1; 4989, 1; Brass, L.J. 2266, 1; 7603, 2; 7936, 1;
7951, 1: 8576, 1; 8638, 1; Brooker, M.1. 4093, 1; Clark, M.J. 1597, 1: Clarkson, J.R. 1046, 1; 2878, 1; Constable,
E.F. NSW22205, 1: Coveny, R. 347, 1; 3525, i; 6954, 1; Dunlop, CR. 3448, 2. 4399, 2; Durrington, L. 1441, |;
Dodson, J. s.n., x.1971, 1; Enoh, M. 327, 2; Fensham, R. 262, 2: 316, 2: Flecker, H. AQ200288, 2; NSW52582,
1: Fosberg, F.R. 62369, 2; Harrold, A.G. 182, 1; Hearne, D.A. 1544, 2: Henshall, T.S. 1835, 2; 3811, 2; Henty,
E.E. NGF38676, 1; NGF49406, 1; Higgins, R.L. s.n., ii.1919, 1; Himson, A. 55, 1; Hubbard, C.E. 3350, 1;
Ingram, C.K. NSW63344, 1; Johnson, L.A.S. NSW17733, 1: Latz, P.K. 3407, 2; Leach, GJ. 1400, 2; 1529, 2
2172, 2; 2307, 2; Levitt, D. 409, 2; Lucas, O.H. NSW58390, 1; Maiden, J.H. NSW58392, 1; Mckee, H.S. 9499,
i: McGillivray, D.J. 1892, 1; Michael, N. 370, 1; Mondi 278, 2: Morton, A.G. 755, 1; Must, J. 1642, 2; O°Gower,
A.K. NSW209445, 1; Ridsdale, C.E. NGF33583, 1; Robinson, A.C. s.n., 20.1.1975, 1; Royen, P. van 4872, 1;
Russell-Smith, J. 2101, 2; 2756, 2; 4422, 1; Scarth-Johnson, V. 833A, 1: Scortechini, s.n., s.d., 1; Shah, M. 571,
2: Sharpe, P.R. 129, 1; 1149, 1; 1646, 1; 1829, 1; 3970, 1; Simmonds, J.H. s.n., xii.1887, 1; Specht, R.L. 890, 1;
Stocker, G. 680, 2; Thorsborne, A. 185, 1; 268, 2; Waddy, J. 562, 2; White, C.T. s.n., 111.1915, 1; 1v.1915, 1;
Whitehouse, F.W. s.n., s.d., 2; Wightman, G.M. 908, 2: Womersley, J.S. NGF6882, 1; NGF9344, 1; NGF9354,
1: Wrigley, J.W. 368, 1.

Acknowledgements

The Directors of BRI, NSW and PERTH are gratefully acknowledged for the loan
of specimens.

160 Austrobaileya 3(1): 1989

126 130

#40 T&O0

— —=
1290 139

Map 1. Distribution of FE. australe (A) and E. longifolium (B).

Leach, Eriocaulon longifolium 161

References
BENTHAM, G. (1878). Eriocaulon. Floa australiensis 7: 190-198. London: Lovell Reeve & Co.
ROYEN, P. VAN (1959). Sertuluim papianum LL. Eriocaulaceae. Nova Guinea (n.s.) 10: 21-44.

RUHLAND, W. (1903). Eriocaulaceae. In Engler, A. (ed.), Das Pflanzenreich. IV, 13. Facsimile ed. Weinheim:
Engelmann-Cramer.

G.J. Leach

Northern Territory Herbarium
Conservation Commission of the Northern Territory
P.O. Box 496 Palmerston Northern Territory 0831

Accepted for publication 14 April 1989

Austrobaileya 3(1): 163-165 (1989) 163

PLANT PROFILE

Austrobaileya scandens C. White

Among the extant primitive angiosperms, the Austrobaileyaceae constitute, accord-
ing to Endress (1980), one of the systematically most isolated and geographically most
restricted Dicotyledonous families. Austrobaileya scandens, its only member, is a woody
vine endemic to the tropical rainforest of north Queensland. It was first discovered in
1929 by S.F. Kajewski when he was collecting for the Arnold Arboretum, USA. White
(1933) described it as a new genus and species of the family Magnoliaceae, but after
detailed research Croizat (1943) erected the new family Austrobaileyaceae for it in
recognition of its unique systematic position. Subsequent authors have followed this
suggestion. White (1948) described a second species, but, for reasons detailed below, it
is considered conspecific with A. scandens.

Austrobaileya scandens C. White, Contr. Arn. Arb. 4: 29 (1933). Type: Queensland.
Cook District: Boonjie, Atherton Tableland, 10 October 1929, S.F. Kajewski
1629 (holo: BRI).

Austrobaileya maculata C. White, J. Arn. Arb. 29: 255 (1948). Type: Queensland.
Cook District: Mt Spurgeon, September 1936, C.T. White 10734 (holo: BRI’).

Robust glabrous vine of rainforest canopy. Stems and older branches lenticellate; young
branchlets smooth, flattened, sometimes somewhat glaucous, often with narrow decurrent
lines, sometimes extending into narrow wings. Leaves opposite or subopposite, simple;
lamina leathery, smooth, bluish green, + shiny above, oblong-ovate or oblong to narrowly
ovate, 4.5-20 cm long, 2-9 cm wide; apex abruptly acuminate, the point itself 5-10 mm
long; base generally rounded to broadly cuneate; margins narrowly recurved; nervation
of midrib + flat or slightly recessed above, broad and somewhat raised below, 4.8
secondary nerves looping out towards margin on each side of midrib, and interstitial
reticulate venation prominent on both sides of lamina. Flowers solitary in leaf axils,
pedunculate; peduncles 7-15 mm long, clothed in small scattered bracts; bracts broadly
ovate, 1-2 mm long. Flowers + pendent, with a foetid odour to attract pollenator flies,
c. 5-6 cm diameter, exhibiting spiral phyllotaxis, bisexual. Tepals 11-23, greatly over-
lapping, varying much in shape and size, though outermost smallest, 5.5-30 mm long,
6-21 mm wide: outer tepals glossy green, + circular, inner creamish green with red-
purple dots, broadly oblong to obovate, margins very thin, irregularly denticulate.
Androecium consisting of stamens and (inner) staminodes, all creamish, papillose,
irregularly dotted with dark purple; stamens 7-11, laminar, varying in shape from
relatively flat to strongly boat-shaped depending on the position in the whorl, 14-17
mm long, 5-9 mm broad, with apex from almost acute or truncate to rounded, innermost
stamens approach staminodes in outline; ventral surface of stamen with 2 bisporangiate
thecae 3-5 mm long, each theca opening by a_ longitudinal slit, staminodes
9-16, 15-17 mm long, narrow, plicate or irregularly ridged. Gynoecium consisting of 8
or 9 free carpels 6.5-9 mm long; ovary c. 3 mm long, containing 4~14 ovules arranged
alternately in 2 longitudinal rows; styles c. 6 mm long, with canal exuding mucilage, all
converging apically and at anthesis tips embedded in a massive mucilage cap; stigma
transversely bilobed. Only a few carpels in each flower develop to maturity. Fruits
stalked, fleshy, orange-yellow, ellipsoid to globose berries, c. 5-7 cm long, 4..5 cm broad;
stalk 1.5-2 cm long. Seeds whitish, often lenticular. Fig. 1.

Distribution and habitat: This species has been collected from Mt Spurgeon in the Mt
Carbine Tableland area, south to the Atherton Tableland and vicinity of Ravenshoe,
and to the Downey Creek area and south to the Walter Hill Range near Tully. Though
collections of it are not numerous, it could be expected to occur over a wide area of the
wet tropics in both lowland and higher altitude rainforest. It is usually found as a canopy
vine growing to the crowns of trees 10-15 m tall.

Conservation status: In a number of areas the species has been recorded as locally
common, but since it is a vine of the canopy it is not often noticed. It is not considered

164

Austrobatleya 3(1): 1989

‘
-#
rf

i
:

3

trp Mert has Parnas ataae,
~~
gy
anh

re Re die Mert oa a

eats

RLS

iy

as
“a4 ereese

Fig. 1. Austrobaileya scandens. A. leaf X 0.5. B. flower X 1. C. fruit X 1. A, Webb & Tracey 6562; B, Jessup 884
& McDonald {in spirit); C, Jessup & Guymer GJM 2774 (in spirit). All at BRI.

Ross, Austrobaileya scandens 165

to be rare and threatened, and has been recorded from national parks through much of
its range.

Notes: Although Telford (1983) mentions two species, viz A. scandens and A. maculata,
other authorities (e.g. Endress 1980) consider there to be only one in the genus. Collections
at BRI show the key characters listed by White (1948) for separating his two species are
unreliable. The leaf shape varies from lanceolate to oblong on the one branchlet; flowers
on both type specimens have densely papillose stamens and staminodes; and as Endress
(1980) pointed out, the shape of each stamen’s apex depends on its position within the
spiral. On the basis of this evidence, the taxa previously included under A. scandens
and A. maculata are thus regarded as conspecific, and A. scandens, being the older name,
should be used for the species concerned.

Acknowledgements

I extend my thanks to L.W. Jessup, W.J.F. McDonald and G.P. Guymer for
providing excellent specimens from which W.A. Smith prepared the illustrations, and
for constructive comment on the manuscript.

References
CROIZAT, L. (1943). New families. Cactus and Succulent Journal (Los Angeles) 15: 64.

ENDRESS, P.K. (1980). The reproductive structures and s stematic position of the Austrobaileyaceae. Botanische
Jahrbucher fiir Systematic, Pflanzengeschichte und P. anzengeographie, 101: 393-433.

TELFORD, I. (1983). Austrobaileyaceae. In Morley, B.D. & Toelken, H.R. (eds), Flowering Plants in Australia.
Adelaide: Rigby.

WHITE, C.T. (1933). Ligneous plants collected for the Arnold Arboretum in North Queensland by S.F. Kajewski
in 1929. Contributions from the Arnold Arboretum 4. {-1%3.

WHITE, C.T. (1948). A new species of Austrobaileya (Austrobaileyaceae) from Australia. Journal of the Arnold
Arboretum 29: 255-256.

E.M. Ross
Queensland Herbarium, Meiers Road, Indooroopilly, Qld 4068

Austrobaileya 3(1): 167-171 (1989) 167

BOOK REVIEWS

W.D. Clayton & S.A. Renvoize. Genera Graminum — Grasses of the World, Kew Bulletin
Additional Series XIII. London: Her Majesty’s Stationery Office, 1986. 389 pp,
24 figures. No price quoted, but available in Australia for c. $58.

A review of this book has previously been published with reference to the effect
‘t has on the classification of the grass genera of southern Africa (de Winter 1987). A
further review with reference to the Australian genera is appropriate. As mentioned by
de Winter the Kew herbarium is certainly an excellent venue at which to compile a
contemporary account of the world’s genera, combining as it does a long tradition of
taxonomic and anatomical research together with a fine world-wide collection and library.
The considerable fund of knowledge established by agrostologists Stapf, Hubbard and
Bor and anatomist Metcalf have been brought to fruition in a succinct way by the
morphological and general contributions of Clayton and the anatomical summaries of
Renvoize.

The circumscription of genera of this world treatment nevertheless needs closer
scrutiny, particularly in regions where local experience and field knowledge has been
accumulated. In the case of southern Africa for example, de Winter questions the wisdom
of placing fairly unique genera such as Stiburus and Diandrochloa into synonymy with
Eragrostis or the recently established segregate arundinoid genera of southern Africa
(Conert 1987) into synonymy with Rytidosperma, itself a fairly dubious split from the
cosmopolitan genus Danthonia. In the Australian context there are similar examples e.g.
the placing of Thellungia into synonymy with Eragrostis and the sinking of Pseudopo-
gonatherum with Eulalia despite the fact that the former genus has paired pedicelled
spikelets and a distinctly awned upper slume. Overall the tendancy seems to have been
to adopt an attitude of combining genera ©.g. Diplachne with Leptochloa, Scrotochioa
with Leptaspis, Beckeropsis with Pennisetum, Diectomis with Andropogon and the exam-
ples given above, although the cases for segregating these genera in the first place have
been fairly lucidly and logically gone into e.g. Judziewicz (1984) where Scrotochloa 1s
separated from Leptaspis by 1) seemingly good characters. The recent generic splits in
the Triticeae (Léve 1984, Dewey 1984), based to a large extent on genomic characters,
although with a slight degree of morphological back-up, have been virtually ignored.

On the other hand the lumping process may not have gone far enough e.g. the
recent treatment of Mnesithea (Veldkamp et al. 1987), where four of the genera treated
as separate in Genera Graminum have been sunk under Mnesithea, but the rationale
for doing so is fairly clearly explained. In a similar vein is the recent placing of most
of Australia’s Brachiaria species and Panicum maximum in Urochloa (Webster 1987),
which at first examination does not seem plausible, but has some support at the level
of type of C4 leaf anatomy (Hattersley 1987). However the last three papers were
published since Genera Graminum, giving an indication of the amount of work currently
being undertaken on generic limits in the Poaceae.

In the main however Clayton and Renvoize have done a magnificent task of
summarizing the contemporary scene of how the world’s genera are constituted and
classified. They present their own views and would be the first to admit that their work
does not represent the last word in this area. Since their book was published a fair
number of genera have either been published or are in the pipeline and others have
been placed in synonymy. The trend will probably continue as generic limits are currently
being researched with increasing vigour, a reflection of this activity being the symposium
on grass classification and evolution in Washington in 1986 (Soderstrom ef al, 1987). A
recently published review on taxonomy in the tropics (Ng 1988) maintains there is a
high degree of subjectivity in the way we circumscribe our taxa, and this is certainly
reflected by the lack of agreement by grass taxonomists as to generic boundaries in many
genera. Ng asserts that the percentage disagreement of taxonomists working on the same
sroup of plants “may range between 5 and 30%” This subjective element certainly
accounts for the difference of opinion between agrostologists on the question of generic
limits.

168 Austrobaileya 3(1): 1989

In his review de Winter summmarizes well the contents of the book that there is
not much point repeating that here. Some final comments of the review in terms of the
great benefits of this compilation are however worth re-stating. “The authors have in
the Genera Graminum brought together a vast amount of information and successfully
summarized progress made during the last decades in grass taxonomy. All agrostologists
are in their debt. In the light of the strongly traditional, and not entirely consistent
approach followed, a ‘modern’ synthesis of the generic classification of the grasses has,
in my opinion, not been fully achieved. This is most likely what the authors themselves
meant to convey when they advised the reader that ‘...there is something here to annoy
everyone, so do not bother to chastise - think rather to improve.’ We await a definitive
treatment of grass genera in the future.” Whether this is a possible goal in view of the
degree of subjectivity in our methods of circumscribing genera, is debatable. However
techniques of accumulating data (Watson 1987) and analyzing it cladistically (Kellogg
1987) and phenetically (Baum 1987) with the aid of computers, give hope for the future.

References

BAUM, B.R. (1987). Numerical taxonomic analyses of the Poaceae. In Soderstrom, T.R., Hilu, K.W., Campbell,
C.S. & Barkworth, M.E., Grass Systematics and Evolution, pp 334-342. Washington, D.C. & London:
Smithsonian Institution Press.

CONERT, H.J. (1987). Current concepts in the systematics of the Arundinoideae. In Soderstrom, T.R., Hilu,
K.W., Campbell, C.S. & Barkworth, M.E., Grass Systematics and Evolution, pp 239~250. Washington, D.C.
& London: Smithsonian Institution Press.

DEWEY, D.R. (1984). The genomic system of classification as a guide to intergeneric hybridization with the
perennial Triticeae. In Gustafson, J.P., Gene manipulation in plant improvement, pp 209-279. Plenum
Publishing Corporation.

DE WINTER, B. (1987). Review of Genera graminum ~ Grasses of the World by Clayton, W.D. & S.A. Renvoize.
Bothalia 17: 299-300.

HATTERSLEY, P.W. (1987). Variations in photosynthetic pathway. In Soderstrom, T.R., Hilu, K.W., Campbell,
C.S. & Barkworth, M.E., Grass Systematics and Evolution, pp 49-64. Washington, D.C. & London:
Smithsonian Institute Press.

JUDZIEWICZ, E.J. (1984). Scrotochloa, a new genus of paleotropical grasses. Phytologia 56: 299-304.

KELLOGG, E.A. (1987). Phylogenetic analyses of the Gramineae. In Soderstrom, T.R., Hilu, K.W., Campbell,
C.S. & Barkworth, M.E., Grass Systematics and Evolution, pp 310-322. Washington, D.C. & London:
Smithsonian Institution Press.

LOVE, A. (1984). Conspectus of the Triticeae. Feddes Repertorium 95: 425-521.

NG, F.S.P.(1988). Problems in the organisation of taxonomic work. Australian Systematic Botany Society Newsletter
54: 2

—
=

SODERSTROM, T.R., HILU, H.W., CAMPBELL, C.S. & Barkworth, M.E. (1987). Grass systematics and evolution.
Washington, D.C. & London: Smithsonian Institution Press. oe

VELDKAMP, J.F., DE KONING, R. & SOSEF, M.S.M. (1986). Generic delimitation of Rotthoellia and related
genera (Gramineae). Blumea 31: 281-307.

WATSON, L. (1987). Automated descriptions of grass genera. In Soderstrom, T.R., Hilu, H.W., Campbell, C.S.
& Barkworth, M.E., Grass systematics and evolution, pp 343-351. Washington, D.C. & London: Smithsonian
Institution Press.

WEBSTER, R.D.(1987). The Australian Paniceae, 322 pp. Berlin and Stuttgart: J. Cramer.

B.K. Simon
Queensland Herbarium, Meiers Road, Indooroopilly, Qid 4068

R.D. Webster. The Australian Paniceae (Poaceae). Berlin and Stuttgart: J. Cramer, 1987.

This book is a precursor to the Flora of Australia account of the Paniceae. The
descriptions and keys have been computer generated using the DELTA package of M.
Dallwitz (1986) and are an extension of those produced by the same author for the
genus Digitaria (Webster 1984), both a result of three years spent at the Taxonomy Unit
of the Research School of Biological Sciences on a grant from the Australian Bureau of
Flora and Fauna, through the Australian Biological Resources Study scheme. Thus far

Book Reviews 169

these are the only two accounts of grasses to be produced by the DELTA system at
species level and this is the first floristic account. The computer generated keys and
descriptions of grass genera from the Taxonomy Unit are now well known documents
of the state of the art of the DELTA system (Watson & Dallwitz 1985 & 1988).

The 308 taxa of Australian Paniceae recognized by Webster are described against
4 character list of 208 characters, of which 42 are confined to vegetative characters, 52
to inflorescence characters, 104 to spikelet characters and one for chromosome number,
where known. Other information given includes native or introduced status, distribution
by state and the botanical regions of Barlow (1984) and vegetation regions of Moore
(1975), moisture requirements, flowering period, general remarks, representative specimen
Citations and relevant literature citations. In addition the full character list is given and
distribution maps of all entities are arranged alphabetically at the end of the book.

The arrangement of the text is alphabetical by genus and species, with keys to
species (where appropriate) being placed after the generic descriptions and remarks on
the genera. The DELTA descriptions are of the usual form, in that they are comparative,
and follow a standard character sequence, punctuation and terminology. A full stop
indicates the end of a sequence of characters describing a primary feature, a semicolon
separates the characters and a comma is used where more than one character state
occurs. Additional comments covering information not mentioned by the character list
are contained in brackets.

The publication of this floristic compilation within a relatively short time of the
completion of the research work is indeed a very impressive feat, both a function of
efficiency of the DELTA system and the author’s good manipulation of the system and
of the diligence of the author himself.

Before any comments are made concerning observed deficiencies of the work a
few comments on the DELTA system and my experiences in using it, both with Websters’s
data, generously provided on tape from the Taxonomy Unit, and my own data set on
the Andropogoneae, are worth making. Webster uses one items file (data base on taxa)
for both genera and species whereas for the Andropogoneae | created separate files for
the species of each genus and for the genera; in this way | found the generation of
descriptions and keys to be more manageable in terms of the different sorts of characters
that constitute genera from those that make up species. Furthermore extreme care has
to be taken in weighting the characters required for key generation as opposed to those
needed for inclusion in a description. I feel that in the Webster keys some characters
which are difficult to discern have been included in the keys (in some cases the only
differentiating character) whereas they are essentially only should be included in descrip-
tions. Examples are couplet 14 of the key to genera (concerning disarticulation of the
spikelet) which cannot always be used as it depends on the age of the plant, and couplet
29 of the same key (concerning the shape of the pedicel apex) as the character is difficult
to apply (see comments on the same character later in the discussion on the key to the
species of Digitaria). An anomaly I discovered generating keys with the Webster Paniceae
data set was that sometimes a genus keyed out separately from its constituent species
when a key to both was generated. An example is the key to the species of Cenchrus
when the genus and four species ( C. echinatus, C. robustus, C. incertus and C. longispinus)
are keyed out as having the upper lemma smooth whereas the remaining species (C.
elymoides, C. biflorus, C. setiger, C. ciliaris, C. brownii and C. australis) are keyed as
having the upper lemma muricate. Although the anomaly is not reflected in any keys
in the book itself, it does show up in the descriptions of the relevant taxa.

In a number of cases Webster places together species of Paniceae where he regards
them as being glabrous and hairy forms of one species, however the grounds for such a
decision seem rather flimsy to me without any experimental evidence, and I prefer not
to follow this move. Examples are the placing of Brachiaria notochthona with glabrous
spikelets with B. gilesii (treated as a Urochloa by Webster), and the placing of B. windersii
with hairy pedicels in synonymy with B. praetervisa (as a Urochioa).

In a few cases Webster presents results which are not taxonomically correct. For
example he confuses the identity of Digitaria milanjiana with the form of D. eriantha
subsp. pentzii previously called D. decumbens and commonly known as Pangola Grass.

170 Austrobaileya 3(1): 1989

The former species is in Australia but is only known in cultivation, whereas the latter
cultivar has escaped and become established in a number of foredune regions in
Queensland. The species Pseudochaetochloa australiensis, a dioecious species previously
only thought to have been collected in the male state, is now known to be represented
in the female state by the grass Pennisetum arnhemicum (Macfarlane, pers. comm.).
This fact was apparently known to Webster but there is no indication of it in his text.
Two species of Oplismenus (O. aemulus and O. imbecillis) are placed together in synonymy
under O. hirtellus (also incidentally treated in the same way by Davey & Clayton 1978),
but the two are distinctly different, particularly the leaf shapes, and grow sympatrically
in the rain-forest floors of south-eastern Australia. Within the same genus the variety
mollis of O. undulatifolius is placed in synonymy with the type variety from Italy.
However the entity is morphologically different, probably a function of its phytogeography,
if indeed it is the same species. For the time being I am following the status given it
by Vickery (1975).

In a few cases I think Webster incorporates too much variation in spikelet size
in some of his species and the situation is better expressed taxonomically if recognition
is given to the more discrete entities. An example which comes to mind is the native
Hymenachne acutigluma which has bigger spikelets than the South American H. amplex-
icaulis but is placed in synonymy with the latter species. The neotropical species has
recently been established as a pasture plant in Australia and is likely to become naturalised
so it is useful to be able to tell them apart. Another example are the two native species
Eriochloa australiensis and E. longiglumis, which have been placed together by both
Webster and Vickery (1975) but the latter has spikelets which are distinctively longer.
Cenchrus pennisetiformis is placed in synonymy with C. ciliaris but the character of
basal bristle fusion, disclaimed by Webster, can be applied consistently. On the other
hand C. australis R. Br. is resurrected from C. caliculatus on the grounds of having
smaller involucres and spikelets but I do not consider these characters to be of sufficient
magnitude to warrant specific delimitation.

There are a few examples of Webster’s names being replaced by more correct
names, these may not have been known to the author or were published at about the
same time. Examples are Holcolemma dispar W. Clayton for Paspalidium inaequale (F.
Muell.) Hughes (Clayton 1987), Panicum laevinode Lindley for P. whitei J. Black (Jacobs
1984) and Ichnanthus pallens var. majus (Nees) Stieber for I. vicinus (Bailey) Merr.
(Stieber 1987). Lazarides (1980) has Panicum luzonense Presl (a name not mentioned
by Webster) as a synonym of P. cambogiense Balansa but this was published 60 years
before Balansa’s name; if they apply to the same species the former is the correct name.
Cenchrus setigerus is wrong botanical Latin and must be corrected to C setiger.

Species omitted by Webster include Arthragrostis aristiglumis B. Simon, Echin-
ochloa picta (Koenig) Michael and Paspalum wettsteinii Hackel. Spinifex X alterniflorus
Nees is mentioned in the text but not keyed out.

The key to Digitaria is essentially the same as Webster (1984). The basic flaw of
this key is that the character of pedicel apex is confused in definition (for example I
find the distinction between the three states of truncate, cupuliform and discoid, as
illustrated by the photographs referred to, very difficult to apply) and yet is used as the
first entry to the key.

The big taxonomic decision made in this book, resulting in many nomenclatural
changes, involves the transfer of all species of Brachiaria except B. eruciformis and of
Panicum maximum to the genus Urochloa. This is based on a group of characters
previously not considered instead of the classical differences, the main one of which is

__ Spikelet orientation. These new characters include texture of the upper lemma, point of
_ disarticulation of the spikelet and whether the upper lemma is mucronate. Other characters

based on leaf anatomy (Hattersley 1987) support these findings to a certain extent. While
Webster's ideas certainly appear to have merit I feel to accept the changes for Australia
before a study has been made on the generic limits of these genera as a whole is slightly
premature. Such work 1s currently being undertaken at the University of Oklahoma and
the outcome of the research based on many characters and all relevant species is awaited.

The distribution maps at the end of the book give a good impression with the
actual distributions laid over the geobotanical regions of Barlow (1984). Flora of Australia

Book Reviews 171

policy is now to follow Barlow’s revised botanical regions (Barlow 1986) so that Webster’s
maps will have to be redrafted for the official Flora account. In most maps the area
occupied by an entity is completely blocked out but in others (e.g. some species of
Digitaria) the area is covered by a mosaic of dots; one technique would have sufhced.
The method of construction of the maps is rather a mystery in terms of the numbers
of specimens that were examined. In the case of the Brisbane material that was loaned
to Webster only about 10% of the Paniceae was actually requested. A comparison with
the HERBRECS generated maps from the Queensland Herbarium revealed a fairly close
similarity for most species that were compared.

The native or introduced status of a number of species is different from that
previously recorded. Two examples are Pennisetum alopecuroides (L.) Sprengel and
Panicum incomtum Trin., both regarded by Webster to be introduced but thought of by
others to be native.

References

BARLOW, B.A. (1984). Proposal for Delineation of Botanical Regions in Australia. Brunonia 7: 195-201.
BARLOW, B.A. (1986). A revised natural regions map for Australia. Brunonia 8: 387-392.

CLAYTON, W.D. (1987). Miscellaneous notes on panicoid grasses. Kew Bulletin 42: 401-403.

DALLWITZ, M.J. (1986). User’s guide to the DELTA system. A general system for coding taxonomic descriptions.
CSIRO, Division of Entomology.

DAVEY, J.C. & Clayton, W.D. (1978). Some multiple discriminant function studies on Oplismenus (Gramineae).
Kew Bulletin 33: 147-157.

HATTERSLEY, P.W. (1987). Variations in photosynthetic pathway. In Soderstrom, [.R., Hilu, K.W., Campbell,
C.S. & Barkworth, M.E. (eds), Grass systematics and evolution. Washington D.C. & London: Smithsonian
Institution Press.

JACOBS, S.W.L. (1984). An earlier name for Panicum whitei (Gramineae). Kew Bulletin 40: 662.

LAZARIDES, M. (1980). The tropical grasses of Southeast Asia. Vaduz: J. Cramer.

MOORE, R.M. (1975). Australian Grasslands. Canberra: Australian National University Press.

Sas M. (1987). Revision of Jchnanthus sect. Foveolatus (Gramineae: Panicoideae). Systematic Botany 12:
{ 6

VICKERY, J.W. (1975). Flora of New South Wales, Gramineae 19: 125-306.

WATSON, L. & DALLWITZ, M.J. (1985). Australian Grass Genera: anatomy, morphology, keys, and classification
(Second Edition). Canberra: Australian National University, Research School of Biological Sciences.

WATSON, L. & DALLWITZ, M.J. (1988). Grass Genera of the World: ifiustrations of characters, descripttons,
classification, interactive identification, information retrieval.

scree Pittal (1984). A revision of the genus Digitaria Haller (Paniceae: Poaceae) in Australia. Brunontia 6:
131-216.

B.K. Simon
Queensland Herbarium, Meiers Road, Indooroopilly, Qld 4068

172 Austrobaileya 3(1): 1989

CORRIGENDUM

Austrobaileya 2(5) 1988

p 514. The caption for the distribution maps should read A. D. nummularia. B. D.
ovata. C. D. major.

S. R. Hampson, Government Printer, Queensiand—1989

104400

Notes for Authors

Austrobaileya publishes original papers in systematic_botany and related fields.
Preference will be given to papers relating to the flora of Queensland. All papers are
refereed and the editorial committee reserves the right to reject papers.

Manuscripts must be submitted in duplicate to The Editor, Austrobaileya, Queens-
land Herbarium, Department of Primary Industries, Meiers Road, Indooroopilly, Qld
4068. They must be double-spaced typewritten, with 2.5 cm margins in the first instance.
After refereeing, the corrected manuscript may alternatively be submitted on an MSDOS
disk as an unformatted ASCII file (DOS Text File Format, e.g. CTRL-F5, | in WordPerfect
Version 4.2 or 5.0), saved twice, accompanied by a printed copy. All illustrations should
he submitted with the text. One set of proofs will be sent to authors.

For style and layout the most recent number of Austrobaileya should be followed,
particularly in the use of subheadings for distribution, etymology, etc. A detailed guide

may be obtained from the editor. Papers must be concise and illustrations should make
economical use of available space.

Where possible illustrations should be submitted in a size suitable for reproduction
without reduction in size. The maximum size of a printed illustration is 12.5 cm X 18.5
cm. Do not put lettering on the illustration but indicate separately on a photocopy or
overlay. Illustrations should be cited in the text. Typed captions should be supplied on
a separate page. All illustrations, both line drawings and photographs, are to be numbered
as figures in a common sequence. Published illustrations remain the property of the
Queensland Herbarium.

Tables should be as simple as possible. Long and/or complicated tables should
be avoided. The tables should be numbered consecutively and each should be cited in
the text.

Standard Abbreviations

Titles of journals cited in the text should be abbreviated according to Botanico-
Periodicum-Huntianum (Hunt Botanical Library, 1968) while names of books should be

abbreviated in accordance with F.A. Stafleu and R.S. Cowan, Taxonomic Literature, edn
2 (W. Junk, 1976-1988). In bibliographies the titles of both journals and books should
be given in full. All entries in the bibliography are to be referred to in the text and all
references to papers or books in narrative text are to be included in the bibliography.

Author abbreviations should follow the Draft Index of Author Abbreviations
compiled at the Herbarium, Royal Botanic Gardens, Kew (HMSO, 1980). Acronyms used
for herbaria cited in the text should be in accordance with P.K. Holmgren, W. Kreuken
and E.K. Schofield (1981), Index Herbariorum Pt 1, edn 7, Regnum Vegetabile 106.

Other abbreviations which may be used in citing specimens are S.F. (State Forest),
S.F.R. (State Forest Reserve), L.A. (Logging Area), T-.R. (Timber Reserve) and an AQ
‘number: This number is the computerised collection number situated on the sheet and/
or on the label of specimens housed in the Queensland Herbarium (BRI). It is distinct
from the BRI number which is a framed sheet number associated with the name
“Queensland Herbarium Brisbane”, stamped on the sheet.

Austrobaileya 3(1): 1-172 (1989)

CONTENTS

A new species of Coix L. aaeeae) from Australia
B.K. Simon .... depts df Bike 4

Ceropegia cumingiana. Decne (Asclepiadaceae)
P.V. Bruyns & P.J. Forster |

The genus Zornia J. Gmelin i rear meet in Australia
S.T. Reynolds & A.E..Holland . ea fee

Two new species of Eucalyptus iia aa nae from central EAS PHENDD
A.R. Bean & M.I.H. Brooker mn 7 i 4 :

A new species of Marsdenia R. Br. cand seein ie from eastern Australia
J.B: Williams . ee, | SRY Baye 28 be ata

Kailarsenia Tirvengadum emend. Puttock (Rubiaceae: Gardenieae) in
Australia ,
C.F. Puttock ......

The genus Ancana F. Muell. EAGER? i in Australia
L.W. Jessup 5 Dig ts dah tte

Secamone R. Br. (Asclepiadaceae: hciaipiiastnataacaie Es in Australia
P.J. Forster & K. Harold . ee ore ee

Studies in Australian grasses: 4. Taxonomic and nomenclatural studies
in Australian Andropogoneae
B.K. Simon eT

Revision of the Australian Vitaceae, 4. Clematocissus Planchon
B.R. Jackes oP. bid thay Sn

Bryophytes in a sub- Prats MENS eapengiey
J. Windolf . | MD ewe ain se) e ins

Notes on hecienbadincete- |
P.]. Forster

Notes on Trachymene pina pr Eont yt in LER, l
A.E. Holland . or ia

New species of Bulbophyllum section Oxysepalum (Orchidaceae) in

Australia
B. Gray & D.L. Jones . .

Revision of the Australian Vitaceae, 5. Sena gees ORE) Planchon
B.R. Jackes Se See 7

NOTES

Eriocaulon longifollum Nees ex Kunth (Eriocaulaceae), a new record for
Australia
2 eg) Bc) a

PLANT PROFILE

Sani scandens C. White ©
. Ross .. .. a

Book Reviews ........

Corrigendum

101

103

109

135

141

149

159

163
167
172