Volume 5
Number 4 2000
AUSTAOBAIICVA
A Journal of Plant Systematics
Queensland Herbarium
&
Queensland Government
Environmental Protection Agency
Editorial Committee
L W. Jessup (editor)
R. J.F. Henderson (technical advisor)
B.K. Simon (technical advisor)
Desktop Publishing
YC. Smith
T.A. Kerr
J.P. Ward
Austrobaileya
Vol. 1, No 1 was published on 1 December 1977
Vol. 5, No.3 was published on 29 December 1999
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ISSN 0155-4131
© Queensland Herbarium 2000
Austrobaileya is the journal of the Queensland Herbarium and is devoted to publication of
results of sound research and of informed discussion on plant systematics, with special
emphasis on Queensland plants.
Opinions expressed by authors are their own and do not necessarily represent the policies or
views of the Queensland Herbarium.
Caution: this version of Austrobaileya was produced
before final corrections to bromides were applied.
Date of Publication of Austrobaileya 5(4) was 15 December 2000
Contents
A revision of Stylidium subg. Andersonia (R.Br. ex G.Don)
Mildbr. (Stylidiaceae)
A.R.Bean. 589
Chromosome numbers of someAcanthaceae from Papua New Guinea
Thomas F. Daniel. 651
Wahlenbergia celata (Campanulaceae), a new species from central Queensland
Paul I. Forster. 661
The identity of Flindersia pimenteliana and F. oppositifolia
(Rutaceae): Evidence fromDNA Sequences
Kirsten D. Scott, Wayne K. Harris & Julia Playford. 667
Cereus uruguayanus (Cactaceae) and its naturalised occurrence in
Queensland, Australia
Paul I. Forster & Miriam Schmeider. 671
A revision of Eucalyptus normantonensis Maiden & Cambage
(Myrtaceae) and its allies
A.R. Bean. 679
Homoranthus coracinus (Myrtaceae), a new species from Queensland
A.R. Bean. 687
Two new species of Apatophyllum McGillivray (Celastraceae) from
Queensland
A.R. Bean & L.W. Jessup. 691
Jasminum domatiigerum subsp. australis (Oleaceae), a new subspecies
from North Queensland
Wayne K. Harris & William J. McDonald. 699
A new species and lectotypification in Campanulaceae: Lobelioideae
DavidAlbrecht. 705
Chamaesyce ophiolitica (Euphorbiaceae), a new and endangered
species endemic to serpentine vegetation in central Queensland
Paul I. Forster. 711
Clausena smyrelliana (Rutaceae: Aurantioideae), a new and critically
endangered species from south-east Queensland
Paul I. Forster. 715
The first botanical record for Australia
John F.P Windolf. 721
Rediscovery of Dischidia torricellensis (Schltr.) P.I.Forst., an unusual epiphytic
asclepiad fromNew Guinea.
Pauli. Forster.725
(continued)
Notes
Cryptolepis papillata P.I.Forst. & Sarcolobus porcatus P.I.Forst.
(Asclepiadaceae), newly recorded from West Papua
Forster, PI. 729
A new combination in Morinda L. (Rubiaceae) for Australia
Halford, DavidA. & Henderson, Rodney J.F. 731
A new combination inProstantheraLabiW. (Lamiaceae)
Bean,A.R. 733
A new combination in Corymbia section “Politaria”: C. citriodora subsp.
variegata (Myrtaceae)
McDonald, M.W. & Bean,A.R. 735
A revision of Stylidium subg . Andersonia (R.Br. ex G. Don)
Mildbr. (Stylidiaceae)
A.R. Bean
Summary
Bean, A.R. (2000). A revision of Stylidium subg. Andersonia (R.Br. ex GDon) Mildbr. (Stylidiaceae).
Austrobaileya 5(4):589-649. Descriptions, distribution maps and notes on habitat, conservation
status and taxonomic affinities are provided for the 37 species of Stylidium subg. Andersonia
which are indigenous to mainland south-east Asia (China, Vietnam, Malaya, Cambodia, Laos,
Thailand, Burma, Bangladesh, India), Sri Lanka, Malesia (Sumatra, Philippines, Sulawesi, Java,
New Guinea, Aru Islands) and northern Australia (Western Australia, Northern Territory, Queensland,
New South Wales). A key to the subgenera of Stylidium occurring in these areas is provided as well
as keys to the sections and species of S. subg. Andersonia. Eight new species; namely S. confertum,
S. longissimum, S. aquaticum, S. oviflorum, S. stenophyllum, S. accedens, S. divergens and S.
ensatum are described and illustrated. A cladistic analysis of the group is provided. Five taxonomic
sections are proposed of which two are new. S. subg. Alsinoida Mildbr. is reduced to sectional rank.
The name Stylidium tenerum Spreng. is resurrected for the taxon formerly known in Australia as
S. uliginosum Sw. ex Willd. S. dunlopianum Carlquist is newly recorded for Western Australia. S.
prophyllum Lowrie & Kenneally, S. fluminense F.L.Erickson & J.H.Willis and S. diffusum R.Br.
are newly recorded for the Northern Territory. S. cordifolium W.Fitzg. is newly recorded for
Queensland.
Keywords: Stylidium, taxonomy, Stylidiaceae, Australia, Malesia, south-east Asia, trigger-plants,
keys, new species, Stylidium subg. Andersonia, Stylidium subg. Alsinoida
A.R. Bean, Queensland Herbarium, Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha Road,
Toowong, Queensland 4066, Australia
Introduction
The genus Stylidium is characterised by flowers
having a column bearing both anthers and
stigma which moves rapidly in response to
physical stimulus, usually provided by an
alighting insect. This feature has given rise to
the common name of trigger-plant. The genus
as a whole reaches its greatest diversity in south¬
western Western Australia. Comprehensive
taxonomic accounts of the genus have been
provided by Brown (1810), DeCandolle (1839),
Mildbraed (1908) and Erickson (1958).
The group of species treated here was
first named (as Andersonia) by Brown (1810),
but he did not specify the rank of this taxon. G.
Don in 1834 treated th q Andersonia group as a
section of Stylidium. Mueller (1859) and
Bentham (1868) included some of the species
treated here within S. sect. Nitrangium (Endl.)
Accepted for publication 14 April 2000
Sonder. Mildbraed (1908) raised S. sect.
Nitrangium and S. sect. Andersonia to
subgeneric level. The former was confined to
south-western Western Australia, the latter to
tropical Australia and south-east Asia.
Mildbraed (loc.cit.) established S. subg.
Alsinoida for S. alsinoides and its allies. While
this is a distinctive group because of the
laterally fused petals, it does not otherwise
differ greatly from groups in S. subg.
Andersonia and is, therefore reduced here to
sectional rank. Cladistic analysis has been used
to place the species in phylogenetic context
and to provide justification for the sectional
classification used.
S. subgenus Andersonia is distinguished
by its linear capsules which are sessile or almost
so, labellum without basal appendages, sepals
which are never all free, brown seeds and
mostly annual habit.
590
Terminology
The terms scapose and scapiform are used
when the inflorescence is borne on a leafless
scape. The scape is recognisable as it is
distinctly different from the leaf-bearing stem
in texture, shape, diameter, and often its
indumentum. In a scapose inflorescence, the
scapes arise from ground level, and the leaves
are in a basal rosette (e.g. Fig. 3E). In a
scapiform inflorescence, the scapes arise from
the apex of the leafy stem (e.g. Fig. 3A), where
there is often a terminal rosette of leaves. In
some species, there is no discernable scape;
there is no discontinuity in the stem tissue
between the lower (vegetative) part and the
upper (fertile) part (e.g. Fig. 7D).
The flowers of all Stylidium species are,
strictly speaking, gamopetalous, as they
possess a corolla tube, albeit usually rather
short and inconspicuous. However, the 4
largest corolla lobes (as they should strictly be
called) are termed petals in this paper, while the
fifth, always much smaller, has for many years
been called the labellum (see Fig. 3C, 6C,6F).
The four large corolla lobes are said to be “free”
if they are not united beyond the distal end (or
“throat”) of the corolla tube. This interpretation
was also used by Erickson (1958) and avoids
confusion when referring to the apical
ornamentation of the corolla parts, which would
otherwise be known as “lobes of the corolla
lobes”. Because the pattern of petal fusion is
very valuable in diagnosing Stylidium species,
a further special terminology is used here. A1
and A2 refer to the anterior petals (on either
side of the labellum); PI and P2 refer to the
posterior petals, with A1 being adjacent to PI.
The ‘+’ operator indicates that the petals
indicated are free (sensu Erickson), while a
operator indicates that the petals indicated are
fused e.g. A1 + A2 + (P1&P2) means that the
anterior petals are free from the posterior petals
and from each other, but that the posterior
petals are fused to each other (e.g. Fig. 3C).
The reduced fifth corolla lobe in Stylidium
is called the labellum. It is found between the
anterior petals, and is attached either at the top
of the corolla tube (F ig. 8B, 8F), or on the outside
of the tube (Fig. 5B).
Austrobaileya 5(4): 589-649 (2000)
The term paracorolla, used here, was
introduced by Slooten (1954), and refers to
small lobes or flanges of tissue attached to the
corolla at the throat (e.g. Fig. 5C). This term is
roughly equivalent to the more commonly used
term “throat appendages”, but is more
comprehensive, as it covers situations where
there is a continuous raised ring of tissue.
The “paracorolla glands” referred to in
the key and descriptions are conspicuous
globular to ellipsoidal glands, commonly
orange or golden in colour, attached to the
paracorolla between the anterior and posterior
petals (e.g. Fig. 4G). They occur in only a few
species, most notably in S. schizanthum and
its allies, and can usually be readily seen even
on dried herbarium specimens.
Corolla colour in this paper refers to the
adaxial surface only, the surface which is most
readily seen from above. While the other surface
may often be a different colour, this is rarely
recorded.
The term “corona” is used here for the
cluster of white hair-like structures radiating
from the anthers of some Stylidium species (see
Erickson (1958: plate 53, no. 2)). Acorona is not
often found amongst species belonging to S.
subg. Andersonia , but occurs commonly in
some other groups within Stylidium.
Materials and methods
This revision is based on the examination of
herbarium material fromAAU, BM, BRI, CANB,
DNA, L, K, MEL, MO, NSW, P, QRS and RSA,
and was supplemented by field work by the
author throughout Queensland and in north¬
eastern Northern Territory.
Floral measurements were based on
material preserved in spirit, or where this was
not available, on herbarium material
reconstituted by boiling in water. Details of the
collections used for these measurements are
given in each species treatment. Leaves, stems,
scapes, capsules and seeds were measured
from dried material. Sepal dimensions are taken
from flowering material; it is apparent that the
sepals lengthen after flowering in some species.
591
Bean, Stylidium subgen. Andersonia
Morphological data for all taxa have been
recorded as a DELTA dataset (Dallwitz et al.
1993), and species descriptions have been
generated by DELTA from these data. 102
morphological characters were measured for
each species, with detailed measurements of
several flowers.
The accuracy of dimensions given for
various plant parts are: glandular hairs and
seeds, ±0.05 mm; scape diameters, sepals,
corolla, petals, labellum, paracorolla and
capsule width, ±0.1 mm; bracts, capsule length
and column, ±0.5 mm; larger leaves and
petioles, ±1 mm; smaller leaves ±0.2 mm.
Measurements are given as circa only when
there was difficulty in providing accurate
measurements e.g. petiole length for some
species, or when less than 3 measurements are
possible for the character, because of lack of
available material.
Details of the type localities of species
described by Robert Brown have been gleaned
from Vallance (1990).
Distribution maps are provided for all
species. Symbols indicate that the taxon has
been recorded within that 1° x 1° square. Solid
(or filled) symbols indicate that there is at least
one recent collection (1960-present). Outline
(or unfilled) symbols indicate that the most
recent collection for that l°xl° square was
before 1960. The vast majority of records were
from herbarium specimens seen by the present
author, but a few additional reliable locational
records have been added to the maps from
Lowrie and Kenneally (1997) for S. perizostera
and S. prophyllum, from Haridasan et al. (1983),
Huq (1986), Baaia & Gogoi (1995) for S. kunthii ;
and from Bhaskar & Kushalappa (1992) for S.
tenellum. Latitudes and longitudes for the non-
Australian specimens were determined only
after considerable effort by the present author,
and hence in the specimen citations they have
been retained, to assist readers to pinpoint the
localities.
Phylogenetic relationships were
examined with the cladistic program PAUP
(Version 3.1.1) (Swofford 1993). The most
parsimonious interpretation of the data was
sought (Parris 1983). Heuristic searches were
used, with the following options: Addition
sequence = Random, number of replications =
10, Mulpars in effect; TBR Branch swapping.
A subset of the 37 taxa was chosen for the
cladistic analyses, because using the full set
of taxa made analyses extremely slow and did
not improve resolution. Other preliminary
analyses were rejected because of errors
detected in the dataset or because some
characters included in them were found to be
unsuitable for this type of analysis. For example,
leaf shape was used in initial analyses, but later
deleted because it displays continuous
variation for the taxa involved. Morphological
data from 19 ingroup taxa (± 1 outgroup taxon)
and 22 characters were used for the final
cladistic analysis. All distinctive or unusual
taxa were included, and where there were
groups containing two or more obviously
related species, a selection of these were made.
The characters used for the analysis are given
in Table 1. All characters used were unweighted
and their states unordered (Table 2). Bootstrap
analysis (Felsenstein 1985) with 100 replicates
(heuristic, 10 random addition relicates,
Mulpars in effect, and Steepest descent on)
was used to assess the relative support for
various branches of the tree. Forstera
bellidifolia Hook. (Stylidiaceae) was selected
as the outgroup, as it displays many
unspecialised character states and probably
belongs to a sister group of Stylidium.
Results of Cladistic Analysis
The unweighted analysis resulted in 156 most
parsimonious trees each of 47 steps; ci= 0.708,
ri= 0.684, rc= 0.484. The strict consensus tree
showing bootstrap values, 50% majority rule
consensus tree and one of the most-
parsimonious trees are shown (Fig. 1,2). Various
tree topologies and the cost of modifying
topologies were investigated using MacClade
vers. 3.07 (Maddison and Maddison 1992). Five
clades are considered sufficiently distinct for
their recognition as taxonomic sections. S. subg.
Alsinoida Mildbr. is reduced in this paper to
sectional rank. S. sect. Tenella is recognised
on the basis of its strong support on the
majority rule tree. Some of its distinctive
features (the largely bare stems, the small bract¬
like leaves and the very short glandular hairs)
were not included in the analysis as the
592 Austrobaileya 5(4): 589-649 (2000)
characters are continuously variable in the Andersonia may be paraphyletic, based on the
subgenus. The residual section, S. sect. majority rule tree, but further study is desirable.
Table 1. Characters and character states used for the cladistic analysis
1. Leaf arrangement (#11): basal rosette (0); mostly in
terminal rosette (1); scattered along stem (2).
2. Leaf base (#17): truncate (0); cuneate, obtuse or
cordate (1).
3. Scapes (#21): present (0); absent (1). All species
can be classified unambiguously for this
character.
4. Inflorence type #27): determinate, monochasially
cymose (0); 1-flowered (1); umbellate (2). This
character is assessed on reasonably mature
individuals. Very young plants will all be 1-
flowered for a time.
5. Hypanthium indumentum (#39): glandular-hairy
throughout (0); glabrous (1); glandular-hairy at
distal end only (2).
6. Fusion pattern of sepals (#41): fused into 2 entire
obtuse lips (0); all free (1); with 3 free and 2
fused for more than half their length (2); fused
into 2 emarginate lips (3). These patterns are
constant within each species.
7. Sepal indumentum (#44): glandular-hairy (0); glabrous
(1). Specimens are coded as glandular-hairy even
when there are just a few scattered glands on
the margin.
8. Sepal apex (#45): acute (0); obtuse (1).
9. Corolla indumentum (#47): glandular-hairy on petals
only (0); glabrous (1); glandular-hairy on tube
only (2); glandular-hairy on tube and petals
(3).
10. Corolla tube incision (#49): with sinus on anterior
and posterior sides (0); without sinus (1); with
sinus on anterior side only (2).
Ecology and Distribution
Most species belonging to Stylidium subg.
Andersonia occur in tropical areas with a
monsoonal climate, where the rainfall is strongly
seasonal. Most species are annuals, and
typically germinate on recently receded waters
in swampy habitats dominated (in Australia at
least) by Melaleuca spp., but also on
creekbanks, in seepage areas in ranges or on
sandstone plateaux on damp sand. Sites are
typically open and sunny.
11. Paracorolla presence (#50): present, continuous
(0); absent (1); present, discontinuous (2).
12. Paracorolla glands presence (#60): present (0);
absent (1).
13. Labellum position (#63): sheathing the column (0);
attached to outside of corolla tube (1); attached
at base of anterior sinus of corolla tube (2).
14. Fusion pattern of petals (#74): laterally fused (0);
petals all free (1); petals with posterior ones
fused (2).
15. Anterior petals lobing (#77): entire (0); bilobed
(1).
16. Lobing of posterior petals (#81): entire (0); bilobed
(1).
17. Column dilation (#84): conspicuously dilated near
distal end forming pouch for the stigma and
anthers (0); of uniform width throughout (1);
slightly dilated near distal end (2).
18. Column indumentum (#85): glandular-hairy only (0);
glabrous (1).
19. Capsule ribbing (#93): with raised longitudinal ribs
(0); without raised longitudinal ribs (1).
20. Capsule dehiscence (#96): halves coherent distally
(0); halves detaching distally (1). This character
applies to mature, but relatively fresh capsules.
All capsule-halves will eventually detach
distally, through weathering.
21. Seed shape (#98): ellipsoidal (0); globose (1).
22. Seed surface micro-features (#102): colliculate (0);
smooth (1).
The subgenus has no representatives
in south-western Western Australia, and
instead has its stronghold in tropical Australia,
especially the Northern Territory. 24 species
are recorded from the Northern Territory (12
spp. endemic); 13 species are recorded from
Queensland (4 spp. endemic); 11 species are
recorded from Western Australia (3 spp.
endemic); 9 species are recorded from outside
Australia (5 species not occurring in Australia)
Bean, Stylidium subgen. Andersonia
593
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
11
17
21
27
39
41
44
45
47
49
50
60
63
74
77
81
84
85
93
96
98
102
1
Forstera
bellidifolia
2
1
0
1
1
1
1
1
1
1
1
?
?
1
?
0
1
1
1
1
?
?
2
capillare
0
0
0
0
2
2
1
0
2
2
0
1
1
1
1
1
1
1
1
0
0
1
3
schizanthum
0
1
0
0
0&2
2
0
1
2
2
0
0
1
2
1
1
2
1
0&1
1
0&1
0&1
4
rotundifolium
0
1
0
0
2
0
0&1
1
3
2
1
?
2
1
0
0&1
0
1
1
1
0
0
5
diffusum
2
0
1
0
1&2
2
0&1
0&1
3
2
0&2
1
1
1
0&1
1
1
1
1
0
0
0
6
candelabrum
2
1
0
0
1&2
3
1
0&1
3
0
2
1
1
1
0&1
1
1
1
1
1
1
1
7
pedunculatum
1
0
0
1
1
2
0&1
0&1
1&2
1&2
1
?
1
2
0
1
0
0
0
1
0&1
1
8
cordifolium
2
1
1
0
1&2
2
0&1
0
0&3
1
1
?
2
0
0
0
1
1
1
0
0
0
9
longissimum
2
1
1
0
2
2
0
0
3
2
0
1
2
1
1
1
2
1
1
1
0
0
10
trichopodum
2
0
0
1
0
2
0
1
2
2
1&2
1
1
2
0&1
1
0
0
1
1
0
1
11
confertum
2
0
1
0
2
2
0&1
0
3
2
2
1
1
1
1
1
1
0&1
1
0
0
0&1
12
lobuliflorum
0
1
0
0
0
2
0
1
3
2
0
0
1&2
1&2
0&1
1
1
1
1
1
0
1
13
fissilobum
2
0
1
0
2
2
0
0&1
2&3
2
0
0&1
1
1&2
1
1
1
1
1
0
0&1
0&1
14
simulans
0
1
0
0
0
2
0
0&1
1
2
2
1
1
2
0&1
1
1
1
1
1
0
1
15
muscicola
1
1
0
0
0
2
0
1
1
2
1&2
1
2
1
1
1
1
1
1
1
0
1
16
fluminense
2
1
1
0
0
2
1
0
3
1
1
?
2
0
0
0
1
1
1
0
0&1
0
17
alsinoides
2
1
1
0
1&2
2
1
1
1&3
1
1
?
2
0
0
0
1
1
1
0
0
0
18
claytonioides
1
0
0
1
1
0
0
1
3
2
1
?
2
2
0
1
0
1
?
?
?
?
19
tenerum
0
1
0
0
0
2
0
1
1
0
2
1
1&2
1
0
1
1
1
1
1
0
1
20
dunlopianum
2
1
0
0
2
0
0
1
3
2
1
?
2
1
0
0
0
1
1
1
0
0
Table 2 - Matrix of character states. Characters are numbered according to Table 1
58
63
58
64
71
88
Forstera
bellidifolia
capillare
schizanthum
lobuliflorum
rotundifolium
dunlopianum
diffusum
candelabrum
pedunculatum
trichopodum
claytonioides
longissimum
confertum
fissilobum
simulans
muscicola
tenerum
cordifolium
fluminense
alsinoides
Forstera
bellidifolia
capillare
diffusum
longissimum
confertum
fissilobum
schizanthum
lobuliflorum
simulans
muscicola
tenerum
candelabrum
pedunculatum
trichopodum
claytonioides
rotundifolium
dunlopianum
cordifolium
fluminense
alsinoides
Fig. la. Strict Consensus Tree, showing Bootstrap
values.
Fig. lb. 50% Majority-rule Consensus Tree
7 : 1->0
10 : 1->2
594
Austrobaileya 5(4): 589-649 (2000)
Forstera
bellidifolia
capillare
fissilobum
confertum
diffusum
longissimum
schizanthum
lobuliflorum
simulans
muscicola
tenerum
candelabrum
pedunculatum
trichopodum
claytonioides
rotundifolium
dunlopianum
cordifolium
fluminense
alsinoides
Fig. 2. One of 156 most parsimonius trees, showing character numbers and their state changes at each branch
595
Bean, Stylidium subgen. Andersonia
and one species is recorded from New South
Wales. All extra-Australian species of Stylidium
belong to S. subg. Andersonia. None of the 37
species in the subgenus is endemic outside
the tropics. Only three species ( S. tenerum, S.
Taxonomy
diffusum and S. rotundifolium) extend south
of the Tropic of Capricorn, while two ( S. kunthii
and S. tenellum ) extend north of the Tropic of
Cancer.
Key to the subgenera of Stylidium occurring in northern and north-easternAustralia,
Malesia and south-eastAsia
1. Hypanthia and capsules linear, 8-20 times longer than wide. S. subg. Andersonia
Hypanthia and capsules globose to ellipsoidal, 1^4 times longer than wide.2
2. Hypanthia and capsules globose; stigma stipitate.S. subg. Centridium
Hypanthia and capsules ellipsoidal to obconical; stigma sessile .... S. subg. Tolypangium
Key to the species of Stylidium subg .Andersonia
1. Inflorescence borne on leafless scape(s), generally arising from a cluster of
leaves, either basal or cauline; leaves not resembling floral bracts.2
Inflorescences not borne on leafless scape(s); leaves always cauline,
tending to grade into floral bracts.26
2. Scapes bearing 1 flower only.3
Scapes bearing cymose inflorescence, 2-many-flowered.7
3. Calyx fused into two obtuse lips.4
Calyx consisting of 5 lobes, some partly fused.5
4. Scapes and hypanthia glandular-hairy. 17. S. perizostera
Scapes and hypanthia glabrous. 18. S. claytonioides
5. Leaves glabrous; corolla yellow. 14. S. trichopodum
Leaves with eglandular hairs; corolla pink.6
6. Scapes 0.15-0.2 mm wide; posterior petals 2.4-3.1 mm long;
capsules 7-9.5 mm long; sepals partly fused, forming two
emarginate lips. 15. S. ericksoniae
Scapes 0.1-0.15 mm wide; posterior petals 0.5-0.7 mm long;
capsules 5-6.5 mm long; 2 sepals fused for more than half
their length, 3 free. 16. S. pedunculatum
7. Sepals fused into two obtuse lips.8
Sepals consisting of 5 lobes, some or all partly fused.10
8. Paracorolla present, lobes acute; posterior petals c. 5.5 mm long. 32. S. fimbriatum
Paracorolla absent; posterior petals 1.3-2.1 mm long.9
9. Corolla tube 2-3 mm long; column 3.5-5 mm long;
leaves in basal rosette. 30. S. rotundifolium
Corolla tube 4.5-6 mm long; column 7-8.5 mm long;
leaves mostly cauline. 31. S. dunlopianum
596 Austrobaileya 5(4): 589-649 (2000)
10. Sepals in two basally fused groups (one 2-lobed, the other 3-lobed) .... 9. S. candelabrum
Three sepals free to base, two fused for more than half their length.11
11. Leaves linear, 20-35 times longer than wide. 13. S. stenophyllum
Leaves obovate to orbicular, 1-3 times longer than wide .12
12. Leaves in basal rosette.13
Leaves scattered along a substantial stem.14
13. Two orange paracorolla glands present.18
Paracorolla glands absent.20
14. Leaves sessile, orbicular; paracorolla lobes 6-9, linear, acute. 5. S. ensatum
Leaves petiolate, oblanceolate to orbicular; paracorolla lobes 0-4, broad, obtuse.15
15. Orange paracorolla glands present.12. S. pachyrrhizum
Paracorolla glands absent.16
16. Scapes glabrous; corolla glandular-hairy;
posterior petals fused; capsules ribbed . 7. S. divergens
Scapes glandular-hairy; corolla glabrous; all petals free; capsules not ribbed.17
17. Capsules 6.5-11 mm long; native of Asia. 1. S. kunthii
Capsules (10—) 13—24 mm long; native ofN.T. andW.A. 4. S. muscicola
18. Hypanthia densely glandular-hairy; glands pale. 10. S. lobuliflorum
Hypanthia sparsely glandular-hairy or glabrous; glands dark.19
19. Stem base thickened; petals glandular-hairy;
column 7.5-8.5 mm long.12. S. pachyrrhizum
Stem base not thickened; petals glabrous;
column 5-7.5 mm long. 11. S. schizanthum
20. Sepals glabrous, acute.21
Sepals sparsely glandular-hairy, obtuse.22
21 Bracts 2—4.7 mm long; corolla tube 0.5-1.6 mm long;
corolla glandular-hairy on tube and petals;
leaves with petioles 1-4.5 mm long. 19. S. nominatum
Bracts 1-2 mm long; corolla tube 1.6-2 mm long;
corolla glandular-hairy on tube only; leaves sessile. 20. S. capillare
22. Petals all free [A1+A2+P1+P2] .23
Posterior petals fused [A1+A2+(P1&P2)].24
23. Corolla tube with sinus on anterior and posterior sides;
paracorolla present; labellum usually attached to the
outside of the corolla tube; Aust. & N.G. . 3. S. tenerum
Corolla tube with sinus on anterior side only; paracorolla
absent; labellum attached at base of anterior sinus of
corolla tube; Sri Lanka to China. 2. S. uliginosum
24. Corolla tube glabrous.6. S. si mu la ns
Corolla tube glandular-hairy.25
597
Bean, Stylidium subgen. Andersonia
25. Column 2.5-3 mm long; capsule 4—8 mm long; sepals 0.1-0.25 mm wide.8. S. accedens
Column 5.5-7 mm long; capsule 10-17 mm long; sepals 0.25-0.4 mm wide . . 7. S. divergens
26. Leaves 0.1-1.7 mm wide.27
Leaves 1.7-8 mm wide.35
27. Leaf base cuneate.28
Leaf base truncate.29
28. Petals laterally fused (A1 &PI + A2&P2); sepals glabrous. 36. S. javanicum
Petals all free (A1+A2+P1+P2); sepals glandular-hairy. 27. S. tenellum
29. Leaves clustered near base of stem.30
Leaves ±evenly scattered along stem.31
30. Sepals oblanceolate; paracorolla continuous;
posterior petals 3^1.1 mm long.22. S. aquaticum
Sepals debate; paracorolla discontinuous;
posterior petals 1.5-3.3 mm long.25. S. confertum
31. Capsules 1.0-1.6 mm wide, pedicels absent or up to 5 mm long. 35. S. tenerrimum
Capsules 0.4-1.0 mm wide, pedicels absent or rudimentary.32
32. Posterior petals entire; labellum attached at base of anterior sinus.21. S. prophyllum
Posterior petals bilobed; labellum attached to outside of corolla tube.33
33. Anterior petals 0.5-1.0 mm long. 26. S. diffusum
Anterior petals 1.3-2.2mmlong.34
34. Corolla yellow and white; 4-6 paracorolla lobes opposite posterior petals . 24. S. oviflorum
Corolla white or pink; 2-4 paracorolla lobes opposite posterior petals .... 23. S. fissilobum
35. Leaf base cordate.37. S. cordifolium
Leaf base cuneate.36
36. Sepals 1.0-2.0 mm long at anthesis.37
Sepals 2.0^1.1 mm long at anthesis.39
37. Petals laterally fused (A1&P1 +A2&P2); column 2.5-3 mm long.33. S. alsinoides
Petals all free (A1+A2+P1+P2); column 3-5 mm long.38
38. Leaf apex acute; corolla glabrous; column 3-3.5 mm long.29. S. inconspicuum
Leaf apex obtuse; corolla glandular-hairy; column 4-5 mm long. 27. S. tenellum
39. Sepals glandular; capsules 26^18 mm long;
corolla tube 2-2.3 mm long.28. S. longissimum
Sepals glabrous; capsules 11-16 mm long;
corolla tube 1.1-1.5 mm long. 34. S. fluminense
Stylidium subg.Andersonia(R.Br. exGDon) Annual or perennial herbs. Stem base
Mildbr. in Engl., Pflanzenr. 35: 31, 34 unthickened, or rarely thickened. Glandular
(1908); S. sect. AndersoniaRBr. ex GDon, hairs present, but often on one or two plant
Gen. hist. 3: 721 (1834).Type: S. tenellum parts only. Glands globose. Leaves entire; in
R.Br. (=S. tenerum Spreng.) (lecto, here basal rosette, terminal rosette, or spirally
chosen). arranged along elongate stems. Inflorescence
598
Austrobaileya 5(4): 589-649 (2000)
scapose or scapiform, 1-flowered or cymose;
or borne on upper part of stem, cymose. Bracts
opposite, sometimes leaf-like and then
inflorescence appearing to be solitary, axillary.
Labellum glabrous, attached at base of anterior
sinus or on outer wall of corolla tube; labellum
basal appendages absent. Sepals never all free;
often with 2 sepals fused for over half their
length and 3 free, but sometimes fused into
two entire or emarginate lips. Stigma sessile,
cushion-shaped. Column lacking eglandular
hairs, spur absent. Anther loculi 4. Capsules
Key to the Sections of Stylidium subg .Andersonia
1. Inflorescences 1-flowered. S. sect. Uniflora
Inflorescences cymose, 2-many flowered.2
2. Calyx fused into 2 obtuse lips.S. sect. Biloba
Calyx consisting of 5 lobes, some partly fused.3
3 Petals laterally fused (A1 &P1 +A2&P2). S. sect. Alsinoida
Petals all free (A1+A2+P1+P2) or posteriors fused (A1+A2+(P1&P2)).4
4 Inflorescences borne on leafless scape(s),
generally arising from a cluster of leaves
(either basal or cauline). Leaves not resembling floral bracts.S. sect. Andersonia
Inflorescences not borne on leafless scape(s).
Leaves always cauline, tending to grade into floral bracts. S. sect. Tenella
linear or linear-lanceolate, pedicels absent or
rudimentary (except S. tenerrimum). Ovules
numerous. Seeds brown.
37 species, occurring from far northern
New South Wales to tropical Western Australia,
and into Malesia and south-east Asia.
The subgenus is here divided into 5
sections; S. sect. Andersonia, S. sect. Uniflora,
S. sect Tenella, S. sect. Biloba and S. sect.
Alsinoida.
A. Stylidium sect. Andersonia R.Br ex G.Don,
Gen. hist. 3: 721 (1834).
Leaves broad, in basal rosette or scattered on
short stem. Scapes present. 2 sepals fused for
more than half their length and 3 free. Corolla
white, pink or mauve; petals all free or posterior
ones fused to each other. Paracorolla usually
present; orange paracorolla glands present or
absent. Labellum usually attached to outside
of corolla tube. Column of uniform width
throughout, glabrous. Capsules sometimes
ribbed; seeds smooth.
13 species, SE Asia, Malesia, tropical
Australia, southern Queensland, northern New
South Wales.
1.StylidiumkunthiiWall exDC ,Prodr 7:335
(1839). Type: [Bangladesh] Sylhet, [24°—
’ N 91°— 5 E], undated, N. Wallich herb,
no. 3759 (holo: K, microfiche BRI; iso:
BM).
Stylidium brunonis Griff., Not. pi. asiat. 4:
275-276 (1854). Type: Burma. Mergui,
[12°—’ N 98°—’ E], undated, W. Griffith
577 (holo: ?K«.v.; iso: BM).
Annual, 8-20 cm high. Glandular hairs 0.05-0.1
mm long; glands globose, dark. Stem base not
thickened. Stems elongate, glabrous. Leaves 5-
15 per plant, mostly in terminal rosette, with
some scattered along stems, oblanceolate or
obovate or orbicular, 4-15 mm long including
petiole, 2-9 mm wide, glabrous; apex obtuse;
base cuneate; margins entire. Petioles 1-6 mm
long. Scapes 1-5 per plant, 0.4-0.7 mm in
diameter, glandular-hairy; sterile bracts absent
or present. Inflorescences 5-14 cm long,
determinate, monochasially cymose; branches
glandular-hairy. Bracts debate or lanceolate,
1.5-3.5 mm long, glabrous or glandular-hairy,
obtuse. Bracteoles absent. Pedicels absent or
rudimentary. Hypanthium linear, glabrous
throughout or glandular-hairy throughout.
599
Bean, Stylidium subgen. Andersonia
Sepals oblanceolate or elliptical, with 3 free and
2 fused for more than half their length, 1.3-2.5
mm long, 0.3-0.5 mm wide, glandular-hairy,
obtuse. Corolla white or pink, glabrous; tube
1.2-1.6 mm long, with sinus on anterior side
only. Paracorolla discontinuous, thin, glabrous,
0.1-0.2 mm high. Paracorolla lobes or
appendages 2, both similar, acute, none
opposite the anterior petals, 2 opposite the
posterior petals. Paracorolla glands absent.
Labellum attached at base of anterior sinus of
corolla tube, lanceolate, 0.6-0.7 mm long, thick,
glabrous, acute or acuminate, terminal
appendage absent. Petals all free,
A1+A2+P1+P2. Anterior petals 0.7-1.2 mm
long, 0.4—0.8 mm wide, bilobed or entire, obtuse.
Posterior petals 1-2 mm long, 0.6-1.3 mm wide,
bilobed, obtuse. Column 4-5 mm long, of
uniform width throughout, glabrous; lateral
lobes absent. Corona absent. Capsule linear,
6.5-11 mm long excluding sepals, 0.6-0.9 mm
wide, without raised longitudinal ribs; halves
detaching distally, strongly recurved or not
recurved. Seeds ellipsoidal, c. 0.2 mm long,
brown; surface convex, smooth.
Specimens examined: India. Motijharan hill,
Sambalpur, [21°N 84°E], Nov 1948, Mooney 3139
(K); Siliguri, North Bengal, [26°N 88°E], Dec 1875,
Clarke 26567 (BM). Bangladesh. Chittagong, [22°N
91°E], undated, Thomson (L, P). Thailand. Poo
Kradeng, [16°N 101°E], Nov 1958, Sorensen et al.
6181 (L). Burma. Pegu, [17°N 96°E], Dec 1957,
McKee 5837 (NSW, P). Vietnam. Phuc Yen, [21°N
105°E], Nov 1935, Petelot 5517 (P); Ben Cat, [11°N
106°E], Dec 1865, Pierre 4533 (AAU, P). Unplaced.
Cochinchine, 1862-66, Thorel 916 (P).
Reconstituted material examined: Petelot
5517(6 fls); Sorensen etal 6181 (1 fl).
Distribution and habitat: Stylidium kunthii
is endemic to south-east Asia, extending from
eastern India to Vietnam (Map 1). From the
localities recorded, it appears to be confined to
altitudes of less than 200 metres. S. kunthii is
recorded from moist road cuttings in forests,
and on open wet soil in rice fields and natural
grassland.
Phenology: Flowers and capsules have been
recorded for November and December.
Affinities: S. kunthii is closely related to S.
uliginosum, but differs by the cauline leaves
(confined to basal rosette for S. uliginosum ),
labellum 0.6-0.7 mm long (0.4-0.5 mm for S.
uliginosum ), column 4-5 mm long (3-3.5 mm
long for S. uliginosum ), presence of a
paracorolla, and corolla tube 1.2-1.6 mm long
(0.9-1.3 mm for A uliginosum).
At L, there is a mixed collection of S.
kunthii and S. uliginosum from Phu Kradung
in northern Thailand, which indicates that these
species, at least occasionally, grow together.
Typification: The holotype of S. brunonis is
presumably at K. One of the sheets received
on loan to BRI had the note “specimen from
Mergui - Griffith - Herb. Hook. - removed from
here 8 iii 1933”. However the latter was not
received.
Conservation status: Not evaluated.
2. Stylidium uliginosum Sw. exWilld., Sp. PI. 4:
147 (1805); Swartz, Ges. Naturf. Freunde
Berl. Mag. NeuestenEntdeck. Gesammten
Naturk. 1: 52 (1807); Candollea uliginosa
(Sw. ex Willd.) F.Muell., Syst. Census
Aust. PI. 86 (1883). Type: Ceylon, undated,
J.G. Koenig( holo: B, microfiche BRI; iso:
BM).
Stylidium sinicum Hance in Walp., Ann.
Bot. Syst. 2: 1030 (1852). Type: Hong
Kong, [22°—’ N 114°—’] E, undated,
collector unknown (syn: BM).
Annual, 4—15 cm high. Glandular hairs 0.05-0.1
mm long; glands globose, dark. Stem base not
thickened. Stems compressed (with leaves in
basal rosette). Leaves 7-16 per plant, elliptical
or obovate, 4.5-11 mm long including petiole,
2.0^1.5 mm wide, glabrous; apex obtuse; base
cuneate; margins entire. Petioles 0.5^1 mm long.
Scapes 1-7 per plant, 0.4-0.5 mm in diameter,
glabrous or glandular-hairy; sterile bracts
absent or present. Inflorescences 4-15 cm long,
determinate, monochasially cymose; branches
glandular-hairy. Bracts debate or lanceolate or
ovate, 1.5-2 mm long, glabrous or glandular-
hairy, obtuse. Bracteoles absent. Pedicels
absent or rudimentary. Hypanthium linear,
glandular-hairy throughout. Sepals
oblanceolate, with 3 free and 2 fused for more
than half their length, 1.2-2 mm long, 0.3-0.4
mm wide, glandular-hairy, obtuse. Corolla white,
600
glabrous; tube 0.9-1.3 mm long, with sinus on
anterior side only. Paracorolla absent. Labellum
attached at base of anterior sinus of corolla
tube, lanceolate, 0.4-0.5 mm long, thin,
glabrous, acute or acuminate, terminal
appendage absent. Petals all free,
A1+A2+P1+P2. Anterior petals 0.5-0.8 mm
long, 0.3-0.5 mm wide, bilobed or entire, acute
or obtuse. Posterior petals 0.8-1.6 mm long,
0.5-1.1 mm wide, bilobed or entire, obtuse.
Column 3-3.5 mm long, of uniform width
throughout, glabrous; lateral lobes absent.
Corona absent. Capsule linear, 6-8 mm long
excluding sepals, 0.5-0.8 mm wide, without
raised longitudinal ribs; halves detaching
distally, strongly recurved or not recurved.
Seeds ellipsoidal, c. 0.2 mm long, brown; surface
convex, smooth.
Specimens examined: China. Shipai Farm near
Guangzhou, Nov 1955, Huang 160717 (MO); Baiyun
Mountain, near Guangzhou, Oct 1963, Huang
165266 (MO); White Cloud Hills, Canton, 23N 113E,
Mar 1885, Sampson 368 (BM); Port Island, Hong
Kong, Oct 1973, Hu 12255 (K); Hong Kong, 1850,
Champion (BM, K); Hong Kong, [22°N 114°E],
1853—6, Wright (L, P). Thailand. Loei, Phu Krading,
Nov 1958, Smitinand 4928 (K. L); Phu Kradung, S of
Loi, [16° 53’N 101° 53’E], Nov 1970, Chaoenphol
4608 et al. (AAU); Loei, Phu Krading, Nov 1965,
Tagawa 827 et al. (AAU, L); Phukradung, Loei, Dec
1971, Beusekom 4498 et al. (L, MO, P); Pak Thong
Chai, Korat, [14°N 102°E], Dec 1923, Kerr 8112 (BM);
Ubon at Kemarath, Me-Kong, [15°N 104°E], 1866-
8, Thorel (P). Vietnam, near Kip, Bac Giang (Ha
Bac) Province, [21°N 106°E], Jan 1936, Petelot 5662
(P); Hue, Annam, [16°N 107°E], Mar 1927, Squires
186 (P); Baie de Tourane, [16°N 108°E], Feb 1908,
d’Alleizette 24 (P); Vinh-yen, Tonkin, [21°N 105°E],
undated, Eberhardt 3808 (AAU, P); Nha-trang and
vicinity, [12°N 109°E], Mar 1911, Robinson 1057
(P). Cambodia. Kep, Kampot, [10°N 104°E], Oct
1904, Geoffray 449 (P). Sri Lanka, [near Colombo]
Ceylon, [6°N 80°E], 1868, Thwaites 3505 (BM, P);
Ceylon, 1830-37, Walker (L, P). Unplaced. Cu Phap,
Tonkin, Feb 1887, Balansa 3617 (P); Mat Lon,
Tonkin, Feb 1892, Bon (P); Dai Khoi, Tonkin, Feb
1893, Bon (P); Ouonbi, Tonkin, Nov 1885, Balansa
1396 (P).
Reconstituted material examined:
Charoenphol et al. 4608 (3 fls); Petelot 5662 (3
fls).
Distribution and habitat: Stylidium
uliginosum is endemic to south-east Asia. The
type was collected from Sri Lanka, but it may
be extinct there now (Wadhwa 1997). It is
Austrobaileya 5(4): 589-649 (2000)
otherwise found from Thailand to the
Guangdong province of southern China (Map
2). From the localities recorded, it appears to
be confined to altitudes of less than 200 metres.
S. idiginosum has been recorded from sandy
moist places in open savannah, beside a pool
on a dried stream, and on sandy earth banks.
Phenology: Flowers and capsules have been
recorded from October to March.
Notes: S. uliginosum is most closely related to
S. kunthii and S. tenerum. See notes under those
species. Only one type specimen for S. sinicum
was seen, but others may exist. As this name is
a synonym only, I have not attempted to
designate a lectotype for it.
Conservation status: Not evaluated.
3. Stylidium tenerum Spreng., Syst. Veg. 3: 749
(1826)
Stylidium tenellumRBr., Prodr. 570 (1810),
nom. illeg., non Sw. exWilld. (1805). Type:
East Coast Port 1 [Queensland. Port Curtis
District: Curtis Island, 23 0 4-’S 151°1-’E],
6 August 1802, R. Bmwn (lecto: BM), here
chosen.
Illustration: R. Erickson, Triggerplants, plates
51,54 (1958), as S. uliginosum.
[S. uliginosum auct. Australiense non. Sw. ex
Willd.]
Annual, 3-20 cm high. Glandular hairs 0.05-0.2
mm long; glands ellipsoidal or globose, dark.
Stem base not thickened. Stems compressed
(with leaves in basal rosette). Leaves 4-10 per
plant, obovate or orbicular, 4-17.5 mm long
including petiole, 3-8 mm wide, glabrous; apex
obtuse; base obtuse, or cuneate; margins
entire. Petioles 1.5-7 mm long. Scapes 1-7 per
plant, 0.2-0.6 mm in diameter, glabrous or
glandular-hairy; sterile bracts present.
Inflorescences 3-20 cm long, determinate,
monochasially cymose; branches glabrous, or
glandular-hairy. Bracts debate or lanceolate, 1-
1.5 mm long, glabrous or glandular-hairy,
obtuse or acute. Bracteoles absent. Pedicels
absent or rudimentary. Hypanthium linear,
glandular-hairy throughout. Sepals
oblanceolate, with 3 free and 2 fused for more
601
Bean, Stylidium subgen. Andersonia
than half their length, 1-1.6 mm long, 0.3-0.5
mm wide, glandular-hairy, obtuse. Corolla white,
glabrous; tube 1-1.6 mm long, with sinus on
anterior and posterior sides. Paracorolla
discontinuous, thin, glabrous, 0.1-0.3 mm high.
Paracorolla lobes or appendages 4-8, all similar,
obtuse, 2{-A) opposite the anterior petals, 2-4
opposite the posterior petals. Paracorolla
glands absent. Labellum attached to outside
of corolla tube or attached at base of anterior
sinus of corolla tube, ovate, 0.3-0.5 mm long,
thick, glabrous, acute or acuminate, terminal
appendage absent. Petals all free,
A1+A2+P1+P2. Anterior petals 0.3-1.1 mm
long, 0.3-0.8 mm wide, entire, obtuse. Posterior
petals 1.1-1.9 mm long, 0.8-1.5 mm wide,
bilobed, obtuse. Column 2.5-3.5 mm long, of
uniform width throughout, glabrous; lateral
lobes absent. Corona absent. Capsule linear,
5-8 mm long excluding sepals, 0.5-0.8 mm wide,
without raised longitudinal ribs; halves
detaching distally, strongly recurved. Seeds
ellipsoidal, 0.2 mm long, brown; surface convex,
smooth.
Selected specimens: Papua New Guinea. Wuroi,
Oriomo River, [8°S 143°E], Jan-Mar 1934, Brass 5820
(BRI). Northern Territory. Wessel Islands, Oct 1972,
Latz 3512 (DNA). Queensland. Cook District:
Yeldham Creek, Etheridge River, undated, Armit 614
(MEL); Emu Creek, 10.2 km from Peninsula
Development road, towards ‘Dixie’, Jul 1998, Bean
13549 (BRI, DNA); Browns Creek, Pascoe River, Jul
1948, Brass 19640 (BRI, L); Daintree N.P., Little
Daintree River, May 1998, Forster PIF22800 et al.
(BRI, QRS); Walsh River gorge, Sep 1985, Godwin
C2866 (BRI). North Kennedy District: Alligator Creek,
Bowling Green Bay N.P, Jun 1991, Bean 3276 (BRI);
Ten Mile Creek, 22.5 km from Proserpine towards
Bowen, Jul 1997, Bean 12088 (BRI); c. 8 km NE of
“Wairuna”, Aug 1997, Bean 12175 (BRI); Stony
Creek, W of Ingham near Wallaman Falls, Aug 1951,
Blake 18804 (BRI). South Kennedy District: Mount
Mandurana N.P., 20 km NNW of Mackay, May 1991,
Bean 3141 (BRI). Port Curtis District: 2 km from
Tannum Sands, old Tannum road, Dec 1989, Aspland
1032 (BRI); between Pineapple Gap and Raspberry
Creek HS, Shoalwater Bay Military Reserve, Jul 1977,
Clarkson 746 & Stanley (BRI); Deepwater Creek N.P,
about 3 km SW of campsite, Sep 1992, Sharpe 5374
& Tan (BRI). Burnett District: Mt Perry, Jun 1889,
Keys (BRI). Wide Bay District: 1.5 km S of Kingfisher
Bay, Fraser Island, Jul 1997, Bean 12055 (BRI, DNA,
MEL); Granite Bay, Noosa N.P, Sep 1985, Sharpe
3909 & Batianoff (BRI); 2.3 km from Tewantin on
road to Boreen Point, Oct 1988, Wannan & Quinn
(BRI). Moreton District: Wappa Dam Pumping station,
W of Yandina, Sep 1997, Bean 12406 (BRI);
Caloundra, Aug 1932, Blake 4246 (BRI); My ora. North
Stradbroke Island, Aug 1969, Coveny 2018 (BRI, MEL,
NSW). New South Wales. Coolangatta Airport,
Tweed Heads West, Jun 1999, Bean 15226 (BRI, NSW);
Cudgen Creek, 1892, Finselbach 106 (MEL).
Reconstituted or spirit material examined:
Bean 12055 (2 fls); Bean 12088 (2 fls); Bean
12175 (2 fls); Bean 12194 (2 fls); Clarkson 746
& Stanley (2 fls); Forster 5444 (2 fls).
Distribution and habitat: Stylidium tenerum
is widespread in eastern Queensland,
especially in the tropics. It is also known from
Papua-New Guinea, the Wessel Islands in
Northern Territory, and from the extreme north¬
east of New South Wales (Map 4). It grows on
seepage areas or consolidated creekbanks, or
sometimes on coastal lowlands in open
Melaleuca woodland. It grows in sandy soils,
and is often associated with species of
Drosera, Eriocaulon and Utricularia.
Phenology: Flowers and capsules have been
recorded mainly between April and October,
but there are a few records from December and
January.
Notes: S. tenerum is closely related to S.
uliginosum. S. tenerum differs by having a
sinus on both the anterior and posterior sides
of the corolla tube, by the presence of a
paracorolla, and by having the labellum usually
attached to the outside of the corolla tube.
Two specimens were available to Brown
when drawing up his description; his own, and
one collected by Banks and Solander from
Endeavour River. Brown’s collection is selected
as lectotype as he indicates it in his Prodromus
as “(T.) v.v.”, meaning “seen living from
Tropical Australia”. This infers that the
description was based primarily on his own
specimen.
Conservation status: The extent of occurrence
of S. tenerum is very large. Within this area it is
confined to relatively undisturbed creekbanks
or open lowland communities on sand. All
known stands are small and scattered, but S.
tenerum is not considered to be rare or
threatened.
602
4. Stylidium muscicola F.Muell., Fragm. 1:153
(1859); Candollea muscicola (F.Muell.)
F.Muell., Syst. census Austral, pi. 86
(1883). Type: [Northern Territory],
Cataracts, Victoria River, [15°—’S 131°—
’E], 26 June 1856, F. Mueller (holo: MEL
[MEL1061541]).
Illustrations: R. Erickson, Triggerplants Plates
51, 53 (1958); J.R. Wheeler (ed.), FI. ofKimb.
Region 882, t. 272B(1992).
Annual, 5-33 cm high. Glandular hairs 0.1-0.15
mm long; glands globose, dark. Stem base not
thickened. Stems elongate, glabrous. Leaves
4-20 per plant, mostly in terminal rosette, with
some scattered along stems, obovate or
orbicular, (6-) 10-33 mm long including petiole,
(5-)9-28 mm wide, glabrous; apex obtuse; base
obtuse, or cuneate; margins entire. Petioles 5—
29 mm long. Scapes 1-8 per plant, 0.2-0.5 mm
in diameter, glandular-hairy; sterile bracts
present or absent. Inflorescences 3-17 cm long,
determinate, monochasially cymose; branches
glandular-hairy. Bracts linear or lanceolate, 0.5-
2.0 mm long, glabrous or glandular-hairy, acute.
Bracteoles absent. Pedicels absent or
rudimentary. Hypanthium linear, glandular-
hairy throughout. Sepals elliptical, with 3 free
and 2 fused for more than half their length, 1.5-
2 mm long, 0.3-0.7 mm wide, glandular-hairy,
obtuse. Corolla white or pink or mauve,
glabrous; tube 2.1-2.5 mm long, with sinus on
anterior side only. Paracorolla absent or
discontinuous, thin, glabrous, 0.1-0.15 mm
high. Paracorolla lobes or appendages 0^1, all
similar, obtuse, 0-2 opposite the anterior petals,
0-2 opposite the posterior petals. Paracorolla
glands absent. Labellum attached at base of
anterior sinus of corolla tube, lanceolate, 0.4-
0.7 mm long, thick, glabrous, acute or
acuminate, tenninal appendage usually present;
0-0.4 mm long. Petals all free,Al+A2+Pl+P2.
Anterior petals 1.4-1.8 mm long, 0.8-1.6 mm
wide, bilobed, obtuse. Posterior petals 2.3-3.5
mm long, L4-2.2 mm wide, bilobed, obtuse.
Column 5-6.5 mm long, of uniform width
throughout, glabrous; lateral lobes present,
0.1-0.2 mm wide. Corona absent. Capsule linear,
(10—) 13—24 mm long excluding sepals, 0.4-0.6
mm wide, without raised longitudinal ribs;
halves detaching distally, strongly recurved.
Austrobaileya 5(4): 589-649 (2000)
Seeds ellipsoidal, 0.15-0.2 mm long, brown;
surface convex, smooth.
Selected specimens: Western Australia. Lennard
River Gorge, King Leopold Range, Jun 1976,
Beauglehole 52606 (DNA); The Grotto, c. 30 km
SSE of Wyndham, Jun 1976, Beauglehole 54083
(DNA); King Edward River, Amax road crossing, Jun
1971, Byrnes 2327 (DNA); S side of Cockburn Range,
c. 13 km W of King River, Jul 1974, Carr 3286 &
Beauglehole 47064 (DNA); Galvins Gorge, c. 200
km E of Derby, Jul 1974, Carr 4141 & Beauglehole
47919 (DNA); Wonjarring Falls, Carson River
escarpment, Jun 1984, Chesterfield 420 (MEL); Whale
Mouth Cave, Osmond Range, Jul 1991, Cowie 1924
(DNA, PERTH). Northern Territory. Pinkerton
Range, Auvergue Station, Mar 1989, Brock 566 &
Russell-Smith (DNA); near Mt Gilruth, Mar 1984,
Craven 8308 & Wightman (DNA, MEL); Little
Nourlangie Rock, Mar 1978, Dunlop 4773 (CANB,
DNA); Mt Boulder, Feb 1989, Dunlop 7999 & Leach
(DNA); Macadam Range, Mar 1989, Dunlop 8073 &
Leach (DNA); Angalarri River catchment. May 1994,
Dunlop 10042 & Latz (DNA, MEL); 10 km SW of
Redbank Mine, Wollogorang Station, Jun 1987, Latz
10505 (DNA, CANB, MEL, NSW); catchment of
Hayward Creek, Mar 1989, Leach 2480 & Dunlop
(DNA); tributary of Fitzmaurice River, Feb 1994,
Leach 4202 (DNA); Katherine Gorge N.P., May 1977,
Parker 799 (DNA, CANB); Victoria River area, Jasper
Gorge, May 1988, Smith 1128 (DNA); South Bay,
Bickerton Island, Jun 1948, Specht 554 (BRI, MEL);
2 km W of Victoria River bridge, Gregory N.P, Mar
1986, Thomson 1328 (DNA); Vanderlin Island, Sir
Edward Pellew group, Jul 1988, Thomson 2456 (DNA);
Yambarran Range, 19 km NE of Mt Milik Monmir,
May 1994, Walsh 3819 & Leach (DNA, MEL).
Reconstituted or spirit material examined:
Parker 199 (3 fls); Smith 1128 (2 fls).
Distribution and habitat: Stylidium muscicola
extends from the western Kimberley of Western
Australia to northern parts of the Northern
Territory, eastward almost to the Queensland
border (Map 5). It grows in sandy soil in
sheltered places in sandstone ranges (e.g. base
of cliffs), in places which receive seepage after
the wet season.
Phenology: Flowers and capsules have been
recorded between February and September.
Notes: S. muscicola is distinguished by the pale
green petiolate cauline leaves, the white flowers
with free posterior petals, and the relatively long
capsules.
603
Bean, Stylidium subgen. Andersonia
There is only one Mueller collection of
this taxon at MEL, hence it is treated as the
holotype.
Conservation status: A. muscicola is a
widespread species and not considered to be
rare or threatened.
5. Stylidium ensatum ARBeansp. nov. affmis
A. muscicola , sed differens foliis
sessilibus, scapis latioribus, bracteis
obtusis, corolla glandulosa et faucis
appendicibus ensatis 6-9. Typus:
Northern Territory. Shoal Bay, 12°22’S
130°57’E, 20 June 1974, C.R. Dunlop 4105
(holo: BRI; iso: CANB, DNA, K, L, NSW).
Annual, 14-22 cm high. Glandular hairs 0.05-
0.1 mm long; glands globose, dark. Stem base
not thickened. Stems elongate, glabrous.
Leaves 6-17 per plant, scattered along stems,
obovate or orbicular, 7-12 mm long, 5.5-12 mm
wide, glabrous; apex obtuse; base cuneate;
margins entire. Petioles absent. Scapes 1-3 per
plant, 0.6-0.8 mm in diameter, glandular-hairy;
sterile bracts absent. Inflorescences 8-16 cm
long, determinate, monochasially cymose;
branches glandular-hairy. Bracts lanceolate or
ovate, 1-2.0 mm long, glandular-hairy, obtuse.
Bracteoles absent. Pedicels absent or
rudimentary. Hypanthium linear, glandular-
hairy throughout. Sepals oblanceolate, with 3
free and 2 fused for more than half their length,
1.8- 2.1 mm long, 0.4-0.6 mm wide, glandular-
hairy, obtuse. Corolla pink or mauve, glandular-
hairy on tube and petals; tube 1.9-2.4 mm long,
with sinus on anterior side only. Paracorolla
continuous, thin, glabrous, 0.2-0.5 mm high.
Paracorolla lobes or appendages 6-9, all similar,
acute, 2-4 opposite the anterior petals, 4-5
opposite the posterior petals. Paracorolla
glands absent. Labellum attached to outside
of corolla tube, lanceolate, 0.4-0.5 mm long,
thin, glabrous, acute or acuminate, terminal
appendage usually present; 0-0.2 mm long.
Petals all free, A1+A2+P1+P2. Anterior petals
1.8- 2.5 mm long, 1.2-2 mm wide, bilobed,
obtuse. Posterior petals 3^1.4 mm long, 1.8-3.3
mm wide, bilobed, obtuse. Column 5.5-7.5 mm
long, of uniform width throughout, glabrous;
lateral lobes present, 0.1-0.2 mm wide. Corona
absent. Capsule linear, 8-12 mm long excluding
sepals, 0.5-0.7 mm wide, without raised
longitudinal ribs; halves detaching distally,
strongly recurved. Seeds ellipsoidal, c. 0.2 mm
long, brown; surface lacunose, smooth. Fig.
3A.
Specimens examined: Northern Territory, near
Hayes Creek, Jul 1946, Blake 16409 (BRI, DNA);
Port Darwin, 1885, Holtze 514 (MEL); Port Darwin,
1888, Holtze 827 (MEL); Port Darwin, 1888, Holtze
831 (MEL); Port Darwin, 1888, Holtze 865 (MEL);
Lee Point road, 7.5 miles [12 km] N of Darwin, Jun
1964, Nelson 1159 (DNA); North Australia, 1886,
Tenison-Woods & Holtze (MEL); edge of McMinns
Lagoon, Jun 1968, Wheelwright DW44 (DNA).
Reconstituted or spirit material examined:
Dunlop 4105 (3 fls); Nelson 1159 (1 fl);
Wheelwright DW 44 (1 fl).
Distribution and habitat: Stylidium ensatum
is endemic to the Darwin area of the Northern
Territory (Map 5). For one collection, the habitat
is given as a Melaleuca viridiflora-
Lophostemon lactifluus forest, with damp peaty
soil.
Phenology: Flowers and capsules have been
recorded from June and July.
Affinities: S. ensatum is most closely related
to S. muscicola , but differs by the sessile
leaves, scapes 0.6-0.8 mm wide (0.2-0.5 mm for
S. muscicola ), bracts obtuse (acute for S.
muscicola ), corolla glandular-hairy (glabrous
for S. muscicola ), paracorolla lobes acute
(obtuse or absent for A muscicola) and labellum
attached to outside of corolla tube (attached at
base of anterior sinus for S. muscicola ).
Notes: This taxon was mentioned by Erickson
(1958) in her discussion of the variation in
5. muscicola.
Conservation status: Data deficient (IUCN
1994)
Etymology: The species epithet is from the
Latin ensatus, meaning “sword-like”. This is in
reference to the slender sword-like throat
appendages, which are a feature of this species.
6. Stylidium simulans Carlquist, Aliso 9: 426
(1979). Type: Northern Territory. Camp 2,
Arnhem Land, 13°04’S 133°01’E, 8 June
1978, A. Carlquist 15435 (holo: RSA; iso:
DNA).
604
Annual, 7-11 cm high. Glandular hairs 0.05-0.1
mm long; glands globose, pale. Stem base not
thickened. Stems compressed (with leaves in
basal rosette). Leaves 4-10 per plant, orbicular,
3-5.5 mm long including petiole, 1.5-3.5 mm
wide, glabrous; apex obtuse; base cuneate;
margins entire. Petioles 1-2 mm long. Scapes 1
or 2 per plant, 0.2-0.3 mm in diameter, glandular-
hairy; sterile bracts present. Inflorescences 7-
11 cm long, determinate, monochasially
cymose; branches glandular-hairy. Bracts
deltate, 0.5-1 mm long, glandular-hairy, obtuse
or acute. Bracteoles absent. Pedicels absent or
rudimentary. Hypanthium linear, glandular-
hairy throughout. Sepals oblanceolate or
elliptical, with 3 free and 2 fused for more than
half their length, 1.4-1.6 mm long, 0.2-0.3 mm
wide, glandular-hairy, obtuse or acute. Corolla
mauve or pale pink, glabrous; tube 1.6-2 mm
long, with sinus on anterior side only.
Paracorolla discontinuous, thin, glabrous, 0.2-
0.4 mm high. Paracorolla lobes or appendages
6-8, all similar, acute, 2-4 opposite the anterior
petals, 4 opposite the posterior petals.
Paracorolla glands absent. Labellum attached
to outside of corolla tube, lanceolate, 0.3-0.4
mm long, thick, glabrous, acute or acuminate;
0.1-0.2 mm long. Petals with posterior ones
fused, A1+A2+(P1&P2). Anterior petals 1.1-
1.3 mm long, 0.4—0.7 mm wide, bilobed or entire,
obtuse. Posterior petals 1.7-2.2 mm long, 0.7-
1.2 mm wide, bilobed, obtuse. Column 3.5^1
mm long, of uniform width throughout,
glabrous; lateral lobes absent. Corona absent.
Capsule linear, 5.5-8.5 mm long excluding
sepals, 0.4-0.5 mm wide, without raised
longitudinal ribs; halves detaching distally,
strongly recurved or not recurved. Seeds
ellipsoidal, 0.15-0.2 mm long, brown; surface
convex, smooth.
Specimens examined: Northern Territory. Mt
Gilruth area, Jun 1978, Dunlop 4910 (DNA); Arnhem
Land, Mt Gilruth area, Jun 1978, Henshall 1887B
(DNA); ditto, Henshall 1887A (DNA).
Reconstituted or spirit material examined:
Dunlop 4910 (3 fls).
Distribution and habitat: Stylidium simulans
is endemic to the Arnhem Land plateau, and is
currently known only from around the type
locality (Map 7), where it grows on a sandstone
plateau in shallow sandy soil.
Austrobaileya 5(4): 589-649 (2000)
Phenology: Flowers and capsules have been
recorded in June only.
Affinities: S. simulans is closely related to
S. accedens. See notes under that species.
Typification: The type of S. simulans was
incorrectly cited as Carlquist 15434 in the
protologue (Carlquist, pers. comm.). The type
collection for S. simulans is Carlquist 15435;
this is the collection received and annotated
as holotype from RSA.
Conservation status: Data deficient (IUCN
1994).
7. Stylidium divergensARBeansp. nov. affinis
S. accedenti sed differens scapis glabris,
sepalis latioribus longioribusque, labello
ad sinum tubi corollae affixo, columna
multo longiore et capsulis 10-17 mm
longis. Typus: Northern Territory. Mt
Brockman Outlier, 15 km SE of Jabiru, 12°
47’S 132° 57 5 E, 19 April 1989, R IE Johnson
4635 (holo: BRI; iso: AD, DNA, CANB,
RSA).
Annual, 7-27 cm high. Glandular hairs 0.05-
0.15 mm long; glands globose, dark. Stem base
not thickened. Stems elongate, glabrous or
leaves in basal rosette. Leaves 2-6 per plant,
elliptical, obovate or orbicular, 4-21 mm long
including petiole, 3-5 mm wide, glabrous; apex
obtuse; base obtuse or cuneate; margins entire.
Petioles 0.5-8 mm long. Scapes 1-6 per plant,
0.3-0.6 mm in diameter, glabrous; sterile bracts
absent. Inflorescences 4-24 cm long,
determinate, monochasially cymose; branches
glabrous or glandular-hairy. Bracts deltate or
ovate, 0.5-1 mm long, glabrous, acute.
Bracteoles absent. Pedicels absent or
rudimentary. Hypanthium linear, glandular-
hairy throughout or at distal end only. Sepals
oblanceolate, with 3 free and 2 fused for more
than half their length, 1.2-1.7 mm long, 0.25-
0.4 mm wide, glabrous or glandular-hairy,
obtuse. Corolla white or mauve, glandular-hairy
on tube and petals; tube 1.5-2.4 mm long, with
sinus on anterior side only. Paracorolla
discontinuous or continuous, thin, glabrous,
0.1-0.5 mm high. Paracorolla lobes or
appendages 2-4, all similar, obtuse, 2 opposite
the anterior petals, 0-2 opposite the posterior
605
Bean, Stylidium subgen. Andersonia
petals. Paracorolla glands absent. Labellum
attached at base of anterior sinus of corolla
tube, ovate, 0.4-0.5 mm long, thick, glabrous,
acute or acuminate, terminal appendage usually
present; 0-0.5 mm long. Petals with posterior
ones fused, A1+A2+(P1&P2). Anterior petals
1.3-2 mm long, 0.5-1.6 mm wide, bilobed,
obtuse. Posterior petals 2.5-3.7 mm long, 2-3.5
mm wide, bilobed, obtuse. Column 5.5-7 mm
long, of uniform width throughout, glabrous;
lateral lobes absent or present, 0-0.2 mm wide.
Corona absent. Capsule linear, 10-17 mm long
excluding sepals, 0.5-0.6 mm wide, with raised
longitudinal ribs; halves detaching distally, not
recurved. Seeds ellipsoidal, 0.25-0.3 mm long,
brown; surface convex, colliculate. Fig 3B-D.
Specimens examined: Northern Territory, near
Kurundie Creek, Kakadu National Park, Apr. 1990,
Leach 2842 & Cowie (DNA).
Reconstituted or spirit material examined:
Johnson 4635 (2 fls); Leach 2842 & Cowie (3
fls).
Distribution and habitat: Stylidium divergens
is endemic to the Kakadu National Park in
Northern Territory (Map 4). It occurs on
sandstone slopes or gullies.
Phenology: Flowers and capsules have been
recorded only for April.
Affinities: S. divergens is closely related to S.
accedens, but differs by the glabrous scapes
(glandular-hairy for S. accedens ); labellum
attached to base of anterior sinus of corolla
tube (attached to outside of tube for S.
accedens ); much larger posterior petals, column
5.5-7 mm long (2.5-3 mm long for S. accedens )
and capsules 10-17 mm long (4-8 mm long for
S. accedens).
Conservation status: Data deficient (IUCN
1994).
Etymology: From the Latin divergens meaning
diverging or separating. This is in reference to
the widely spreading posterior petals.
8. Stylidium accedens A . R. Bean sp. nov. afFmis
S. simulanti , sed differens tubo corollae
glanduloso, paracorolla absens
continuave, labello breviore, absentia
appendicis terminalis in labello, petalis
anticis et posticis brevioribus, columna
2.5-3 mm longa et seminibus colliculatis.
Typus: Northern Territory. Katherine
Gorge National Park, 14°19’S 132°25 , E,23
June 1975, C.R. Dunlop 3797 (holo: BRI;
iso: CANB, DNA).
Annual, 4-15 cm high. Glandular hairs 0.05-0.1
mm long; glands globose, pale. Stem base not
thickened. Stems compressed (with leaves in
basal rosette). Leaves 4-10 per plant, obovate,
5-12 mm long including petiole, 2.5-5.5 mm
wide, glabrous; apex obtuse; base cuneate;
margins entire. Petioles 1.5-3.5 mm long. Scapes
1-2 per plant, 0.2-0.4 mm in diameter, glandular-
hairy; sterile bracts present. Inflorescences 4-
15 cm long, determinate, monochasially
cymose; branches glandular-hairy. Bracts linear
or lanceolate, 0.5-1 mm long, glandular-hairy,
acute. Bracteoles absent. Pedicels absent or
rudimentary. Hypanthium linear, glandular-
hairy throughout. Sepals oblanceolate, with 3
free and 2 fused for more than half their length,
1.2-1.4 mm long, 0.1-0.25 mm wide, glandular-
hairy, obtuse. Corolla white, glandular-hairy on
tube only; tube 1.2-1.7 mm long, with sinus on
anterior side only. Paracorolla absent or
continuous, thin, glabrous, 0.1 mm high.
Paracorolla lobes (when present) 6, all similar,
obtuse, 2 opposite the anterior petals, 4
opposite the posterior petals. Paracorolla
glands absent. Labellum attached to outside
of corolla tube, ovate or lanceolate, 0.3-0.4 mm
long, thick, glabrous, acuminate, terminal
appendage absent. Petals with posterior ones
fused, A1+A2+(P1&P2). Anterior petals 0.5-
0.7 mm long, 0.4-0.5 mm wide, bilobed, obtuse.
Posterior petals 0.6-1 mm long, 0.5-0.8 mm
wide, bilobed, obtuse. Column 2.5-3 mm long,
of uniform width throughout, glabrous; lateral
lobes absent. Corona absent. Capsule linear,
(4-)6-8 mm long excluding sepals, 0.3-0.5 mm
wide, without raised longitudinal ribs; halves
detaching distally, not recurved. Seeds
ellipsoidal, 0.2 mm long, brown; surface convex,
colliculate. Fig. 3E-G
Specimens examined: Northern Territory. Edith
Falls, Edith River, 17 miles [27 km] by road E of
Stuart Highway, Jul 1965, Beauglehole ACB10878
(DNA); Edith Falls, Jul 1965, Beauglehole ACB43215
(DNA); Arnhem Land plateau, Jun 1978, Carlquist
15445 (BRI); Katherine Gorge N.P., Jun 1975, Dunlop
3760 (CANB, DNA); upper Katherine River, Arnhem
606
Land, Jul 1996, Dunlop 10303 & Mangion (DNA);
above UDP Falls, Jul 1971, Gittins 2637 (BRI); sources
of Goomadeer R., Arnhem Land, Jun 1978, Henshall
1993 (DNA).
Reconstituted or spirit material examined:
Dunlop3760 (3 fls ),Dunlop 10303 & Mangion
(2 fls).
Distribution and habitat: Stylidium accedens
occurs around and to the north-east of
Katherine in Northern Territory (Map 6), where
it grows on sandstone plateaux with shallow
sandy soil.
Phenology: Flowers and capsules have been
recorded for June and July
Affinities: S. accedens is closely related to S.
simulans, but differs by the obovate leaves
(orbicular for S. simulans ), corolla tube
glandular-hairy (glabrous for S. simulans ),
paracorolla absent or continuous
(discontinuous for S. simulans ), the shorter
labellum, the absence of a terminal appendage
on the labellum, the shorter anterior and
posterior petals, column 2.5-3 mm long (3.5-4
mm for S. simulans ), and minutely colliculate
seeds (smooth for S. simulans).
Conservation status: Data deficient (IUCN
1994).
Etymology: The specific epithet is from the
Latin accedens , an indeclinable participle
meaning “resembling”. This is in reference to
the superficial resemblance of this species to
S. uliginosum, with which it was confused in
the past.
9. Stylidium candelabrum Lowrie & Kenneally,
Nuytsia 13: 251 (1999). Type: Northern
Territory. Headwaters Katherine River,
Arnhemland, 13°47’S 133°05’E, 11 July
1996, C.P.Mangion 245 & C.R.Dunlop
(holo: DNA; iso: PERTH).
Annual, 6-20 cm high. Glandular hairs 0.05-0.1
mm long; glands globose, dark. Stem base not
thickened. Stems elongate, glabrous. Leaves
11-100 per plant, scattered along stems,
elliptical, 2.5—13(—18) mm long including petiole,
1.5—5(—9) mm wide, glabrous; apex obtuse; base
obtuse, or cuneate; margins entire. Petioles 2-
6(-8) mm long. Scapes 1-13 per plant, 0.2-0.7
Austrobaileya 5(4): 589-649 (2000)
mm in diameter, glabrous; sterile bracts absent.
Inflorescences 3-16 cm long, determinate,
monochasially cymose; branches glabrous, or
glandular-hairy. Bracts debate or lanceolate,
0.5-1.5 mm long, glabrous, obtuse or acute.
Bracteoles absent. Pedicels absent or
rudimentary. Hypanthium linear, glabrous
throughout or glandular-hairy at distal end
only. Sepals debate or ovate, fused into 2
emarginate lips, 0.8-1.4 mm long, 0.3-0.4 mm
wide, glabrous, obtuse or acute. Corolla white,
glandular-hairy on tube and petals; tube 1.2-
1.8 mm long, with sinus on anterior and
posterior sides. Paracorolla discontinuous,
thin, glabrous, 0.2-0.4 mm high. Paracorolla
lobes or appendages 4, all similar, obtuse, 2
opposite the anterior petals, 2 opposite the
posterior petals. Paracorolla glands absent.
Labellum attached to outside of corolla tube,
ovate or lanceolate, 0.4-0.6 mm long, thick,
glabrous, acute or acuminate, terminal
appendage absent. Petals all free,
A1+A2+P1+P2. Anterior petals 0.9-1.3 mm
long, 0.6-1.1 mm wide, bilobed, trilobed or
entire, obtuse. Posterior petals 1.6-2.3 mm long,
1.5-2.3 mm wide, bilobed, or 3 or 4-lobed,
obtuse. Column 5.5-6.5 mm long, of uniform
width throughout, glabrous; lateral lobes
absent. Corona absent. Capsule linear, 6-15 mm
long excluding sepals, 0.4-0.7 mm wide, without
raised longitudinal ribs; halves detaching
distally, not recurved. Seeds globose, 0.2-0.25
mm long, brown; surface convex, smooth. Fig.
4A-D.
Selected specimens: Northern Territory, near Kub-
O-Wer Hill, Arnhem Land reserve, Jun 1978,
Carlquist 15408 (DNA, K); near Ranford Creek,
Wandie Mine road, Mary River station, Jul 1987, Clark
807 (DNA); Melville Island, near Snake Bay airstrip
plantation. Mar 1994, Cowie 4698 (DNA); Bathurst
Island, Runka, Mar 1995, Cowie 5419 (DNA); Flinders
Peninsula, east side opposite S end Probable Isle, Apr
1996, Cowie 6698 & Bokarra (DNA); 7.5 km S of
Cooinda on Pine Creek road. May 1980, Craven 5656
(CANB, DNA); Mt Gilruth area, Jun 1978, Dunlop
4885 (DNA, MEL); Kakadu N.P, Apr 1990, Dunlop
8577 & Munns (CANB, DNA); Mt Brockman, Kakadu
N.P., Mar 1995, Egan 4591 (DNA); road to
Umbrawarra Gorge, Apr 1995, Egan 4643 (DNA);
above U.D.P. Falls, May 1975, Gittins 2864 (BRI);
Holmes Jungle, Apr 1995, Harwood 100 (DNA);
Arnhem Land, 19 km E of Jabiru, Apr 1989, Johnson
4554 (BRI, DNA, NSW); above Twin Falls, Kakadu
N.P, Jul 1983, King 324 (DNA); Rum Bottle Creek,
Bean, Stylidium subgen. Andersonia
607
Fig. 3. A: Stylidium ensatum. A. habitx0.7. B-D: Stylidium divergens. B. habit* 1; C. anterior view of flower*9;
D. corolla, opened outx9. E-G: Stylidium accedens. E. habit><2; F. anterior view of flowerxlS; G. corolla,
opened outxl8. A: Holtze 514; B-D: Johnson 4635; E-G; Dunlop 3797.
608
Jun 1972, Must 1070 (BRI, CANB, DNA, K); Marrakai
Station, Mar 1974, Must 1197 (BRI, DNA); 10 km
SSW of ElSharana, Kakadu N.P., Apr 1990, Orr 354
(DNA); Koongarra area, 1 km S of jump-up, Apr 1979,
Rankin 1979 (CANB, DNA, L); Upper Koolpin Creek,
Kakadu N.R, Jun 1988, Russell-Smith 5488 & Lucas
(DNA); Ngilipitji area, E Arnhem Land, Jul 1981,
Scarlett 406 (DNA); 8 km E of Goyder River crossing,
Jun 1972, Symon 7732 (DNA).
Reconstituted or spirit material examined:
Dunlop 4885 (3 fls );Egan 4591 (2 fls); Wightman
3906 (2 fls).
Distribution and habitat: Stylidium
candelabrum is endemic to the northernmost
parts of the NorthemTerritory (Map 11). It often
grows in shallow sand on sandstone
pavements, but may occur on other substrates.
It appears to be confined to the higher rainfall
areas.
Phenology: Flowers and capsules have been
recorded from March to July.
Affinities: S. candelabrum is a distinctive
species readily identified by its petiolate,
elliptical cauline leaves and sepals in 2 basally
fused groups. It differs from S. capillare by its
petiolate cauline leaves, sepals in 2 basally
fused groups, glandular-hairy petals, sinus on
both the anterior and posterior sides of the
corolla, and the discontinuous paracorolla.
Conservation status: S. candelabrum is quite
widely distributed in the north of the Northern
Territory, and much of its occurrence is within
National Park. It is not considered to be rare or
threatened.
10. Stylidium lobuliflorum F.Muell., Fragm. 1:
153 (1859); Candollea lobuliflora
(F.Muell.) F.Muell, Syst. census Austral,
pi. 86 (1883). Type: [NorthemTerritory].
sandy flats near the Main Camp, Victoria
River, [15° —’S 130° —’E], May 1856,
F. Mueller (lecto: MEL [MEL242996]),
here chosen.
Illustration: R. Erickson, Triggerplants, plate
54 (1958), as S. lobuliferum.
Annual, 12-20 cm high. Glandular hairs 0.1-0.2
mm long; glands globose, pale. Stem base not
thickened. Stems compressed (with leaves in
Austrobaileya 5(4): 589-649 (2000)
basal rosette). Leaves 4-15 per plant, obovate
or orbicular, 6-9.5 mm long including petiole,
4-6.5 mm wide, glabrous; apex obtuse; base
cuneate; margins entire. Petioles 1-4 mm long.
Scapes 1-3 per plant, 0.4-0.7 mm in diameter,
glandular-hairy; sterile bracts absent.
Inflorescences 12-20 cm long, determinate,
monochasially cymose; branches glandular-
hairy. Bracts linear or lanceolate or ovate, 1-2
mm long, glandular-hairy, acute. Bracteoles
absent. Pedicels absent or rudimentary.
Hypanthium linear, glandular-hairy throughout.
Sepals oblanceolate, with 3 free and 2 fused for
more than half their length, 2.3-3 mm long, 0.4-
0.5 mm wide, glandular-hairy, obtuse. Corolla
pink or mauve, glandular-hairy on tube and
petals; tube 2.1-3.3 mm long, with sinus on
anterior side only. Paracorolla continuous, thin
or thick, glabrous, 0.2-0.6 mm high. Paracorolla
lobes or appendages 2—4, all similar, obtuse, 0-
2 opposite the anterior petals, 2 opposite the
posterior petals. Paracorolla glands 2. Labellum
attached to outside of corolla tube or attached
at base of anterior sinus of corolla tube, ovate
or lanceolate, 0.4-0.6 mm long, thick, glabrous,
acuminate or obtuse, terminal appendage
usually present; c. 0.2 mm long. Petals all free,
A1+A2+P1+P2 or with posterior ones fused,
A1 +A2+(P 1 &P2). Anterior petals 1.4-1.9 mm
long, 0.7-1.1 mm wide, bilobed or entire, obtuse.
Posterior petals 2.3-3.4 mm long, 1.2-2.2 mm
wide, bilobed, obtuse. Column 6.5-7.5 mm long,
of uniform width throughout, glabrous; lateral
lobes absent. Corona absent. Capsule linear,
10-12 mm long excluding sepals, 0.6-0.8 mm
wide, without raised longitudinal ribs; halves
detaching distally, not recurved. Seeds
ellipsoidal, 0.15-0.2 mm long, brown; surface
convex, smooth.
Specimens examined: Western Australia, c. 32 km
WSW of Kununurra, Jul 1976, Beauglehole 54297
(DNA); 59 km by road N of Gibb River road, Meda-
Oobagooma road, Jun 1976, Beauglehole 52687
(DNA). Northern Territory. 8 km west of Koongarra,
May 1978, Carlquist 15392 (DNA); Wilderness Trail,
Katherine Gorge N.R, 8 km from Park Headquarters,
Jun 1978, Carlquist 15470 (BRI, DNA); Plot 426,
Arnhem Land, [12° 09’S 133° 55’E], Jul 1987, Clark
1327 (DNA, NSW); tributary of Barramundie Ck, 33
km WSW of Twin Falls, Jun 1980, Craven 6419
(CANB, DNA); Kakadu N.P., Apr 1990, Dunlop 8560
& Munns (DNA); Fitzmaurice River basin. May 1994,
Dunlop 9950 & Late (DNA, NSW); Fitzmaurice River,
upper catchment. May 1994, Dunlop 10084 & Barritt
609
Bean, Stylidium subgen. Andersonia
(BRI, DNA, MEL); Arnhem Land, Mt Gilruth area,
Jun 1978, Henshall 1878 (DNA, MEL); headwaters
of Baroalba Creek, Kakadu Park, May 1981, Henshall
3604 (DNA); 12.5 km SW of Jabiru on Pine Creek
road, Jun 1980, Lazarides 9177 (BRI, CANB, DNA,
MEL); Mt Bundey Training area, Jul 1992, Leach
2972 (BRI, CANB, DNA); Yamburran Range, 19 km
NE Mt Millikmonmir, May 1994, Leach 4514 &
Walsh (DNA, MEL); Litchfield Park, road to Lost
City, Jun 1998, Michell 1583 & Risler (DNA); Magela
Creek, May 1978, Rice 2641 (DNA); Arnhem Land,
13.1 km E of Murgenella Settlement along road to
Brogden Point, Aug 1987, Ross 3273 (MEL); Bamboo
Ck, tribuatry of McKinley, 1882, Tate (MEL).
Reconstituted or spirit material examined:
Dunlop 10084 & Barritt (2 fls); Leach 2972 (2
fls); Michell 1583 & Risler (2 fls).
Distribution and habitat: Stylidium
lobuliflorum is found in the Kimberley region
of Western Australia and adjacent areas of
Northern Territory (Map 8). It inhabits
sandstone pavement, sandy depressions,
seepage areas or streamlines. It may be
associated with Pandanus spp., Triodia sp.,
Melaleuca viridiflora or various Eucalyptus
spp.
Phenology: Flowers and capsules have been
recorded from April to August.
Typification: There are two collections of S.
lobuliflorum at MEL which were made by
Mueller from the vicinity of the Victoria River.
One collection has several relatively intact
plants (chosen here as lectotype) while in the
other, only fragments remain.
Affinities: S. lobuliflorum is most closely
related to S. schizanthum , but differs by
densely glandular hypanthia with pale glands
(glabrous or sparsely glandular with dark
glands for S. schizanthum ), absence of sterile
bracts on scapes, petals glandular-hairy
(glabrous for S. schizanthum ), column of
uniform width throughout (slightly dilated for
S. schizanthum ), corolla tube 2.1—3.3 mm long
(1.3-2.2 mm for S. schizanthum) and 2-4
paracorolla lobes (4-6 for S. schizanthum).
Conservation status: S. lobuliflorum is a
widespread species, and not considered to be
rare or threatened.
11. Stylidium schizanthum F.Muell., Fragm. 1:
152 (1859); Candollea schizantha
(F.Muell.) F.Muell., Syst. census Austral,
pi. 86 (1883). Type: Victoria River
[Northern Territory, 15°—’S 130°—’E],
May 1856, F. Mueller (holo: MEL
[MEL1061485]).
Illustration: R. Erickson, Triggerplants, plates
51, 54 (1958); J.R. Wheeler (ed.), FI. ofKimb.
Region 882, t. 272E(1992).
Annual, 9-30 cm high. Glandular hairs 0.05-0.2
mm long; glands globose, dark. Stem base not
thickened. Stems compressed (with leaves in
basal rosette). Leaves 3-13 per plant,
oblanceolate or obovate or orbicular, 3.5-23 mm
long including petiole, 1.5-12 mm wide,
glabrous; apex obtuse; base cuneate; margins
entire. Petioles 0.5-3(-6) mm long. Scapes 1-
2(^1) per plant, 0.3-0.8 mm in diameter, glabrous
or glandular-hairy; sterile bracts present.
Inflorescences 9-30 cm long, determinate,
monochasially cymose; branches glabrous, or
glandular-hairy. Bracts debate or lanceolate or
ovate, 1-2.5 mm long, glabrous or glandular-
hairy, obtuse or acute. Bracteoles absent.
Pedicels absent or rudimentary. Hypanthium
linear, glandular-hairy at distal end only or
glandular-hairy throughout. Sepals
oblanceolate, with 3 free and 2 fused for more
than half their length, 1.5-2.7 mm long, 0.4-0.7
mm wide, glandular-hairy, obtuse. Corolla white
or pink or mauve or yellow, glandular-hairy on
tube only; tube 1.3-2.2 mm long, with sinus on
anterior side only. Paracorolla continuous, thin,
glabrous, 0.3-1 mm high. Paracorolla lobes or
appendages 4-6, dimorphic or all similar, acute
or obtuse, 2 opposite the anterior petals, 2^1
opposite the posterior petals. Paracorolla
glands 2. Labellum attached to outside of
corolla tube, ovate or orbicular, 0.3-0.6 mm long,
thick, glabrous, acuminate; 0.1-0.2 mm long.
Petals with posterior ones fused,
A1+A2+(P1&P2). Anterior petals 0.7-2.7 mm
long, 0.5-1.4 mm wide, bilobed, acute or obtuse.
Posterior petals 2.1-4.9 mm long, 1.6-3.1 mm
wide, bilobed, obtuse. Column 5-7.5 mm long,
slightly dilated near distal end, glabrous; lateral
lobes absent. Corona absent. Capsule linear,
8-15 mm long excluding sepals, 0.5-0.8 mm
wide, without raised longitudinal ribs or with
raised longitudinal ribs; halves detaching
610
distally, strongly recurved or not recurved.
Seeds globose or ellipsoidal, 0.15-0.25 mm
long, brown; surface convex, smooth or
colliculate.
Selected specimens: Irian Jaya. c. 15 km NE of
Koembe village on N bank of Koembe River, Merauke
district, [8°S 140°E], Sep 1954, van Royen 4854 (L).
New Guinea. Wuroi, Oriomo River, Western
Division, [8°S 143°E], Jan-Mar 1934, Brass 5796,
5797 (BRI); Lake Daviumbu, Middle Fly River, [7°S
141°E], Sep 1936, Brass 7825 (L); c. 1 mile [1.6 km]
S of Morehead Patrol Post, Western District [9°S
141°E], Aug 1967, Pullen 7159 (L); near Weam,
Western District, [8°S 141°E], Aug 1967, Ridsdale
NGF33681 (L). Western Australia. Mitchell
Plateau, near mining camp, Aug 1978, Beauglehole
59004 & Errey 2704 (DNA); King Edward River, c.
50 km NE of Mitchell River HS, Aug 1978,
Beauglehole 58879 & Errey 2579 (DNA); Gibb River-
Kalumburu Mission road, 11.3 km NNW of Drysdale
River crossing. May 1976, Beauglehole 51707 (DNA);
Gibb River road, 1.5 km W of Lennard River Gorge
turnoff, Jul 1974, Carr 4063 & Beauglehole 47841
(DNA); Mount Parker plateau, Jul 1991, Cowie 1933
(DNA); 6 km W of Mitchell River Falls, Apr 1993,
Cowie 4343 & Brubaker (CANB, DNA); Camp Ck,
next to Amax campsite, Mitchell Plateau, May 1978,
Kenneally 6638 (L); Flying Fox Creek, SW of Lake
Argyle, May 1980, Weston 12215 (DNA). Northern
Territory. Malabanbandju camping ground. Kakadu
N.P, May 1995, Booth 902 (DNA); McArthur River
area, c. 15 km N of Borroloola, Jun 1976, Craven
4228 (L); Katherine Gorge N.P, Jun 1975, Dunlop
3788 (DNA, NSW); Fitzmaurice River basin. May
1994, Dunlop 9940 & Latz (DNA, MEL); Winchelsea
Island, May 1993, Egan 2461 (DNA); Vanderlin Island,
Jul 1988, Latz 10820 (DNA); on Oenpelli road, c.
10.5 miles [16.8 km] from Mudginbarry HS, Jul 1972,
Lazarides 7591 (BRI, DNA); Bickerton Island, South
Bay, May 1993, Leach 3723 & Cowie (DNA); Deaf
Adder Basm, Jun 1972, Schodde AE97 (BRI, CANB,
DNA, K, L); Magela Creek, Mudginberri Stn, May
1982, Wightman 48 (CANB, DNA). Queensland.
Burke District: “Esmeralda” about SSE of Croydon,
Jul 1954, Blake 19651 (BRI); 12 miles [19 km] N of
Esmeralda station, Jul 1954, Speck 4732 (BRI). Cook
District: One Hundred Mile swamp, near Undara resort,
E of Mt Surprise, Jul 1998, Bean 13762 & Fox (BRI);
Pascoe River crossing. Iron Range-Wenlock road, Jul
1948, Brass 19663 (BRI, L); Archer River, Wenlock-
Coen road, Jul 1948, Brass 19736 (BRI, L); Moa
Island, Aug 1985, Budworth 178 (BRI); 1.8 km N of
the track from Agnew to Bertiehaugh, Aug 1983,
Clarkson 5000 (BRI, K, NSW, PERTH, QRS); 33 km
from Wakooka on the track to Bathurst Bay and
Cape Melville N.P., Jun 1984, Clarkson 5379 (BRI,
DNA, L, QRS); Cowal Creek crossing on the Bamaga
to Mutee Head road, Sep 1985, Clarkson 6209 (BRI,
DNA, MBA, PERTH, QRS); 9.7 km N of Morehead
River, Jun 1989, Clarkson 8077 (BRI); Pascoe River
Austrobaileya 5(4): 589-649 (2000)
at Wattle Hills, Aug 1991, Clarkson 9087 & Neldner
(BRI, MBA, PERTH); c. 5 km ENE of the mouth of
MacDonald River, c. 68 km WNW of Heathlands,
Apr 1993, Clarkson 9875 & Neldner (BRI, DNA,
MBA, PERTH); 6 km N of Lilyvale on the track to
Running Creek, Jun 1993, Clarkson 10095 & Neldner
(BRI, MBA, PERTH); 29.3 km E by road of Maloneys
Springs, Jun 1989, Forster 5448 (BRI); 2 km N of Mt
Tozer, Jul 1991, Forster PIF9007 (BRI); 4 miles [6
km] N of Hopevale mission, Sep 1970, Gittins 2183
(BRI); along Kennedy road, c. 2 km S of Moreton
Telegraph station, Aug 1978, Kanis 2036 (BRI,
CANB, L); 23 km NE of “Violet Vale” HS, Aug 1978,
Paijmans 2912 (BRI); c. 60 miles [97 km] W of
Musgrave Telegraph Office, Jun 1968, Pedley 2679
(BRI).
Reconstituted or spirit material examined:
Booth 902 (2 fls); Clarkson 5379 (2 fls);
Clarkson 8077 (2 fls); Clarkson 9087 &
Neldner (2 fls); Clarkson 9875 & Neldner (2
fls); Clarkson 10095 & Neldner (2 fls); Egan
2461 (2 fls ), Forster 5448 (2 fls); Wightman 48
(3 fls).
Distribution and habitat: Stylidium
schizanthum is a very widespread species,
extending from the Kimberley region of
Western Australia to eastern Northern Territory,
and then in north Queensland, as far south as
Mt Surprise, and in southern New Guinea (Map
9). It grows in damp sand in Eucalyptus or
Melaleuca dominated communities, along
creekbanks, on sandstone escarpments and
amongst sandstone boulders.
Phenology: Flowers and capsules have been
recorded from February to October.
Affinities: It is closely related to S.
pachyrrhizum and S. lobuliflorum. See notes
under those species.
Notes: S. schizanthum as currently
circumscribed, is a very variable taxon, and
detailed field studies may reveal that more than
one species is involved.
There is only one Mueller collection at MEL,
hence this collection is treated as a holotype.
Conservation status: S. schizanthum is a
widespread species, and not considered to be
rare or threatened.
611
Bean, Stylidium subgen. Andersonia
12. Stylidium pachyrrhizum F.Muell., Fragm.
1: 152 (1859); Candollea pachyrrhiza
(F.Muell.) F.Muell., Syst. census Austral,
pi. 86 (1883), nom. illeg., non (Steud.)
Benth. (1863). Type: [NorthemTerritory],
between Macadam Range and
Providence Hill, [14° —’S 129°—’E],
October 1855, F. Mueller (holo: MEL
[MEL1061539]).
Illustration: R. Erickson, Triggerplants, plate
55 (1958); J.R Wheeler (ed.), FI. ofKimb. Region
882, t. 2720(1992).
Perennial, 15-40 cm high. Glandular hairs 0.05-
0.15 mm long; glands globose, dark. Stem base
thickened. Stems elongate, glabrous. Leaves
5-30 per plant, scattered along stems,
oblanceolate or obovate, 13—55(—100) mm long
including petiole, 3.5-18(-24) mm wide,
glabrous; apex obtuse; base cuneate; margins
entire. Petioles 3—13(—35) mm long. Scapes 1-6
per plant, 0.3-1 mm in diameter, glabrous; sterile
bracts absent or present. Inflorescences 13-33
cm long, determinate, monochasially cymose;
branches glabrous, or glandular-hairy. Bracts
debate or ovate, 1-1.5 mm long, glabrous or
glandular-hairy, acute. Bracteoles absent.
Pedicels absent or rudimentary. Hypanthium
linear, glabrous throughout or glandular-hairy
throughout. Sepals oblanceolate, with 3 free
and 2 fused for more than half their length, 2-
2.5 mm long, 0.5 mm wide, glandular-hairy,
obtuse. Corolla white or pink, glandular-hairy
on tube and petals; tube 1.8-2 mm long, with
sinus on anterior side only. Paracorolla
continuous, thin, glabrous, 0.2-0.5 mm high.
Paracorolla lobes or appendages 4, all similar,
obtuse, 2 opposite the anterior petals, 2
opposite the posterior petals. Paracorolla
glands 2. Labellum attached to outside of
corolla tube, ovate, 0.5-0.6 mm long, thick,
glabrous, acute or acuminate, terminal
appendage usually present; 0-0.2 mm long.
Petals with posterior ones fused,
A 1+A2+(P1&P2). Anterior petals 1.2-1.3 mm
long, 0.6-0.8 mm wide, bilobed, acute or obtuse.
Posterior petals 2.5-3.5 mm long, 1.7-2.2 mm
wide, bilobed, obtuse. Column 7.5-8.5 mm long,
slightly dilated near distal end, glabrous; lateral
lobes absent; column extension present, c. 0.5
mm long. Corona absent. Capsule linear, 9-13
mm long excluding sepals, 0.4-0.7 mm wide,
with raised longitudinal ribs; halves detaching
distally, strongly recurved. Seeds globose, 0.2-
0.25 mm long, brown; surface convex, smooth.
Specimens examined: Western Australia. 30 km
W of Drysdale River crossing, Jun 1976, Beauglehole
52227 (DNA); Prince Regent River, 1891, Bradshaw
(MEL); Hidden Island, Buccaneer Archipelago, Jun
1982, Kenneally 8372 (BRI). Northern Territory.
E of Fogg Bay, Aug 1946, Blake 16796 (BRI); tributary
of Allia Ck, May 1994, Cowie 4876 & Albrecht (DNA,
MEL); Goose Creek road, Melville Island, Jun 1987,
Clark 1252 (DNA); sandstone plateau, [12° 40’S 133°
15’E], Mar 1973, Craven 2467 (BRI, DNA); Keep
River N.P., Feb 1981, Dunlop 5780 (DNA, MEL);
Moyle River headwaters. May 1994, Dunlop 9826 &
Latz (DNA); headwaters of Cui-eci Creek, May 1994,
Cowie 4913 & Albrecht (DNA, MEL); Fitzmaurice
River basin. May 1994, Dunlop 9983 & Latz (DNA,
MEL); Eva Valley station. Mar 1991, Evans 3672
(BRI, DNA); Waterfall Creek Falls, Kakadu N.P, Apr
1992, Halford Q1167 (BRI); Melville Island, Apr
1986, Johnson 4143 (BRI); Sculthorpe Pound, Aug
1985, Latz 10172 (DNA); “The Pines”, Douglas River,
Oct 1974, Parker 517 (DNA); Kakadu N.P, 10.5 km
NE of Mt Evelyn, Apr 1990, Slee & Craven 2575
(MEL); Oenpelli, Oct 1948, Specht 1313 (MEL); 35
km E of Goyder River crossing, Jun 1972, Symon
7733 (DNA, K, L); Wallaby Beach, Gove, Jun 1972,
Symon 7779 (DNA); Angurugu River, Groote Eylandt,
Oct 1976, Waddy 601 (DNA); beside Allia Creek, Daly
River Aboriginal Reserve, May 1994, Walsh 3659
(DNA).
Reconstituted or spirit material examined:
Cowie 4876 & Albrecht (2 fls); Evans 3672 (2
fls).
Distribution and habitat: Stylidium
pachyrrhizum is found from the western
Kimberley to Groote Eylandt off the east coast
of Northern Territory (Map 10). It inhabits
swamp edges, or seepage areas near sandy or
rocky creeks. There is one record from a vertical
sandstone rockface.
Phenology: Flowers and capsules have been
recorded from February to October, with most
records from March to June.
Notes: S. pachyrrhizum is very close to S.
schizanthum , and some specimens are difficult
to place. S. pachyrrhizum has a thickened stem
base, glandular-hairy petals, column 7.5-8.5 mm
long (5-7.5 mm for S. schizanthum ) and
spathulate leaves scattered along a
considerable stem (obovate to orbicular leaves
612
in basal rosette for S. schizanthum). It is also
close to S. stenophyllum, see notes under that
species.
There is only one Mueller collection at MEL,
hence this collection is treated as the holotype.
Conservation status: S. pachyrrhizum is a
widespread species, and not considered to be
rare or threatened.
13. Stylidium stenophyllum A. R. Bean sp. nov.
affinis Stylidio pachyrrhizo sed differens
caulorhizanon incrassata, foliis linearibus
1.4-2.6 mm latis, petalis glabris, anticis
longioribus, columna ubique uniformiter
lata, capsulis non costatis et seminibus
ellipsoidalibus. Typus: Northern Territory.
Caranbirini Conservation Park, SW of
Borroloola, 7 June 1999 ,A.R. Bean 15066
(holo: BRI; iso: CANB, DNA, MEL, NSW,
PERTH).
Annual, 30-40 cm high. Glandular hairs 0.05-
0.2 mm long; glands globose, dark. Stem base
not thickened. Stems elongate, glabrous.
Leaves 7-25 per plant, scattered along stems,
linear, 36-86 mm long including petiole, 1.5-2.5
mm wide, glabrous; apex obtuse; base cuneate;
margins entire. Petioles 15-50 mm long. Scapes
3-4 per plant, 0.4-0.7 mm in diameter, glabrous;
sterile bracts absent or present. Inflorescences
28-39 cm long, determinate, monochasially
cymose; branches glabrous. Bracts linear or
deltate, 1-4 mm long, glandular-hairy or
glabrous, acute. Bracteoles absent. Pedicels
absent or rudimentary. Hypanthium linear,
glabrous throughout or glandular-hairy
throughout. Sepals oblanceolate, with 3 free
and 2 fused for more than half their length, 2.2-
3.3 mm long, 0.4-0.6 mm wide, glandular-hairy,
obtuse. Corolla white or pink, glandular-hairy
on tube only; tube 2-2.3 mm long, with sinus
on anterior side only. Paracorolla continuous,
thin, glabrous, 0.1-0.4 mm high. Paracorolla
lobes or appendages 2-4, all similar, obtuse, 0-
2 opposite the anterior petals, 2 opposite the
posterior petals. Paracorolla glands 2. Labellum
attached to outside of corolla tube, ovate, 0.5-
0.7 mm long, thick, glabrous, obtuse or acute,
terminal appendage absent. Petals with
posterior ones fused, A1+A2+(P1&P2).
Anterior petals 1.2-2.0 mm long, 0.6-1.1 mm
Austrobaileya 5(4): 589-649 (2000)
wide, bilobed, acute. Posterior petals 2.9-4.3
mm long, 2-3.4 mm wide, bilobed, obtuse.
Column 7-8 mm long, of uniform width
throughout, glabrous; lateral lobes absent;
column extension present, c. 0.5 mm long.
Corona absent. Capsule linear, 9-13 mm long
excluding sepals, 0.5-0.7 mm wide, without
raised longitudinal ribs; halves detaching
distally, not or strongly recurved. Seeds
ellipsoidal, 0.15-0.25 mm long, brown; surface
convex, smooth. Fig. 4E-G
Specimens examined: Northern Territory, near
Caranbirini waterhole, McArthur River area, Jul 1977,
Craven 4678 (BRI, CANB, MEL).
Reconstituted or spirit material examined:
Bean 15066 (2 fls); Craven 4678 (3 fls).
Distribution and habitat: Stylidium
stenophyllum is known only from the type
locality in far north-eastern Northern Territory
(Map 10). It grows in crevices on a deeply
dissected “beehive” sandstone formation.
Phenology: Flowers and capsules have been
recorded for June and July.
Affinities: S. stenophyllum is most closely
related to S. pachyrrhizum , but differs by the
non-thickened stem base, linear leaves 1.4-2.6
mm wide, glabrous petals (glandular-hairy for
S. pachyrrhizum ), unribbed capsules (ribbed
for S. pachyrrhizum ) and ellipsoidal seeds
(globose for S. pachyrrhizum).
Notes: The type locality of S. stenophyllum is
home to another endemic plant, Calytrix
mimiana Craven (Craven 1980). The latter, while
similarly restricted in distribution, occurs on
nearly all the beehive formations at the site.
Conservation status: The habitat of S.
stenophyllum is very specialised, being eroded
sandstone monoliths often known as
“beehive” formations, because of their dome¬
like shape. S. stenophyllum is known from only
one outcrop at the type locality, and the total
number of plant clumps is less than 100.
However, each clump may comprise up to 10
individual plants. A Conservation status of
“Vulnerable” is recommended, based on the
IUCN Criterion D.
613
Bean, Stylidium subgen. Andersonia
Etymology: From the Greek stenos- narrow and
phyllon- leaf, in reference to the very narrow
leaves of this species compared to its closest
relatives.
B. Stylidium sect. Uniflora A.R.Bean sect,
nov. Folia linearia usque anguste deltata;
flores in scapis solitarii; glandulae
minutae globosae; bracteolae praesentes
vel absentes; paracorolla pro parte
maxima absens; petala postica
coadunata, biloba, antica integra;
columna dilatata marsupium formans ubi
stigma et antherae requiescunt; semina
laevia. Typus: S. pedunculatum R.Br.
Flowers solitary on scapes; glands tiny,
globose; bracteoles present or absent; leaves
linear to narrowly debate; paracorolla mostly
absent; posterior petals fused, bilobed; anterior
petals entire; column dilated forming pouch
where stigma and anthers rest, glandular-hairy
or glabrous; seeds smooth.
5 species; tropical Australia and the Aru
Islands.
14. Stylidium trichopodumF.Muell.,Fragm. 10:
86 (1876); Candollea trichopoda
(F.Muell.)F.Muell., Syst. census Austral,
pi. 86 (1883). Type: Queensland. Cook
District: Etheridge River, [18°— 9 S 143°—
’ E], undated, W.Armit 622 (syn: BM).
Annual, 6-20 cm high. Glandular hairs 0.05-0.1
mm long; glands globose, dark. Stem base not
thickened. Stems elongate, glabrous. Leaves
20-200 per plant, scattered along stems, linear,
3-9 mm long, 0.2-0.7 mm wide, glabrous; apex
obtuse, or acute; base truncate; margins entire,
or crenulate. Petioles absent. Scapes 1-10 per
plant, 0.15-0.2 mm in diameter, glandular-hairy;
sterile bracts absent. Inflorescences 2.6-6 cm
long, 1-flowered. Bracteoles present, 0.8-1.2 mm
long. Pedicels absent or rudimentary.
Hypanthium linear, glandular-hairy throughout.
Sepals oblanceolate or ovate or elliptical, with
3 free and 2 fused for more than half their length,
1-2 mm long, 0.5-0.6 mm wide, glandular-hairy,
obtuse. Corolla yellow or orange, glandular-
hairy on tube only; tube 1.4-2.3 mm long, with
sinus on anterior side only. Paracorolla absent
or discontinuous, thin, glabrous, 0.2-0.4 mm
high. Paracorolla lobes or appendages 0-2, all
similar, obtuse, 0 opposite the anterior petals,
0-2 opposite the posterior petals. Paracorolla
glands absent. Labellum attached to outside
of corolla tube, ovate or lanceolate, 0.4-1 mm
long, thick, glabrous, acute or acuminate,
terminal appendage absent. Petals with
posterior ones fused, A1+A2+(P1&P2).
Anterior petals 1-1.7 mm long, 0.6-1 mm wide,
bilobed or entire, obtuse. Posterior petals 2.7-
3.9 mm long, 1.9-2.5 mm wide, bilobed, obtuse.
Column 5-6 mm long, conspicuously dilated
near distal end forming a pouch for the stigma
and anthers, glandular-hairy; lateral lobes
absent. Corona absent. Capsule linear, 10-15
mm long excluding sepals, 0.5 mm wide, without
raised longitudinal ribs; halves detaching
distally, not recurved. Seeds ellipsoidal, 0.2-
0.25 mm long, brown; surface lacunose, smooth.
Fig 5A-C.
Specimens examined: Queensland. Cook District:
3.4 km S of Musgrave, on road to Laura, Jul 1998,
Bean 13564 (BRI, DNA, MEL, NSW); California
Creek road, 10 km from gorge, Jul 1987, Champion
286 (BRI); 4.8 km N of Kennedy River crossing, on
Peninsula Development road, Jun 1981, Clarkson
3683 (BRI); Bulleringa NP, 80 km NW of Mt Surprise,
Red River track past Donkey Spring, Apr 1998,
Forster 22617 & Booth (BRI, DNA, MEL).
Reconstituted or spirit material examined:
Bean 13564 (2 fls); Champion 286 (2 fls);
Clarkson 3683 (1 ^Forster 22617 (2 fls).
Distribution and habitat: Stylidium
trichopodum is known from a just a few small
populations in northern Queensland (Map 14).
It grows on gentle slopes or flats in moist sandy
soil, sometimes in communities dominated by
Melaleuca spp.
Phenology: Flowers have been recorded for
June and July.
Notes: S. trichopodum is clearly related to S.
pedunculatum , as both have solitary flowers
on filiform peduncles and a pouched column
which holds the resting anthers and stigma. It
is readily distinguished however by the
conspicuous stem and glabrous cauline leaves,
and the much larger yellow corolla. There may
be other type material to be found of this species
and hence no lectotype has been chosen at
this stage.
614
Austrobaileya 5(4): 589-649 (2000)
Fig. 4. A-D: Stylidium candelabrum. A. habit* 1; B. sepals removed from hypanthium, showing the 2 fused
groups* 18; C. posterior view of flower, with column removed*9; D. anterior view of flower and hypanthium*9.
E-G: Stylidium stenophyllum. E. habit, showing leaves and part of inflorescence*0.5; F. anterior view of
flower*9; G corolla, opened out*9. A-D: Wightman 3906; E-G: Craven 4678.
615
Bean, Stylidium subgen. Andersonia
Conservation status: Data deficient (IUCN
1994). S. trichopodum appears to be quite rare.
About 110 years elapsed between the type
collection and the next known collection.
15. Stylidium ericksoniae J.H. Willis, Victorian
Naturalist 73: 43 (1956), as 6 ericksonae
Stylidium pedunculatum var. ericksoniae
(J.H. Willis) Carlquist,Aliso 9:322 (1979).
Stylidium androsaceum O. Schwarz, Repert.
Spec. Nov. RegniVeg. 24:105 (1927 \nom.
illeg., wowLindl. (1839)«ecDC. (1839).
Type: Northern Territory. Koolpinyah, 12°
2-’S 131° l-’E, undated, F.A.K. Bleeser
411 (holo: B, destroyed); between RAAF
emergency landing strip and creek, 40 km
S of Darwin, [12° 4- S 131° 0—’E], 18 June
1978,5. Carlquist 15145 (neo: DNA), here
chosen.
Illustration: R. Erickson, Triggerplants, plates
51, 55 (1958), as S. pedunculatum.
Annual, 6-15 cm high. Glandular hairs 0.025-
0.05 mm long; glands globose, dark. Stem base
not thickened. Stems elongate, with simple
hairs. Leaves 20-100 per plant, mostly in
terminal rosette, with some scattered along
stems, linear or debate, 4.0-9.5 mm long, 0.6-
1.1 mm wide, with eglandular hairs; apex acute,
or mucronate; base truncate; margins entire.
Petioles absent. Scapes 1-20 per plant, 0.15-
0.2 mm in diameter, glabrous or glandular-hairy;
sterile bracts absent. Inflorescences 5-11 cm
long, 1-flowered. Bracteoles present, 0.3-1.2 mm
long. Pedicels absent or rudimentary.
Hypanthium linear, glabrous throughout or
glandular-hairy at distal end only. Sepals
oblanceolate or ovate, fused into 2 emarginate
lips, 0.8-1.3 mm long, 0.3-0.5 mm wide,
glandular-hairy, obtuse. Corolla pink or mauve,
glandular-hairy on tube only; tube 2-2.5 mm
long, with sinus on anterior side only.
Paracorolla absent. Labellum attached to
outside of corolla tube, ovate or lanceolate, 0.3-
0.5 mm long, thick, glabrous, acute or
acuminate, terminal appendage absent. Petals
with posterior ones fused, A1+A2+(P1&P2).
Anterior petals 1-1.7 mm long, 0.7-1.3 mm wide,
bilobed or entire, obtuse. Posterior petals 2.4-
3.1 mm long, 1.4-2.1 mm wide, bilobed, obtuse.
Column 5.5-7 mm long, conspicuously dilated
near distal end forming a pouch for the stigma
and anthers, glandular-hairy; lateral lobes
absent. Corona absent. Capsule linear, 7-9.5
mm long excluding sepals, 0.6-0.8 mm wide,
with raised longitudinal ribs; halves detaching
distally, not recurved. Seeds globose, 0.15-0.2
mm long, brown; surface lacunose, smooth.
Specimens examined: Northern Territory: near
Grove Hill, about 13° 28’S and 131° 35’E, Jul 1946,
Blake 16371 (BRI); between RAAF emergency landing
strip and creek, 40 km S of Darwin, Jun 1978,
Carlquist 15145 (DNA); road heading W off Stuart
Highway, opposite juncture of Shoal Bay road and
Stuart Highway, Jun 1977, Carlquist 15188 (DNA);
13 miles [21 km] SE of Darwin, May 1958,
Chippendale 4433 (DNA, MEL); Arnhem Land, upper
Magela Ck catchment, Apr 1995, Cowie 5601 &
Brennan (DNA); 15 km NNE of Jabiru East, Mar
1981, Craven 6555 (DNA, MEL); Koongarra Jump-
up, May 1978, Dunlop 4848 (DNA); Kakadu N.P.,
Apr 1990, Dunlop 8593 & Munns (DNA); Kakadu
N.P., Mount Brockman, Mar 1995, Egan 4571 (DNA,
MEL); Howard Springs, May 1995, Egan 4987 (DNA);
20 miles [32 km] from Darwin on Stuart Highway, Jul
1955, Erickson s.n. (MEL); Adelaide River, 1890,
Holtze 1174 (MEL); Arnhem Land, 19 km E of Jabiru,
Apr 1989, Johnson 4560 (BRI); near Koongarra
Saddle, 1.5 km N of Koongarra, May 1980, Lazarides
8897 (DNA, MEL); 8 miles [13 km] NW of Humpty
Doo, Jun 1972, McKean B569 (DNA); 1 mile [1.6
km] SE of McMinns Lagoon, Aug 1971, Must 761
(DNA); Koongarra area, 1 km S of jump-up, Apr 1979,
Rankin 1992 (CANB, DNA, K); 1 km N of Elizabeth
R., Stuart Hwy, May 1980, Rankin 2373 (DNA);
Kakadu N.P, Upper Koolpin Creek, Jun 1988, Russell-
Smith 5487 & Lucas (DNA).
Reconstituted or spirit material examined:
Blake 16371 (2 fls);Dw«/op4848 (2 fls ),Egan
4987(3 11s).
Distribution and habitat: Stylidium
ericksoniae is confined to northern parts of
the Northern Territory (Map 13). It grows in
damp sand amongst sandstone rocks
(sometimes with Micraira spp.) or on broad
flat swampy drainage channels with sandy soil
frequently with Pandanus spp.
Phenology: Flowers and capsules have been
recorded from March to August.
Affinities: S. ericksoniae may be distinguished
from S. pedunculatum on dried material by its
scapes 0.15-0.2 mm across (0.1-0.15 mm for S.
pedunculatum ), capsules 7-9.5 mm long (5-
616
Austrobaileya 5(4): 589-649 (2000)
Fig. 5. A-C: Stylidium trichopodum. A. habitx0.9; B. anterior view of flowerx9; C. corolla, opened outx9. A-
B: Clarkson 3683; C: Bean 13564.
617
Bean, Stylidium subgen. Andersonia
6.5 mm for S. pedunculatum), and the sepals
fused into two emarginate lips (3 free and 2
fused for more than half their length for S.
pedunculatum). Its floral details are markedly
different from S. pedunculatum, with both the
posterior and anterior petals being much larger
and the column much longer in S. ericksoniae.
Notes: Two distinct forms of S. ericksoniae are
recognisable: plants from the lowland habitats
near Darwin have densely hairy leaves with
acute apices, while plants from the sandstone
escarpments of Kakadu N.P. have sparsely
hairy leaves with mucronate apices. However
the other characteristics appear to be identical,
and hence no taxonomic distinction has been
made. No isotypes of S. androsaceum
O.Schwarz are known (McKee 1963), hence a
neotype has been selected to fix the application
of the name.
Conservation status: Data deficient (IUCN
1994).
16. Stylidium pedunculatum R.Br., Prodr. 571
(1810); Candollea pedunculata (R.Br.)
F.Muell., Syst. census Austral, pi. 86
(1883), nom. illeg., non DC. (1817). Type:
[Queensland. Cook District :] Endeavour
River [15°2-’S 145°1-’E, June-July 1770],
J. Banks & D. Solander (holo: ?BM n.v.;
iso: L).
Stylidium bryoides F.Muell., Fragm. 6: 91
(1867). Type: Queensland. North
Kennedy District: Rockingham Bay,
[18°—’S 146°—’E], undated,/. Dali achy,
n.v.
Stylidium cur turn Carlquist, Aliso 9: 421
(1979),syn. nov. Type: Northern Territory.
Wilderness trail, Katherine Gorge, 6.8 km
from Park headquarters, [ 14°2- S 132°2-
’E], 24 June 1978,5. Carlquist 15473 (holo:
RSA; iso: BRI).
Illustrations: Carlquist (1979: 422), as S.
curtum, Britten (1901:170C)
Annual, 5-10 cm high. Glandular hairs 0.025-
0.05 mm long; glands globose, dark. Stem base
not thickened. Stems elongate, with simple
hairs. Leaves 20-200 per plant, mostly in
terminal rosette, with some scattered along
stems, linear or debate, 4.5-8.5 mm long, 0.5-
0.8 mm wide, with eglandular hairs; apex
acuminate, or mucronate; base truncate;
margins entire. Petioles absent. Scapes 2-14
per plant, 0.1-0.15 mm in diameter, glabrous;
sterile bracts absent. Inflorescences 4-7 cm
long, 1-flowered. Bracteoles present, 0.4—0.6 mm
long. Pedicels absent or rudimentary.
Hypanthium linear, glabrous throughout.
Sepals oblanceolate or elliptical, with 3 free and
2 fused for more than half their length, 0.8-1
mm long, 0.2-0.4 mm wide, glabrous or
glandular-hairy, obtuse or acute. Corolla white
or pink, glabrous or glandular-hairy on tube
only; tube 0.8-1 mm long, without sinus or with
sinus on anterior side only. Paracorolla absent.
Labellum attached to outside of corolla tube or
attached at top of corolla tube, ovate or
lanceolate, 0.3-0.4 mm long, thick, glabrous,
acute or acuminate, terminal appendage absent.
Petals with posterior ones fused,
A 1+A2+(P1&P2). Anterior petals 0.2-0.5 mm
long, 0.2-0.4 mm wide, entire, obtuse. Posterior
petals 0.5-0.7 mm long, 0.3-0.6 mm wide,
bilobed, obtuse. Column 3-3.5 mm long,
conspicuously dilated near distal end forming
a pouch for the stigma and anthers, glandular-
hairy; lateral lobes absent. Corona absent.
Capsule linear, 5-6.5 mm long excluding sepals,
0.5-0.7 mm wide, with raised longitudinal ribs;
halves detaching distally, strongly recurved or
not recurved. Seeds globose or ellipsoidal,
0.15-0.2 mm long, brown; surface convex,
smooth.
Specimens examined: Moluccas, between Kp.
Meroor and Selarin, P. Trangan, Aru Islands, Jul 1938,
Buwalda 5534 (L). Northern Territory. U.D.P Falls,
Arnhemland, Jul 1971, van Balgooy 1379 & Byrnes
(L); south of mouth of Daly River, Jul 1946, Blake
16548A (BRI); between Lily Pond trail and Smith’s
Rock trail, Katherine Gorge N.P, Jun 1978, Carlquist
15473 (BRI); Little Nourlangie Rock, Mar 1978,
Dunlop 4794 (DNA); 23 miles [37 km] from Darwin
on Stuart Highway, Jul 1955, Erickson s.n. (MEL);
Galiwinku, Elcho Island, Jul 1975, Latz 6185 (DNA);
Banjo Beach, Melville Island, Jul 1967, Stocker 334
(BRI, DNA); 81 km NNW of turnoff junction to
Maningreda, Jun 1972, Symon 7856 (DNA); Rum
Bottle Creek, Jun 1972, Symon 7943 (DNA).
Queensland. Cook District: Peninsula Development
road, 3.3 km N of the Archer River, Jul 1998, Bean
13625 (BRI, DNA); 14.6 km N of ‘Starcke’, N of
Cooktown, Jul 1998, Bean 13715 (BRI, DNA, MEL,
NSW); 71.6 km by road NNW of Coen Post Office,
Sep 1975, Coveny 7063 & Hind (BRI, NSW); c. 60
miles [100 km] W of Musgrave Telegraph Office, Jun
618
1968, Pedley 2678 (BRI); Cape York, 2.5 km S of
lodge on Bamaga road, Jul 1985, Thiele 921 (CANB,
MEL). North Kennedy District: Rockingham Bay, 1869,
Dali achy (MEL).
Reconstituted or spirit material examined:
Bean 13625 (2 fis),Bean 13715 (2 fls )\Coveny
7063 & Hind(2 fls).
Distribution and habitat: Stylidium
pedunculatum is found in the scattered areas
of tropical Queensland and Northern Territory,
as well as in theAru Islands west of New Guinea
(Map 12). It grows in damp sand in open
Melaleuca viridiflora communities with genera
such as Utricularia, Drosera and Schoenus.
Phenology: Flowers and capsules have been
recorded from March to September.
Affinities: S. pedunculatum is most closely
related to S. ericksoniae. See affinities under
that species.
Notes: The description and illustrations of ‘ S.
pedunculatum ’ provided by Erickson (1958) are
referrable to S. ericksoniae. S. pedunculatum ,
as to type, was not described therein. No doubt
this prompted Carlquist (1979) to describe S.
curtum as a new species, when in fact
specimens he identified as S. curtum match S.
pedunculatum in every respect.
Typification: Only one gathering of S.
pedunculatum was available to Brown when
he drew up his description, that made by Banks
and Solander. Hence the BM sheet, which was
not sent on loan has been treated as the
holotype, and the Leiden sheet is therefore an
isotype.
No indisputable type of S. bryoides is
present at MEL or K, but the detailed description
allows its placement as a synonym of S.
pedunculatum. Furthermore there are two
‘topotypes’ collected by Dallachy, and these
are clearly S. pedunculatum.
The type of S. curtum was incorrectly
cited as Carlquist 15467 in the protologue
(Carlquist, pers. comm.). The type collection
for S. curtum is Carlquist 15473; this is the
collection received and annotated as type from
RSA.
Conservation status: Data deficient (IUCN
1994).
Austrobaileya 5(4): 589-649 (2000)
17. Stylidium perizostera Lowrie & Kenneally,
Nuytsia 11(2): 208 (1997). Type: Western
Australia, near Roe River mouth, c. 4 km
north-west of Mount Brookes, Mitchell
Plateau, [15°10' S 125°22 ! E], 2 May 1996,
A. Lowrie 1442 (holo: PEKYH. fide Lowrie
& Kenneally (1997), n.v., iso: DNA, MEL,
fide Lowrie & Kenneally (1997), n.v).
Annual, 5-11 cm high. Glandular hairs 0.025-
0.05 mm long; glands globose, dark. Stem base
not thickened. Stems elongate, glabrous.
Leaves 8-12 per plant, mostly in terminal
rosette, with some scattered along stems, linear,
c. 7.5 mm long, 0.4-0.7 mm wide, glabrous; apex
mucronate; base cuneate, or truncate; margins
entire. Petioles absent. Scapes 1-10 per plant,
c. 0.7 mm in diameter, glandular-hairy; sterile
bracts absent. Inflorescences c. 9 cm long, 1-
flowered. Bracteoles absent. Pedicels absent
or rudimentary. Hypanthium linear, glandular-
hairy at distal end only or glandular-hairy
throughout. Sepals ovate, fused into 2 entire
lips, c. 1.5 mm long, c. 1.2 mm wide, glandular-
hairy, obtuse. Corolla white, yellow, and orange,
glandular-hairy on petals only; tube c. 3 mm
long, with sinus on anterior side only.
Paracorolla absent. Labellum attached at base
of anterior sinus of corolla tube, lanceolate, c.
0.4 mm long, thick, glabrous, acuminate,
terminal appendage usually present. Petals with
posterior ones fused, A1+A2+(P1&P2).
Anterior petals c. 3.5 mm long, c. 2.5 mm wide,
bilobed, obtuse. Posterior petals c. 6 mm long,
c. 3.5 mm wide, bilobed, obtuse. Column c. 8.5
mm long, conspicuously dilated near distal end
forming a pouch for the stigma and anthers,
glabrous; lateral lobes absent. Corona absent.
Capsule linear, c. 27 mm long excluding sepals,
c. 0.7 mm wide, without raised longitudinal ribs.
Seeds c. 0.2 mm long, brown.
Distribution and habitat: Stylidium
perizostera is endemic to the Kimberley region
of Western Australia, where it is known from
the Mitchell Plateau and Bigge Island (Map
12). It grows in drainage lines of sandstone
outcrops (Lowrie & Kenneally 1997).
Note: Only an isotype was available for
examination. The description above is based
largely on Lowrie & Kenneally (1997).
619
Bean, Stylidium subgen. Andersonia
18. Stylidium claytonioides W.Fitzg., J. & Proc.
Roy. Soc.WestemAustralia3:219(1918).
Type: WestemAustralia. “between Isdell
Range and Mt Bartlett” [Barnett?], [16°
—!S 125°—’E], 1905 -6,WV. Fitzgerald
s.n., n.v.
Illustration: J.R. Wheeler (ed.), FI. of Kimb.
Region 876, t. 270B(1992).
Annual, 15-27 cm high. Glandular hairs 0.05-
0.1 mm long; glands globose, dark. Stem base
not thickened. Stems elongate, glabrous.
Leaves 9-30 per plant, mostly in terminal
rosette, with some scattered along stems, linear,
12-24 mm long, 0.7-1 mm wide, glabrous; apex
acute; base truncate; margins entire. Petioles
absent. Scapes 1-6 per plant, 0.3-0.6 mm in
diameter, glabrous; sterile bracts absent.
Inflorescences 10-18 cm long, 1-flowered.
Bracteoles absent. Pedicels absent or
rudimentary. Hypanthium linear, glabrous
throughout. Sepals ovate or orbicular, fused
into 2 entire lips, 2.3-2.4 mm long, 1.5-2.7 mm
wide, glandular-hairy, obtuse. Corolla pink and
mauve, glandular-hairy on tube and petals; tube
2.6-3 mm long, with sinus on anterior side only.
Paracorolla absent. Labellum attached at base
of anterior sinus of corolla tube, lanceolate, 0.6-
1 mm long, thick, glabrous or glandular-hairy,
acute or acuminate, terminal appendage usually
present; 0-0.5 mm long. Petals with posterior
ones fused, A1+A2+(P1&P2). Anterior petals
2.4-3.5 mm long, 1.2-2.8 mm wide, entire,
obtuse. Posterior petals 4.2-5.9 mm long, 1.7-
3.5 mm wide, bilobed, obtuse. Column 7.5-9
mm long, conspicuously dilated near distal end
forming a pouch for the stigma and anthers,
glabrous; lateral lobes absent. Corona absent.
Capsule not seen.
Specimens examined: Western Australia. Barnett
Gorge, c. 250 km SW of Wyndham, Jun 1976,
Beaugehole 52339 (DNA); Vansittart Bay, creek
leading into Pauline Bay, May 1984, Chesterfield 354
(MEL); Wonjarring Falls, Carson River escarpment,
Jun 1984, Chesterfield 425 (MEL); unnamed creek
running into Pauline Bay, May 1984, Forbes 2164
(MEL); northern end of Airfield Swamp, Mitchell
Plateau, Jun 1976, Kenneally 4843 (K); Kalumburu
road, 40 km N from Drysdale River crossing, Jun
1984, Willis s.n. (MEL).
Reconstituted or spirit material examined:
Forbes 2164 (2 fls); Willis s.n. (2 fls).
Distribution and habitat: Stylidium
claytonioides is endemic to the Kimberley
region ofWesternAustralia (Map 11). It grows
along creekbanks or in seasonal swamps near
sandstone outcrops.
Phenology: Flowers have been recorded for
May and June.
Conservation status: Data deficient (IUCN
1994).
C. Stylidium sect. Tenella (Benth.)A.R.Bean
comb, et stat. nov.
S. ser. Tenella Benth. (as ‘ Tenellae ’), FI.
Austral. 4: 22 (1869). Type: S. tenellum Sw. ex
Willd.
Leaves sessile, often minute, bract-like, rarely
in rosettes. Scapes mostly absent. Petals all
free from each other or with posterior ones
fused, all petals bilobed (excepts, prophyllum );
plants glabrous except for hypanthium apex,
sepals and corolla; paracorolla present;
glandular hairs globose, <0.2 mm long; labellum
thick, glabrous, acuminate, attached to outside
of corolla tube; column often bearing lateral
lobes.
11 species; south-east Asia, Malesia,
northern Australia.
19. Stylidium nominatum Carlquist, Aliso 10:
38(1981)
Stylidium mitrasacmoides Carlquist, Aliso
9: 419 (1979), nom. illeg ., non F.Muell.
(1859). Type: Northern Territory, c. 0.5 km
NE of Camp 1, Arnhem Land, 12° 59’S 133°
09’E, 4 June 1978, 5. Carlquist 15409
(holo: RSA; iso: DNA, K/tv, PERTH/tv.).
Annual, 4—15 cm high. Glandular hairs 0.05-0.1
mm long; glands globose, dark. Stem base not
thickened. Stems elongate, glabrous. Leaves
in basal rosette and scattered along stems.
Rosette leaves 5-15, oblanceolate, 2.5-9 mm
long including petiole, 0.5-1.5 mm wide,
glabrous; apex obtuse; base cuneate; margins
entire; petioles 1-4.5 mm long. Stem leaves 4-
11, linear, 2.3-4.7 mm long, 0.3-0.5 mm wide,
glabrous, apex acute, base truncate, sessile.
Scapes absent. Inflorescences 2-6 cm long,
determinate, monochasially cymose; branches
620
glabrous. Bracts linear, 2-3 mm long, glabrous,
acute. Bracteoles absent. Pedicels absent or
rudimentary. Hypanthium linear, glabrous
throughout or glandular-hairy at distal end
only. Sepals debate, all free or with 3 free and 2
fused for more than half their length, 1.2-2.1
mm long, 0.3-0.4 mm wide, glabrous or
glandular-hairy, acute. Corolla white, glandular-
hairy on tube and petals; tube 0.5-1.6 mm long,
with sinus on anterior side only. Paracorolla
continuous, thin, glabrous, 0.3-0.5 mm high.
Paracorolla lobes or appendages 2, all similar,
obtuse, none opposite the anterior petals, 2
opposite the posterior petals. Paracorolla
glands absent. Labellum attached to outside
of corolla tube or attached at base of anterior
sinus of corolla tube, lanceolate, 0.3-0.5 mm
long, thick, glabrous, acute, terminal
appendage absent. Petals all free,
A1+A2+P1+P2. Anterior petals 0.6-1.5 mm
long, 0.6-1.5 mm wide, bilobed, obtuse.
Posterior petals 1.3-3.3 mm long, 1.2-3.3 mm
wide, bilobed, obtuse. Column 2.5-5 mm long,
of uniform width throughout, glabrous; lateral
lobes absent. Corona absent. Capsule linear,
4.5-12 mm long excluding sepals, 0.3-0.5 mm
wide, without raised longitudinal ribs; halves
detaching distally, not recurved. Seeds
ellipsoidal, 0.15-0.2 mm long, brown; surface
convex, smooth.
Specimens examined: Northern Territory. Melville
Island, Apr 1987, Fensham 481 (DNA); Arnhem Land,
19 km E of Jabiru, Apr 1989, Johnson 4557 (BRI);
Cooper Creek area, Nabarlek, Apr 1979, Rankin 2213
(CANB, DNA, K); Nabarlek, Apr 1979, Rice 3204
(DNA); Rola Plains, Yapilika, Melville Island, Apr
1987, Wilson 62 (DNA).
Reconstituted or spirit material examined:
Rankin 2213 (3 fls); Wilson 62 (3 fls).
Distribution and habitat: Stylidium
nominatum is endemic to the Northern Territory
where it is known from the northern parts of
Kakadu National Park and also from Melville
Island (Map 17). It has been recorded from a
Melaleuca viridiflora woodland, from the base
of a sandstone escarpment, and from a flat
treeless area with gravelly yellow soil.
Phenology: Flowers and capsules have been
recorded from April to June.
Austrobaileya 5(4): 589-649 (2000)
Affinities: S. nominatum is very close to S.
capillare. See affinities section under that
species.
Notes: There is large variation between the
available specimens. For example, Rankin 2213
has short capsules, a short corolla tube and
small petals. However, further collections are
needed to determine whether more than one
taxon is involved.
Conservation status: Data deficient (IUCN
1994).
20. Stylidium capillare RBr., Prodr. 570 (1810);
Candollea capillaris (R.Br.) F.Muell.,
Syst. census Austral, pi. 86 (1883). Type:
[Queensland. Cook District:] Endeavour
River [15°2-’S 145°1-’E, June-July 1770],
J. Banks & D. Solander (holo: BM).
Stylidium quadrifurcatum F.L.Erickson &
J.H. Willis, Victorian Naturalist 73:5 (1956),
syn. nov. Type: Northern Territory. Pine
Creek, [13°4-’S 131° 5-E], April 1904,
J.H. Niemann (holo: MEL).
Illustration: R. Erickson, Triggerplants, plate
54 (1958), as S. quadrifurcatum.
Annual, 6-13 cm high. Glandular hairs 0.05-0.1
mm long; glands globose, dark. Stem base not
thickened. Stems compressed (with leaves in
basal rosette). Leaves 4-7 per plant, obovate
or orbicular, 1.5-5 mm long, 1-3 mm wide,
glabrous; apex obtuse, or acute; base truncate;
margins entire. Petioles absent. Scapes l(-2)
per plant, 0.3-0.8 mm in diameter, glabrous;
sterile bracts present. Inflorescences 6-13 cm
long, determinate, monochasially cymose;
branches glabrous. Bracts lanceolate, 1-2 mm
long, glabrous, obtuse or acute. Bracteoles
absent. Pedicels absent or rudimentary.
Hypanthium linear, glandular-hairy at distal end
only. Sepals deltate, with 3 free and 2 fused for
more than half their length, 1.1-1.6 mm long,
0.3-0.5 mm wide, glabrous, acute. Corolla white,
glandular-hairy on tube only; tube 1.6-2 mm
long, with sinus on anterior side only.
Paracorolla continuous, thin, glabrous, 0.1-0.4
mm high. Paracorolla lobes or appendages 4,
all similar, obtuse, 2 opposite the anterior petals,
2 opposite the posterior petals. Paracorolla
621
Bean, Stylidium subgen. Andersonia
glands absent. Labellum attached to outside
of corolla tube, ovate or lanceolate, 0.4-0.8 mm
long, thick, glabrous, acute or acuminate,
terminal appendage absent. Petals all free,
A1+A2+P1+P2. Anterior petals 1.1-1.7 mm
long, 1-2 mm wide, bilobed, obtuse. Posterior
petals 2.8-4.2 mm long, 2.5-3.1 mm wide,
bilobed, obtuse. Column 5-6 mm long, of
uniform width throughout, glabrous; lateral
lobes absent. Corona absent. Capsule linear,
11-20 mm long excluding sepals, 0.4-0.8 mm
wide, without raised longitudinal ribs; halves
coherent distally. Seeds ellipsoidal, 0.15-0.25
mm long, brown; surface lacunose, smooth.
Specimens examined: Northern Territory. 31 km
W of “Wollogorang”, Jun 1999, Bean 15141 (BRI);
O.T. Station, May 1947, Blake 17644 (BRI);
Litchfield N.P., catchment of Aida Ck, Mar 1995,
Cowie 5200 & Taylor (DNA); Kakadu N.P., Mar 1982,
Dunlop 6232 & Taylor (DNA); Berry Springs, May
1977, Parker 851 (DNA); Berry Springs, Apr 1978,
Rankin 1216 (DNA, K); 150 km W of Borroloola,
Apr 1979, Rankin 1880 (DNA); south to tributaries
of McKinley R., Mar 1882, Tate s.n. (MEL).
Queensland. Cook District: Tait and Lynd Rivers, c.
1877, Armit 556 (MEL); near Cooktown, May 1970,
Blake 23355 (BRI); Douglas Creek, c. 7 miles [11
km] SE of Mareeba, Apr 1967, Brass 33528 (BRI);
14.3 km N of Batavia Downs on the Peninsula
Development road, Apr 1990, Clarkson 8345 &
Neldner (BRI, MBA, PERTH, QRS); off Peninsula
road, 52.2 km along main Weipa road, Apr 1988,
Forster PIF4101 & Liddle (BRI); Unigan Nature
Reserve, Weipa, Mar 1990, Forster PIF6509 &
O Reilly (BRI); between Cobra & Levison Creeks, c.
4 miles [6 km] E of Mareeba, Apr 1953, Melville
3737 et al. (MEL); downstream from Hey Point,
Franjum Point, on Embley River, S of Weipa, Mar
1981, Morton AMI 164 (BRI, MEL); near Mareeba,
Apr 1967, Pedley 2274 (BRI); near Ayton,
Bloomfield, Mar 1978, Scarth-Johnson 730A (BRI);
near CSIRO Tobacco Research Institute, 3 miles [5
km] E of Mareeba, May 1959, Thorne 20988 & Jones
(BRI, RSA).
Reconstituted or spirit material examined:
Clarkson 8345 & Neldner (3 fls); Dunlop 6232
& Taylor (2 fls); Morton AMI 164 (2 fls); Pedley
2274(1 fl).
Distribution and habitat: Stylidium capillare
is widepread from Litchfield National Park in
the Northern Territory to Cooktown and
Mareeba in northern Queensland (Map 7). This
ephemeral species grows in eucalypt woodland
or on swamp margins with Melaleuca spp. The
soils are generally sandy.
Phenology: Flowers and capsules have been
recorded between March and June.
Affinities: S. capillare is closely related to S.
nominatum , but differs by having leaves in
basal rosette only (rosette and stem leaves for
S. nominatum ); bracts 1-2 mm long (2-3 mm
for S. nominatum ); corolla glandular-hairy on
tube only (glandular-hairy on tube and petals
for A nominatum ); corolla tube 1.6-2.0 mm long
(0.5-1.6 mm long for S. nominatum ); and
capsules 11-20 mm long (4.5-12 mm for S.
nominatum).
Note: It is remarkable that exactly 200 years
elapsed between Banks and Solander’s original
collection and the next herbarium collection
from the Cooktown area. This is indicative of
the very thorough botanical collecting done
by Banks and Solander during their enforced
stay at Endeavour River.
Conservation status: S. capillare is not
considered to be rare or threatened.
21. Stylidium prophyllum Lowrie & Kenneally,
Nuytsia 11(2): 210-2 (1997). Type:
Western Australia. On the road to Bell
Gorge, 2 km west of Silent Grove camping
area, [17°05’S 125°15’E], 5 June 1995,A.
Lowrie 1180 (holo: PERTH Jide Lowrie &
Kenneally (1997) n.vj iso: DNA, MEL,
fide Lowrie & Kenneally (1997) n. v.).
Annual, 8-30 cm high. Glandular hairs 0.05-0.1
mm long; glands globose, dark. Stem base not
thickened. Stems elongate, glabrous. Leaves
4-10 per plant, scattered along stems, deltate,
0.7-1.5 mm long, 0.3-0.6 mm wide, glabrous;
apex obtuse, or acute; base truncate; margins
entire. Petioles absent. Scapes absent.
Inflorescences 3-14 cm long, determinate,
monochasially cymose; branches glabrous.
Bracts linear or deltate, 1-2 mm long, glabrous,
acute. Bracteoles absent. Pedicels absent or
rudimentary. Hypanthium linear, glandular-
hairy at distal end only. Sepals oblanceolate,
with 3 free and 2 fused for more than half their
length, 1.7-1.9 mm long, 0.4-0.5 mm wide,
glabrous, acute. Corolla pink, glandular-hairy
on tube and petals; tube 2-2.5 mm long, with
sinus on anterior side only. Paracorolla
discontinuous, thin, glabrous, 0.2-0.3 mm high.
622
Paracorolla lobes or appendages 4, all similar,
acute or obtuse, 0 opposite the anterior petals,
4 opposite the posterior petals. Paracorolla
glands absent. Labellum attached at base of
anterior sinus of corolla tube, lanceolate, c. 0.5
mm long, thick, glabrous, acuminate; 0.3-0.7
mm long. Petals all free,A1+A2+P1+P2. Anterior
petals 2.3-2.6 mm long, 1.3-1.6 mm wide, entire,
obtuse. Posterior petals 4.1^1.5 mm long, 2-2.2
mm wide, entire, obtuse. Column 5-6 mm long,
slightly dilated near distal end, glabrous; lateral
lobes absent. Corona absent. Capsule linear,
11-17 mm long excluding sepals, 0.5-0.8 mm
wide, without raised longitudinal ribs; halves
coherent distally. Seeds ellipsoidal, 0.25 mm
long, brown; surface convex, colliculate.
Specimens examined: Western Australia: On the
road to Bell Gorge, 2 km west of Silent Grove camping
area, Jun 1995, Lowrie 1180 (DNA). Northern
Territory: Fitzmaurice River, Macadam Range, Feb
1994, Leach 4126 (DNA).
Reconstituted or spirit material examined:
Leach 4126 (2 fls).
Distribution and habitat: Stylidium
prophyllum occurs in the Kimberley region of
Western Australia and adjacent areas of
Northern Territory (Map 16). It grows on grassy
floodplains, seepage areas and waterways.
Phenology: Flowers and capsules have been
recorded from February to June.
Affinities: S. prophyllum is closely related to
S. fissilobum, but differs by the glabrous sepals
(glandular-hairy for S. fissilobum ); labellum
attached at the base of the anterior sinus of
corolla tube (attached to outside of corolla tube
for S. fissilobum), anterior petals 2.3-2.6 mm
long (1.3-2.2 mm for S. fissilobum ); anterior and
posterior petals entire (bilobed for S.
fissilobum).
Conservation status: Data deficient (IUCN
1994).
22. Stylidium aquaticum A. R. Bean sp. nov.
affinis Stylidio fissilobo sed differens
foliis filiformibus 0.1-0.3 mm latis,
paracorolla 0.1-0.2 mm alta columna
breviore et lobis contra petala antica
nullis.Typus: Northern Territory.
Austrobaileya 5(4): 589-649 (2000)
Headwaters Cui-Eci Creek, [14°28’S
130°06’E], 12 May 1994, ID. Cowie 4906
& D. Albrecht (holo: DNA).
Annual, 18-30 cm high. Glandular hairs 0.05-
0.1 mm long; glands globose, dark. Stem base
not thickened. Stems elongate, glabrous.
Leaves 20-100 per plant, scattered along stems,
linear, 1.7-11 mm long, 0.1-0.3 mm wide,
glabrous; apex acute; base truncate; margins
entire. Petioles absent. Scapes absent.
Inflorescences 3-8 cm long, determinate,
monochasially cymose; branches glabrous.
Bracts linear, 1.5-2.5 mm long, glabrous, acute.
Bracteoles absent. Pedicels absent or
rudimentary. Hypanthium linear, glandular-
hairy at distal end only. Sepals oblanceolate,
with 3 free and 2 fused for more than half their
length, 1.5-1.9 mm long, 0.4-0.5 mm wide,
glandular-hairy, acute. Corolla white or pink,
glandular-hairy on tube and petals; tube 1.5-
1.8 mm long, with sinus on anterior side only.
Paracorolla continuous, thin, glabrous, 0.1-0.2
mm high. Paracorolla lobes or appendages 4,
all similar, obtuse, 0 opposite the anterior petals,
4 opposite the posterior petals. Paracorolla
glands absent or 2. Labellum attached to
outside of corolla tube, ovate or lanceolate, 0.5-
0.6 mm long, thick, glabrous, acuminate; 0.2-
0.3 mm long. Petals all free, A1+A2+P1+P2.
Anterior petals 1.3-1.6 mm long, 0.9-1.3 mm
wide, bilobed, acute or obtuse. Posterior petals
3.3^1.4 mm long, 3-4.1 mm wide, bilobed,
obtuse. Column 4.5 mm long, of unifonn width
throughout, glabrous; lateral lobes present, c.
0.3 mm wide. Corona absent. Capsule linear, 19
mm long excluding sepals, c. 0.4 mm wide,
without raised longitudinal ribs. Seeds not
seen. Fig. 6A-D.
Specimens examined: see type
Reconstituted or spirit material examined:
Cowie 4906 & Albrecht (3 fls)
Distribution and habitat: Stylidium
aquaticum is known only from the type
specimen collected in north-western Northern
Territory (Map 16). It was recorded from a
Melaleuca viridiflora swamp, growing in
shallow water c. 7 cm deep.
623
Bean, Stylidium subgen. Andersonia
Phenology: Flowers have been recorded for
May
Affinities: S. aquaticum is closely related to S.
fissilobum , but differs by the thread-like leaves
0.1-0.3 mm wide (0.3-0.7 mm for S', fissilobum );
paracorolla 0.1-0.2 mm high with no lobes
opposite the anterior petals (0.2-0.5 mm high,
2^1 lobes opposite the anterior petals for S.
fissilobum ); and column c. 4.5 mm long (5-7
mm long for S. fissilobum).
Conservation status: Data deficient (IUCN
1994).
Etymology: From the Latin aquaticus , in
reference to the habitat of this species.
23. Stylidium fissilobum F.Muell., Fragm. 1:154
(1859); Candollea fissiloba (F.Muell.)
F.Muell., Syst. census Austral, pi. 86
(1883). Type: [Northern Territory], Victoria
River, [15° —’S 129°—’E], 1855-56, A
Mueller (lecto: MEL [MEL 1061830]), here
chosen.
Stylidium pseudotenellum O.Schwarz,
Repert. Spec. Nov Regni Veg. 24: 104
(1927). Type: Northern Territory. Port
Darwin, 8 miles [13 km] east, [12° 2-’S
130° 5-’E], undated, F.A.K. Bleeser 466
(holo: B, destroyed).
Illustrations: R. Erickson, Triggerplants, plates
51, 56 (1958); J.R. Wheeler (ed.), FI. of Kimb.
Region 878, t. 271B(1992).
Annual, 5-50 cm high. Glandular hairs 0.05-0.2
mm long; glands globose, dark. Stem base not
thickened. Stems elongate, glabrous. Leaves
4-34 per plant, scattered along stems, linear or
deltate, 1-6 mm long, 0.3-0.7 mm wide,
glabrous; apex obtuse, or acute; base truncate;
margins entire. Petioles absent. Scapes absent.
Inflorescences 3-18 cm long, determinate,
monochasially cymose; branches glabrous.
Bracts linear or deltate or lanceolate, 1-4 mm
long, glabrous, obtuse or acute. Bracteoles
absent. Pedicels absent or rudimentary.
Hypanthium linear, glandular-hairy at distal end
only. Sepals deltate or oblanceolate, with 3 free
and 2 fused for more than half their length, 1.4-
2.3 mm long, 0.2-0.7 mm wide, glandular-hairy,
obtuse or acute. Corolla white or pink or mauve,
glandular-hairy on tube and petals, or on tube
only; tube 1.6-2.3 mm long, with sinus on
anterior side only. Paracorolla continuous, thin,
glabrous, 0.2-0.5 mm high. Paracorolla lobes
or appendages 4-8, all similar, acute or obtuse,
2-4 opposite the anterior petals, 2-4 opposite
the posterior petals. Paracorolla glands absent
or present, 0, 2 or 8. Labellum attached to
outside of corolla tube, ovate or lanceolate, 0.4-
0.7 mm long, thick, glabrous, acute or
acuminate; 0.1-0.3 mm long. Petals all free,
A1+A2+P1+P2 or with posterior ones fused,
A1+A2+(P1&P2). Anterior petals 1.3-2.2 mm
long, 0.6-1.5 mm wide, bilobed, acute or obtuse.
Posterior petals 1.9-4.2 mm long, 1.4-3.4 mm
wide, bilobed, obtuse. Column 5-7 mm long, of
uniform width throughout, glabrous; lateral
lobes absent or present, 0-0.5 mm wide. Corona
absent or not extending beyond anthers.
Capsule linear, 12-21 mm long excluding sepals,
0.4-1 mm wide, without raised longitudinal ribs;
halves coherent distally. Seeds globose or
ellipsoidal, 0.15-0.25 mm long, brown; surface
convex, smooth or colliculate.
Specimens examined: Western Australia. 30 km
NW of Drysdale River crossing, c. 12 km W of Gibb
River-Kalumburu Mission road, Jun 1976,
Beauglehole 52220 (DNA); Mount Brophy Springs,
Gardner Range, 190 km SE of Halls Ck, Jul 1995,
Coate 370B (BRI). Northern Territory. 2 km E of
Berry Springs turnoff, on road to Water Supply Dam,
Jun 1977, Carlquist 15195 (DNA); Mary River,
undated, Clark 1737 (DNA); Malabanbandjii Swamp,
10 km E of Nourlangie Ranger station on Pine Creek
road. May 1980, Craven 5484 (CANB, DNA); 18 km
SSW of Cooinda on Pine Creek road. May 1980,
Craven 5624 (CANB, DNA, MEL); 19 km NNW of
Twin Falls, Jun 1980, Craven 6292 (MEL); Muirella
Park turnoff. May 1974, Fox 409 (DNA); Port
Darwin, 1885, Holtze 482 (MEL); Port Darwin, Oct
1888, Holtze 882 (MEL); Nourlangie Rock area. May
1973, Must 1125 (BRI, DNA); Nourlangie Creek, [12°
50’S 132° 46’E], May 1974, Must 1211 (BRI, DNA);
Berry Springs reserve. May 1977, Parker 852 (DNA);
Berry Springs, Apr 1978, Rankin 1217 (DNA); end
of Gulnare road, Elizabeth River, Mar 1995, van
KerckhofD36 (DNA). Queensland. Cook District :
Robertson River, c. 1878, Armit 797 (MEL); One
Hundred Mile swamp, near Undara resort, E of Mt
Surprise, Jul 1998, Bean 13757 & Fox (BRI, DNA,
MEL, NSW, PERTH, QRS). BurkeDistrict: 21 miles
[34 km] SE of Croydon, Jul 1954, Blake 19581 (BRI,
DNA); c. 29 km NW of old “Corinda” on the road
from Doomadgee Aboriginal Station to Woologorang,
May 1974, Pullen 9149 (BRI); 23 miles [37 km] SE
of Croydon township, Jul 1954, Speck 4723 (BRI,
CANB, DNA).
624
Reconstituted or spirit material examined:
Bean 13757 (3 fls); Blake 19581 (2 fls); Must
1211(2 fls), Parker 852 (3 fls).
Distribution and habitat: Stylidium fissilobum
has a broad distribution across northern
Australia, from the western Kimberley and
Northern Territory to as far east as Mt Surprise
in Queensland (Map 15). It grows in moist to
wet sand at the edge of swamps, often in
association with grassland-sedgeland. The
weak-stemmed S. fissilobum threads its way
through the often dense cover of grasses and
sedges and is supported by them.
Phenology: Flowers and capsules have been
recorded from March to July
Affinities: S. fissilobum is related to S.
oviflorum and S. aquaticum. See notes under
those species.
Typification: The only Mueller specimen of S.
fissilobum at MEL is one which has come via
the O.W. Sonder herbarium. Presumably there
was or is other material at MEL, but it cannot
now be found. Therefore the specimen cited
above is chosen as lectotype, as it is a good
specimen and its details agree with the
protologue.
No isotypes of S. pseudotenellum are
known (McKee 1963). From the description, it
appears to be a synonym of S. fissilobum.
Carl quist (1979: 443) was also of this opinion.
Conservation status: Data deficient (IUCN
1994).
24. Stylidium oviflorum A.R.Bean sp. nov.
affinis S. fissilobo sed differens corolla
alba luteaque, paracorollae lobis 4-6
contra petala postica et capsulis
generaliter longioribus (18-25 mm longis).
Typus: Queensland. Cook District: Gorge
Creek, c. 10 km west of Mareeba on the
Dimbulahroad, [17 o 0ES 145°20’E], 15
April 1989 ,J.R. Clarkson 7 880 & R.J.F.
Henderson (holo: BRI; iso: DNA, MBA,
PERTH, QRS).
Austrobaileya 5(4): 589-649 (2000)
Annual, 6-26 cm high. Glandular hairs 0.1-0.2
mm long; glands globose, dark. Stem base not
thickened. Stems elongate, glabrous. Leaves
3-11 per plant, scattered along stems, linear or
debate, 1.4-3 mm long, 0.25-0.6 mm wide,
glabrous; apex obtuse, or acute; base truncate;
margins entire. Petioles absent. Scapes absent.
Inflorescences 4-15 cm long, determinate,
monochasially cymose; branches glabrous.
Bracts linear or deltate, 1.0-2.0 mm long,
glabrous, acute. Bracteoles absent. Pedicels
absent or rudimentary. Hypanthium linear,
glandular-hairy at distal end only. Sepals
deltate, with 3 free and 2 fused for more than
half their length, 1.5-1.8 mm long, 0.3-0.5 mm
wide, glandular-hairy, acute. Corolla white and
yellow, glandular-hairy on tube only or
glandular-hairy on tube and petals; tube 1.6-2
mm long, with sinus on anterior side only.
Paracorolla continuous, thin, glabrous, 0.3-0.4
mm high. Paracorolla lobes or appendages 6-
8, all similar, obtuse, 2 opposite the anterior
petals, 4-6 opposite the posterior petals.
Paracorolla glands absent. Labellum attached
to outside of corolla tube, ovate, 0.5-0.6 mm
long, thick, glabrous, acuminate, terminal
appendage absent. Petals all free,
A1+A2+P1+P2 or with posterior ones fused,
A1+A2+(P1&P2). Anterior petals 1.4-1.8 mm
long, 0.9-1.5 mm wide, bilobed, obtuse.
Posterior petals 2.7^1 mm long, 1.6-2.7 mm
wide, bilobed, obtuse. Column 5-6 mm long, of
uniform width throughout, glabrous; lateral
lobes present, 0.25-0.3 mm wide. Corona absent
or not extending beyond anthers. Capsule
linear, 18-25 mm long excluding sepals, 0.6-0.8
mm wide, without raised longitudinal ribs;
halves coherent distally. Seeds ellipsoidal, 0.2
mm long, brown; surface convex, smooth. Fig
6E-F.
Specimens examined: Queensland. Cook District:
18 km from Kennedy Highway, on road to ‘Barwidgi’,
Aug 1997, Bean 12212 (BRI); Boyle Creek, NW of
Mareeba, Apr 1962, McKee 9176 (BRI, NSW); Gorge
Creek, 10 miles [16 km] W of Mareeba on Dimbulah
road, Apr 1962, McKee 9228 (BRI, NSW); Cobra
Creek, Mareeba District, Apr 1962, McKee 9386 (BRI);
Herberton-Chillagoe road, c. 8 miles [13 km] from
Herberton, Jun 1958, Pedley 252 (BRI). North Kennedy
District: c. 1 km south of ‘Glen Harding’ turnoff,
south of Mt Garnet, Jul 1997, Bean 12144 (BRI); c. 8
km NE of Wairuna, Aug 1997, Bean 12179 (BRI);
Evelyn Creek Conservation Park, Jun 1999,
McDonald KRM1 & Thompson (BRI).
625
Bean, Stylidium subgen. Andersonia
Reconstituted or spirit material examined:
Bean 12144 (2 fls); Bean 12179 (2 fls); Bean
12212 (2 fls); Clarkson 7880 & Henderson (1
fl>
Distribution and habitat: Stylidium oviflorum
is restricted to north Queensland, from
Mareeba to Wairuna and west to Barwidgi (Map
15). It grows on sandy soils, in seepage areas
on hillsides, or beside creeks. Canopy species
include Eucalyptus camaldulensis, E. cullenii
or Melaleuca viridiflora.
Phenology: Flowers and capsules have been
recorded from April to August.
Affinities: S. oviflorum is closely related to S.
fissilobum , but differs by the corolla colour,
combining white and rich yellow, the erect self-
supporting stems, the 4-6 paracorolla lobes
opposite the posterior petals, and the generally
longer capsules 18-25 mm long.
Conservation status: Data deficient (IUCN
1994).
Etymology: From the Latin ovi- egg ancj florus-
flower, in reference to the corolla colour which
combines rich yellow and pure white, just as in
a fried hen’s egg.
25. Stylidium confertum A.R.Bean sp. nov.
affinis Stylidio fissilobo sed differens foliis
numerosis prope basin plantae dense
aggregatis, paracorolla interrupta lobis 0-2
(0.1-0.3 mm altis) contra petala antica. Typus:
Queensland. Cook District: Walsh’s Pyramid,
east flank of main north ridge, [17° 07’S 145°
48’E], 28April 1998, C. Lyons 194 (holo: BRI;
iso: DNA, JCT, K, L, MEL, MO, NSW, PERTH,
QRS, distribuendi).
Annual, 6-21 cm high. Glandular hairs 0.05-0.1
mm long; glands globose, dark. Stem base not
thickened. Stems elongate, glabrous. Leaves
15-100 per plant, scattered along stems, linear
or deltate, 1.5-4 mm long, 0.2-0.7 mm wide,
glabrous; apex obtuse, or acute, or acuminate;
base truncate; margins entire. Petioles absent.
Scapes absent. Inflorescences 3-13 cm long,
determinate, monochasially cymose; branches
glabrous. Bracts linear or deltate, 1-2.5 mm
long, glabrous, obtuse or acute. Bracteoles
absent. Pedicels absent or rudimentary.
Hypanthium linear, glandular-hairy at distal end
only. Sepals deltate, with 3 free and 2 fused for
more than half their length, 1.1-1.8 mm long,
0.3-0.5 mm wide, glandular-hairy or glabrous,
acute. Corolla white, glandular-hairy on tube
and petals; tube 1.3-1.9 mm long, with sinus
on anterior side only. Paracorolla
discontinuous, thin, glabrous, 0.1-0.3 mm high.
Paracorolla lobes or appendages 2-6, all similar,
obtuse, 0-2 opposite the anterior petals, 2-4
opposite the posterior petals. Paracorolla
glands absent. Labellum attached to outside
of corolla tube, ovate, 0.4-0.7 mm long, thick,
glabrous, acuminate, terminal appendage
absent. Petals all free, A1+A2+P1+P2. Anterior
petals 1-1.7 mm long, 0.5-2 mm wide, bilobed,
acute or obtuse. Posterior petals 2.5-3.8 mm
long, 1.5-3.3 mm wide, bilobed, obtuse. Column
4.5-6 mm long, of uniform width throughout,
glabrous or with glandular hairs; lateral lobes
present, 0.25-0.35 mm wide. Corona absent.
Capsule linear, 16-24 mm long excluding sepals,
0.4-0.7 mm wide, without raised longitudinal
ribs; halves coherent distally. Seeds ellipsoidal,
0.15-0.25 mm long, brown; surface convex,
colliculate or smooth. Fig. 7A-C.
Specimens examined: Queensland. Cook District:
lower western slopes of Walsh’s Pyramid, May 1962,
Blake 21760 (BRI); slopes of Mt Fraser, Apr 1932,
Brass 2534 (BRI); Daintree N.P., Little Daintree
River, May 1998, Forster 22800 et al. (BRI, DNA,
QRS); Davies Creek, 0.4 km E of falls, Apr 1998,
Wannan 725 (BRI).
Reconstituted or spirit material examined:
Blake 21760 (1 fl); Brass 2534 (2 fls); Forster
22800 (2 fls )■ Lyons 194 (3 fls).
Distribution and habitat: Stylidium confertum
is known only from a few localities in the “wet
tropics” of north Queensland, between Tully
and Cooktown (Map 13). Altitudes where found
range from 100-880 metres. The habitat is open
rock faces or rocky creekbanks, where the
species is confined to small islands or pockets
of vegetation which remain damp for a few
months after the wet season, and are dominated
by mosses, Micraira subulifolia or small
herbs.
Phenology: Flowers and capsules have been
recorded for April and May.
626
Austrobaileya 5(4): 589-649 (2000)
Fig. 6. A-D: Stylidium aquaticum. A. habitx0.9; B. lower part of stem and leaves><2; C. anterior view of flower
(column removed)x9; D. column, showing lateral lobesx9. E-F: Stylidium oviflorum. E. habitx0.7; F. anterior
view of flowerx9. A-D: Cowie 4906 & Albrecht ; E-F: Bean 12179.
627
Bean, Stylidium subgen. Andersonia
Affinities: S. confertum is close to S.
fissilobum , but differs by the 15-100 leaves
densely clustered at base of plant (4-34 leaves
scattered along stem for S. fissilobum );
discontinuous paracorolla with 0-2 lobes
opposite the anterior petals (continuous, 2-4
lobes opposite the anterior petals for S.
fissilobum)., paracorolla lobes 0.1—0.3 mm high
(0.2-0.5 mm for S. fissilobum).
Conservation status: Data deficient (IUCN
1994).
Etymology: The specific epithet is from the
Latin confertus , meaning crowded or close
together. This refers to the clustering of leaves
near the base of the stem in this species.
26. Stylidium diffusumRBr, Prodr 571 (1810);
Candollea diffusa (R.Br.) F.Muell, Syst.
census Austral, pi. 86 (1883). Type:
Shoalwater Bay [Queensland. Port Curtis
District: near Pine Mountain, 22° 2- S
150° 1-’ E], 3 September 1802,R. Brown
(holo: BM).
Annual, 3-14 cm high. Glandular hairs 0.025-
0.1 mm long; glands globose, dark. Stem base
not thickened. Stems elongate, glabrous.
Leaves 5-20 per plant, scattered along stems,
linear or deltate, 2-8 mm long, 0.4—1.7 mm wide,
glabrous; apex obtuse, or acute; base truncate;
margins entire. Petioles absent. Scapes absent.
Inflorescences 1.5-8 cm long, determinate,
monochasially cymose; branches glabrous.
Bracts linear or deltate or lanceolate, 1.5-3.5
mm long, glabrous, obtuse or acute. Bracteoles
absent. Pedicels absent or rudimentary.
Hypanthium linear, glandular-hairy at distal end
only or glabrous. Sepals oblanceolate or
elliptical, with 3 free and 2 fused for more than
half their length, 1.1-2 mm long, 0.25-0.6 mm
wide, glabrous or glandular-hairy, obtuse or
acute. Corolla white or pink or mauve,
glandular-hairy on tube and petals; tube 1.3-
1.6 mm long, with sinus on anterior side only.
Paracorolla discontinuous or continuous, thin
or thick, glabrous, 0.1-0.3 mm high. Paracorolla
lobes or appendages 2-6, all similar, acute or
obtuse, 2 opposite the anterior petals, 0-4
opposite the posterior petals. Paracorolla
glands absent. Labellum attached to outside
of corolla tube, ovate or lanceolate, 0.3-0.7 mm
long, thick, glabrous, acute or acuminate; 0.1-
0.2 mm long. Petals all free, A1+A2+P1+P2.
Anterior petals 0.5-1 mm long, 0.5-1 mm wide,
bilobed or entire, acute or obtuse. Posterior
petals 1.5-2.6 mm long, 1-1.9 mm wide, bilobed,
obtuse. Column 3.5-5 mm long, of uniform
width throughout, glabrous; lateral lobes
absent. Corona absent. Capsule linear, 8-18 mm
long excluding sepals, 0.5-1 mm wide, without
raised longitudinal ribs; halves coherent
distally. Seeds ellipsoidal, 0.2-0.25 mm long,
brown; surface convex, colliculate. Fig. 7D-F.
Specimens examined: Northern Territory. Ritjirriur
Swamp, Elcho Is., Jul 1975, Latz 6120 (CANB, DNA,
L, NSW); Hemple Bay, Groote Eylandt, May 1948,
Specht 352 (BRI). Queensland. Cook District:
Embley Range, 13 km SSW of the Batavia Downs HS,
Jul 1985, Clarkson 6060 (BRI, DNA, K, L, MBA,
QRS); Jardine River, May 1948, Brass 18885 (BRI);
Coen River, Aug 1948, Brass 19786 (BRI); southern
end of Temple Bay in upper reaches of an unnamed
creek between Glennie and Hunter Inlets, Jun 1978,
Clarkson 2204 (BRI, NSW); c. 10 km S of Musgrave
Telegraph Station, Jul 1978, Clarkson 2344 (BRI,
NSW); McLeod River, Sep 1936, Flecker 2261 (BRI);
Namelita Creek, on Venture Mine campsite, Apr 1994,
Gunness AG2326 (BRI). Burnett District: Teatree
Paddock, ‘Toondahra’, fence with ‘Manar’, Mar 1984,
Forster 1760 (BRI); 1 km SE of Mt Lorna,
‘Toondahra’, Jun 1998, Forster 22992 (BRI, MEL).
Moreton District: Coolum, south of Noosa, Apr 1974,
Blaxell 1298 (BRI, NSW).
Reconstituted or spirit material examined:
Blaxell 1298 (3 fls); Clarkson 2204 (3 fls);
Clarkson 6060 (2 fls); Forster 1760 (2 fls);
Forster22992 (2 fls).
Distribution and habitat: Stylidium diffusum
has a scattered distribution along the east
coast of Queensland, and is known from Elcho
Island and Groote Eylandt in the Northern
Territory (Map 17). It has been recorded as
growing in Melaleuca viridiflora woodlands,
a soakage area in eucalypt woodland, on
swamp edges and on damp sandy creekbanks.
Phenology: Flowers and capsules have been
recorded from March to September.
Affinities: S. diffusum is most closely related
to S. tenellum (see notes under that species)
Conservation status: Data deficient (IUCN
1994).
628
27. Stylidium tenellum Sw. exWilld., Sp. PL 4:
146 (1805). Type: “Malacca” [Malaya], 18
February 1779, J.G. Koenig (holo: B,
microfiche BRI).
Stylidium roseum Kurz, J. Asiat. Soc.
Bengal, Pt. 2, Nat. Hist. 46(2): 212 (1877).
Type: [Bangladesh] Chittagong, [22°—’
N 91°— 3 E], undated, ?W.S. Kurz (holo:
?CAL n.v.)
Stylidium tenellum var. minimum C.B.Clarke
inHook.f.,Fl. Brit. India 3:420 (1881) (as
‘minima'’). Type: India. ChotaNagpore,
at Hazaribagh, [24°—’ N 85°—’ E], 9
October 1873, C.B. Clarke 20282 (holo:
K).
Epilobium tonkinense H.Lev., Bull. Herb.
Boissier ser. 2, 7: 588 (1907). Type:
[Vietnam] Tonkin, near Quang-Yen, [20°—
‘ N 106°—‘ E], 25 November 1885, B.
Balansa 1398 (iso: E n.v., P).
Illustration: R. Erickson, Triggerplants, plate
56(1958).
Annual, 5-27 cm high. Glandular hairs 0.025-
0.1 mm long; glands globose, dark. Stem base
not thickened. Stems elongate, glabrous.
Leaves 6-14 per plant, scattered along stems,
elliptical or obovate, 3.5-8.5 mm long including
petiole, 0.8-2.5 mm wide, glabrous; apex
obtuse; base cuneate; margins entire. Petioles
0-0.4 mm long. Scapes absent. Inflorescences
2-8 cm long, determinate, monochasially
cymose; branches glabrous. Bracts linear, 2-4
mm long, glabrous, acute. Bracteoles absent.
Pedicels absent or rudimentary. Hypanthium
linear, glandular-hairy at distal end only. Sepals
oblanceolate or elliptical, with 3 free and 2 fused
for more than half their length, 1.3-1.9 mm long,
0.3-0.5 mm wide, glandular-hairy, obtuse or
acute. Corolla white or pink or mauve,
glandular-hairy on tube and petals; tube 1.6-
1.8 mm long, with sinus on anterior side only.
Paracorolla continuous, thin, glabrous, 0.2-0.3
mm high. Paracorolla lobes or appendages 3-
6, all similar, obtuse, 0-2 opposite the anterior
petals, 3-4 opposite the posterior petals.
Paracorolla glands absent. Labellum attached
at base of anterior sinus of corolla tube,
lanceolate, 0.4-0.6 mm long, thick, glabrous,
Austrobaileya 5(4): 589-649 (2000)
acute or acuminate, terminal appendage usually
present; 0-0.1 mm long. Petals all free,
A1+A2+P1+P2. Anterior petals 0.4-0.6 mm
long, 0.2-0.4 mm wide, bilobed or entire, obtuse.
Posterior petals 2.1-3.3 mm long, 1.2-1.3 mm
wide, bilobed, obtuse. Column 4-5 mm long, of
uniform width throughout, glabrous; lateral
lobes absent. Corona absent. Capsule linear,
9-21 mm long excluding sepals, 0.7-0.9 mm
wide, without raised longitudinal ribs; halves
coherent distally. Seeds ellipsoidal, 0.2-0.25 mm
long, brown; surface lacunose, colliculate.
Specimens examined: India. Serampore, East Bengal,
[22°N 88°E], undated, Griffith 3450 (K); Serampore,
undated, coll, unknown (AAU); Geirsoppa Falls, 1400
ft, Oct 1919, Hallberg 7142 (K); Bhatodih, Keonjhar,
Orissa, [21°N 85°E], Oct 1946, Mooney 2763 (K).
China. High Island, N.T., [22°N 114°E], Nov 1969,
Hu 8619 (K); Hong Kong, [22°N 114°E], Nov 1881,
Hance 887 (BM). Bangladesh. Dacca, [23°N 90°E],
Oct 1868, Clarke 7866 (K). Thailand. Kanchanadit,
Surat Thani, [9°N 99°E], Aug 1927, Kerr 13088 (BM);
Hat Yai, Songkhla, [7°N 100°E], Dec 1927, Kerr
14363 (BM); Ban Bua Dut, Ko Samui, 9N 99E, May
1928, Kerr 15718 (BM); between Chanthaburi and
Trat, [12°N 102°E], Sep 1972, Larsen 32405 et al.
(AAU); Sahm Lahn forest, Muang District, Saraburi
Province, [14°N 101°E], Oct 1973, Maxwell 73-440
(AAU). Burma. Mergui, 12N 98E, undated, Griffith
240 (BM); Kyaukpyu, Ramree Is., 19N 93E, Oct 1945,
Wallace 9181 (BM). Vietnam. Phuong Mai, Ninh
Binh province, [20°N 105°E], Jan 1883, Bon (P);
Phu Quoc Island, [10°N 103°E], Sep 1875, Godefroy
876 (P). Cambodia. Kampot, [10°N 104°E], Oct
1903, Geoffray 159 (P). Laos. Sedone, Me-Kong R.,
[14°N 105°E], 1866-8, Thorel (P). Malaya. Penang
Island, [5°N 100°E], Dec 1895, Ridley 7103 (BM);
Setul, Mar 1910, Ridley 14694 (BM); 8.25 miles [13.2
km] Kuala Trengganu-Besut road, Trengganu, [5°N
103°E], Sep 1955, Sinclair 40873 (L). Sumatra.
Taram, Kepala Bandar, E of Pajakumbuh, [0°S 100°E],
Apr 1957, Meijer 5750 (L); Taram, Pajakumbuh
region, [0°S 100°E], Jul 1957, Meijer 7151 (L).
Reconstituted or spirit material examined:
Larsen 32405 et al. (2 fls); Wallace 9181 (1 fl).
Distribution and habitat: Stylidium tenellum
has a broad distribution in south-east Asia from
southern India to southern China and south to
Sumatra (Map 3). The recent collection by
Bhasker and Kushalappa (1992) from southern
India provided a considerable extension of the
known range. It inhabits seasonally swampy
areas and has been on several occasions
reported growing near rice paddies. It often
grows at low altitudes, but has been found as
high as 1100 metres above sea level on Sumatra
629
Bean, Stylidium subgen. Andersonia
(Slooten 1954). Species or genera recorded by
Bhaskar and Kushalappa (1992) to be growing
in association with S. tenellum were Rotala
ilecebroides, Lindernia, Bergia, Canscora
diffusa, Xyris, Commelina and Eriocaulon.
Phenology: Flowers and capsules have been
recorded mostly from September to December,
with a few records from other months of the
year.
Affinities: S. tenellum is very closely related
to S. diffusum , but differs by the cuneate leaf
bases (truncate for S. diffusum ); labellum
attached at base of anterior sinus of corolla
tube (attached to outside of corolla tube for S.
diffusum ); corolla tube 1.6-1.8 mm long (1.3—
1.6 mm for S. diffusum) and anterior petals 0.2-
0.4 mm wide (0.5-1 mm wide for S. diffusum ).
Notes: No specimens from China (except Hong
Kong) have been seen by the present author,
but it is described very adequately in Hong
(1983) where it is reported to occur in the
Guangdong, Yunnan and southern Fujian
provinces.
Typification: The holotype of S. tenellum var.
minimum consists of the specimens on the
upper half of the sheet only. The other
specimens are labelled as originating from
Dacca, and are therefore excluded. From the
protologue of S. roseum, it seems clearly
referrable to S. tenellum.
Conservation status: Not evaluated.
28. Stylidium longissimum ARBeansp. nov.
affinisY inconspicuo seddifferens foliis
obtusis, corolla glandulosa purpurea
(rosacea usque rosinea, HCC 6.29 usque
5.33), paracorolla prominenti, sepalis
petalis columnisque longioribus et
capsulis multo longioribus. Typus:
Queensland. Cook District: Smugglers
track, Cape Melville National Park, north
ofCooktown, [14° 17' S 144°26' E], 19 July
1998, A.R. Bean 13671 (holo: BRI; iso:
DNA, K, L, MEL, NSW).
Annual, 16-30 cm high. Glandular hairs 0.05-
0.1 mm long; glands globose, dark. Stem base
not thickened. Stems elongate, glabrous.
Leaves 4-14 per plant, scattered along stems,
elliptical, 3.5-9.5 mm long including petiole, 1.8-
5 mm wide, glabrous; apex obtuse, or acute;
base cuneate; margins entire. Petioles 0-0.5 mm
long. Scapes absent. Inflorescences 5-20 cm
long, determinate, monochasially cymose;
branches glabrous. Bracts linear or lanceolate,
1.5-4.5 mm long, glabrous, obtuse or acute.
Bracteoles absent. Pedicels absent or
rudimentary. Hypanthium linear, glandular-
hairy at distal end only. Sepals debate or
oblanceolate, with 3 free and 2 fused for more
than half their length, 2-3 mm long, 0.5-0.7 mm
wide, glandular-hairy, acute. Corolla pink or
mauve, glandular-hairy on tube and petals; tube
2-2.3 mm long, with sinus on anterior side only.
Paracorolla continuous, thick, minutely
papillose, 0.4-0.5 mm high. Paracorolla lobes
or appendages 4, all similar, obtuse, 2 opposite
the anterior petals, 2 opposite the posterior
petals. Paracorolla glands absent. Labellum
attached at base of anterior sinus of corolla
tube, ovate, 0.5-0.7 mm long, thick, glabrous,
acuminate; 0.7-0.8 mm long. Petals all free,
A1+A2+P1+P2. Anterior petals 2.1-2.9 mm
long, 1.2-1.5 mm wide, bilobed, acute or obtuse.
Posterior petals 4.1-5.4 mm long, 3-3.2 mm
wide, bilobed, obtuse. Column 7-8 mm long,
slightly dilated near distal end, glabrous; lateral
lobes absent or present, 0-0.2 mm wide. Corona
absent. Capsule linear, 26-48 mm long
excluding sepals, 0.7-1 mm wide, without raised
longitudinal ribs; halves detaching distally, not
recurved. Seeds ellipsoidal, 0.25-0.35 mm long,
brown; surface convex, colliculate. Fig. 8E-H.
Specimens examined: Queensland. Cook District:
11 km S of beach at Bathurst Bay, Cape Melville
N.P., Jul 1998, Bean 13688 (BRI); 33 km from
Wakooka on the track to Bathurst Bay and Cape
Melville National Park, June 1984, Clarkson 5382
(BRI, QRS).
Reconstituted or spirit material examined:
Bean 13671 (3 fls); Clarkson 5382 (3 fls).
Distribution and habitat: Stylidium
longissimum is known only from the Cape
Melville area of Cape York Peninsula in north
Queensland (Map 16). It grows in Melaleuca
viridiflora woodland in sandy soil.
Phenology: Flowers and capsules have been
recorded for June.
Affinities: S. longissimum is related to S.
inconspicuum Slooten, based on the
Austrobaileya 5(4): 589-649 (2000)
Fig. 7. A-C: Stylidium confertum. A. habit* 1; B. anterior view of flower*9; C. corolla, opened out*9. D-F:
Stylidium diffusion. D. habit* 1; E. anterior view of flower and hypanthium*9; F. corolla, opened out*9. A-C:
Lyons 194; D: Brass 19786; E-F: Forster 22992.
631
Bean, Stylidium subgen. Andersonia
description and illustrations provided by
Slooten (1954), but differs by its obtuse leaf
apices, pink to mauve corolla (rosy for S.
inconspiciium ), glandular-hairy corolla tube
and petals (glabrous for S. inconspicuum ),
longer sepals, anterior petals, posterior petals,
column, and much longer capsules (26^18 mm
long vs. c. 15 mm long for S. inconspicuum).
The paracorolla in S. longissimum is much more
prominent than in S. inconspicuum.
Conservation status: S. longissimum has an
extent of occurrence of about 30x30 kilometres,
and the area of occupancy is only a small
fraction of this. A Conservation status of
“Vulnerable” is recommended, based on the
IUCN CriteriaA2(e) and D2.
Etymology: The species epithet is from the
Latin longissimus meaning “very long”. This
refers to the capsules of this species, which
are perhaps the longest of the genus.
29. Stylidium inconspicuum Slooten, Bull. Jard.
Bot. Buitenzorg Ser. Ill, 14: 171 (1937).
Types: Java. Plosokerep near halting-
place Terisi, partition 7 and 56 of forest-
section Indramajoe, [6°—’S 108°—’E],3
March 1935, C.GGJ. van Steenis 6719
(syn: ?BO,«.v.); Plosokerep near halting-
place Terisi, partitions 7,3 and 5 of forest-
section Indramajoe, [6°—’S 108°—’E],3
May 1936, C.GGJ. van Steenis 8214 &
D.F. van Slooten (syn: ?BO, n.v.).
Illustration: R. Erickson, Triggerplants, plate
56(1958).
Annual, 5-20 cm high. Stem base not thickened.
Stems elongate, glabrous. Leaves 5-30 per
plant, scattered along stems, elliptical or
obovate, 4—8 mm long, 2-5 mm wide, glabrous;
apex acute; base cuneate; margins entire.
Petioles absent. Scapes absent. Inflorescences
5-8 cm long, determinate, monochasially
cymose; branches glabrous. Bracts lanceolate
or ovate, glabrous, acute. Bracteoles absent.
Pedicels absent or rudimentary. Hypanthium
linear. Sepals oblanceolate or ovate or elliptical,
with 3 free and 2 fused for more than half their
length, 1-1.5 mm long, obtuse. Corolla pink or
red, glabrous; tube 1-1.5 mm long, with sinus
on anterior side only. Paracorolla
discontinuous or continuous. Paracorolla
glands absent. Labellum attached at base of
anterior sinus of corolla tube, ovate, c. 0.5 mm
long, glabrous, acuminate, terminal appendage
usually present. Petals all free, A1+A2+P1+P2.
Anterior petals c. 1 mm long, bilobed or entire,
obtuse. Posterior petals 1.75-2 mm long,
bilobed, obtuse. Column 3-3.5 mm long, of
uniform width throughout; lateral lobes absent.
Capsule linear, 15 mm long excluding sepals.
Seeds not seen.
Distribution and habitat: Stylidium
inconspicuum is endemic to Java (Map 3),
where it is apparently rare. It was recorded from
“moist places of grass-fields at 20-30 metres”.
Note: No material of this species was available
for examination. The description is based on
Slooten (1937).
Conservation status: Not evaluated.
D. Stylidium sect. Biloba A.R.Bean sect. nov.
Folia in rosula basali vel caulina; calyx
bilabiatus, integer, obtusus; inflorescentia
cymosa; columna glabra, fine distali magnopere
dilatata. Typus: S. rotundifolium R.Br.
Leaves in basal rosette or cauline; calyx 2-
lipped, entire, obtuse; inflorescences cymose;
labellum attached at base of anterior sinus;
column glabrous, strongly dilated at distal end.
3 species; northern Australia.
30. Stylidium rotundifolium R.Br., Prodr. 571
(1810); Candolle a rotundifolia (R.Br.)
F.Muell., Syst. census Austral, pi. 86
(1883). Type: East Coast, Port 1
[Queensland. Port Curtis District: Curtis
Island or Facing Island, 23° 4- S 151° 1-
’ E], 5-9 August 1802,7?. Brown (Bennett
No. 2600) (lecto: BM), here chosen.
Stylidium irriguum W.Fitzg., J. & Proc. Roy.
Soc. WesternAustralia 3:219(1918), syn.
nov. Types: WesternAustralia. Messmate
Creek, 1905-6, W. V. Fitzgerald (syn, n.v);
Isdell & Charnley Rivers, 1905-6, W.V.
Fitzgerald (syn, n.v ).
Stylidium reductum Carlquist,Aliso 9: 313
(1978), syn. nov. Type: Queensland. North
Kennedy District: Millstream Falls Park,
along the Ravenshoe-Mt Garnet road,
[17° 3-’S 145° 2- E], 5 July 1977, S.
Carlquist 15230 (holo: RSA).
632
Annual, 4—18 cm high. Glandular hairs 0.05-0.1
mm long; glands globose, dark. Stem base not
thickened. Stems compressed (with leaves in
basal rosette). Leaves 4-17 per plant,
oblanceolate or obovate, 5-29 mm long
including petiole, 3-10 mm wide, glabrous; apex
obtuse; base cuneate; margins entire. Petioles
1-12 mm long. Scapes 1-10 per plant, 0.2-0.4
mm in diameter, glabrous; sterile bracts absent.
Inflorescences 4-18 cm long, monochasially
cymose; branches glabrous. Bracts debate or
lanceolate, 1-1.5 mm long, glabrous, mucronate.
Bracteoles absent. Pedicels absent or
rudimentary. Hypanthium linear, glandular-
hairy at distal end only. Sepals ovate, fused
into 2 entire lips, 1-1.8 mm long, 0.8-1.3 mm
wide, sparsely glandular-hairy or glabrous,
obtuse. Corolla white or pink, glandular-hairy
on tube and petals; tube 2-3 mm long, with
sinus on anterior side only. Paracorolla absent.
Labellum attached at base of anterior sinus of
corolla tube, lanceolate, 0.4-0.6 mm long, thick,
glabrous, acuminate, terminal appendage
usually present; 0-0.1 mm long. Petals all free,
A1+A2+P1+P2. Anterior petals 1-2 mm long,
0.7-1.2 mm wide, entire, obtuse. Posterior petals
1.3-2 mm long, 0.8-1.3 mm wide, bilobed or
entire, obtuse. Column 3.5-5 mm long,
conspicuously dilated near distal end forming
a pouch for the stigma and anthers, glabrous;
lateral lobes absent. Corona absent. Capsule
linear, 12-30 mm long excluding sepals, 0.5-0.7
mm wide, without raised longitudinal ribs;
halves detaching distally, strongly recurved or
not recurved. Seeds ellipsoidal, 0.2-0.25 mm
long, brown; surface convex, colliculate.
Selected specimens: Western Australia. Gibb River-
Kalumburu road, 15.3 km N of Doggan River, May
1976, Beauglehole 51795 (DNA); Manning Gorge,
c. 275 km SW of Wyndham, Jun 1976, Beauglehole
52581 (DNA); Adcock Gorge, c. 180 km E of Derby,
Jul 1974, Carr 4233 & Beauglehole 48011 (DNA);
Mitchell River, Feb 1980, Dunlop 5275 (DNA).
Northern Territory. UDP Falls, c. 80 km NE of
Pine Creek, Aug 1978, Beauglehole 58538 & Errey
2238 (DNA); south of mouth of Daly River, Jul 1946,
Blake 16548 (BRI); 9.5 km S of Twin Fall, May 1980,
Craven 5856 (DNA); Mt Gilruth area, Jun 1978,
Dunlop 4897 (CANB, DNA); Caledon Bay, Jun 1972,
Latz 2927c (DNA); South Bay, Bickerton Island, Jun
1948, Specht 465 (BRI). Queensland. Burke District :
Westmoreland, Fagoon Creek, off track to Camp
Ridgeway, May 1997, Forster PIF21025 & Booth
(BRI); Murrays Springs, 12.1 km by road west of
Austrobaileya 5(4): 589-649 (2000)
Musselbrook mining camp, 175 km N of Camooweal,
Apr 1995, Thomas MRS32 & Johnson (BRI). Cook
District: 13.8 km from Irvinebank towards Herberton,
Aug 1997, Bean 12192 (BRI); One Hundred Mile
swamp, near Undara resort, E of Mt Surprise, Jul 1998,
Bean 13758 & Fox (BRI, MEF); c. 1 mile [1.6 km]
south of Bamaga, Jul 1977, Carlquist 15250 (BRI,
RSA); 0.5 km SE of Kimba HS, Jun 1981, Clarkson
3737 (BRI, DNA, K, MO, NSW, PERTH, QRS, RSA);
Weipa, Vyse Crossing, Jul 1980, Morton 656 (BRI,
MEL). North Kennedy District: Walkers Creek, Mount
Elliot, S of Townsville, Aug 1991, Bean 3616 (BRI);
Mt Garnet-Wairuna road, 4.8 km S of Princess Hills
T/0,Aug 1997, Bean 12168 (BRI). Leichhardt District:
Mt Rose, Taroom district, Jan 1996, Fensham 2500
(BRI).
Reconstituted or spirit material examined:
Bean 12100 (2 fls); Bean 12114 (2 fls); Bean
12168 (2 fls); Bean 12192 (2 fls); Clarkson 3737
(2 fls); Fensham 2500 (1 fl).
Distribution and habitat: Stylidium
rotundifolium is widespread (though sporadic)
from the Kimberley region ofWesternAustralia
to north-eastern Queensland, with a remarkable
disjunct occurrence near Taroom (Map 18). It
occurs in damp sandy soil, on receding
waterholes, on creekbanks or in Melaleuca
woodlands. The occurrence near Taroom is
associated with mound-springs, locally known
as “boggomosses”.
Phenology: Flowers and capsules have been
recorded from April to October, with one record
for January and one for February.
Affinities: S. rotundifolium is closest to S.
dunlopianum. See notes under that species.
Notes: Specimens of this species collected by
Banks and Solander were available to Robert
Brown when drawing up his description, hence
the need to choose a lectotype. The lectotype
locality for S. rotundifolium is Curtis or Facing
Island near Gladstone but it has not been found
there, nor indeed in the Port Curtis Pastoral
District, since Brown’s collection in 1802.
Fitzgerald (1918) did not explain why he
considered S. irriguum to be specifically
distinct from S. rotundifolium , but by
comparing the description given under both
names, it seems that the distinction was based
on: the pale yellow to white corolla for
S. irriguum (vs. pink with red blotches for
633
Bean, Stylidium subgen. Andersonia
S. rotundifolium ) and a shorter capsule.
However pale yellow to white is the usual flower
colour for S. rotundifolium , and the capsule
length given for S. irriguum (to nearly 0.75
inches [19 mm]) is within the normal range for
S. rotundifolium.
S. reductum represents juvenile forms of
S. rotundifolium which are only 1-flowered.
There is continuous variation between this form
and typical S. rotundifolium.
Conservation status: S. rotundifolium is a
widespread species, and not considered to be
rare or threatened.
31. Stylidium dunlopianum Carlquist, Aliso 9:
431 (1979). Type: Northern Territory.
Cultivated in Darwin, ex pennanent spring
near Munmarlary Station, [ 12° 2- S 132°
3-Tf|, 21 July 1978, C.R. Dunlop 4998
(holo: RSA; iso: ?CANB,w.v., DNA, ?K,
n.v.).
Perennial, 15-50 cm high. Glandular hairs 0.05-
0.1 mm long; glands globose, dark. Stem base
thickened. Stems elongate or leaves in basal
rosette, glabrous. Leaves 6-12 per plant,
scattered along stems, obovate, 20-60 mm long
including petiole, 7-19 mm wide, glabrous; apex
obtuse; base cuneate; margins entire. Petioles
8-35 mm long. Scapes 1-5 per plant, 0.5-0.9
mm in diameter, glabrous; sterile bracts absent.
Inflorescences 13-40 cm long, determinate,
monochasially cymose; branches glabrous.
Bracts linear or ovate, 1-2.5 mm long, glabrous,
acute ormucronate. Bracteoles absent. Pedicels
absent or rudimentary. Hypanthium linear,
glandular-hairy at distal end only. Sepals ovate,
fused into 2 entire lips, 1.9-2.5 mm long, 1.1-
1.6 mm wide, glandular-hairy, obtuse. Corolla
pink or mauve, glandular-hairy on tube and
petals; tube 4.5-6 mm long, with sinus on
anterior side only. Paracorolla absent. Labellum
attached at base of anterior sinus of corolla
tube, lanceolate, 0.7-0.9 mm long, thick,
glabrous, acuminate, terminal appendage
usually present; 0.4—0.5 mm long. Petals all free,
A1+A2+P1+P2. Anterior petals 1.1-2.1 mm
long, 1.1-2 mm wide, entire, obtuse. Posterior
petals 1.3-2.1 mm long, 1-1.8 mm wide, entire,
obtuse. Column 7-8.5 mm long, conspicuously
dilated near distal end forming a pouch for the
stigma and anthers, glabrous; lateral lobes
absent. Corona absent. Capsule linear, 20-32
mm long excluding sepals, 0.5-0.6 mm wide,
without raised longitudinal ribs; halves
detaching distally, not recurved. Seeds
ellipsoidal, 0.2-0.25 mm long, brown; surface
convex, colliculate.
Specimens examined: Western Australia. Gibb
River-Kalumburu road, Ngolalah Creek, 38.6 km NE
of Mitchell River turnoff, Jun 1976, Beauglehole
52145 (DNA); King Edward River, c. 50 km NE of
Mitchell River HS., Aug 1978, Beauglehole 29112 &
Errey 2853 (DNA). Northern Territory. Edith Falls,
Jul 1965, Beauglehole 43214 (DNA); 2 km W of
East Alligator River crossing, Aug 1978, Beauglehole
58648 & Errey 2348 (DNA); Horn Billabong, Sep
1946, Blake 17009 (BRI); Docherty Ck, Sep 1967,
Byrnes NB497 (DNA); 47 miles [75 km] N of
Oenpelli, Jul 1961, Chippendale 8115 (DNA);
Ningalaye Brook, 30 km WNW of Cannon Hill
Ranger Station, May 1980, Craven 6172 (CANB,
DNA); 65 km NE of Pine Creek, Nov 1980, Dunlop
5615 (DNA); Maxwell Creek, Melville Island, Nov
1983, Dunlop 6547 & Wightman (DNA); Point Stuart,
Swim Creek, May 1987, Dunlop 7012 (DNA); S of
Koolpinyah Stn., Sep 1995, Harwood B153 (DNA);
26 km E of Goomadeer River at Nungbalgari Creek,
Oct 1981, Henshall 3816 (DNA); Katherine Gorge
N.P., amphitheatre, Jun 1981, King s.n. (DNA);
headwaters of Florence Creek, May 1989, Leach 2583
& Dunlop (DNA); Oenpelli, Oct 1948, Specht 1313
(BRI); Old Oenpelli road, just N of Lonely Rock,
Magela Ck, Sep 1980, Waterhouse 11210/1 (BRI).
Reconstituted or spirit material examined:
Beauglehole 29112 & Errey 2853 (2 fls ), Dunlop
7012(1 f[)-Leach2583 &Dunlop(\ fl).
Distribution and habitat: Stylidium
dunlopianum is distributed from the Mitchell
River area of Western Australia to Kakadu
Natonal Park in the Northern Territory (Map
19). It inhabits seepage areas in Melaleuca
leucadendra swamps or near rainforest, or on
creekbanks, in sand or black organic soil.
Phenology: Flowers and capsules have been
recorded from May to November.
Affinities: S. dunlopianum is closely related
to S. rotundifolium but differs by its perennial
habit with thickened stem base (annual with
unthickened stem base for S. rotundifolium ),
scapes 0.5-0.9 mm wide (0.2-0.4 mm wide for S.
rotundifolium ), sepals 1.9-2.5 mm long (1.0-
1.8 mm long for A rotundifolium ), corolla tube
4.5-6 mm long (2-3 mm long for S.
rotundifolium ), labellum 0.7-0.9 mm long (0.4-
634
0.6 mm long for S. rotundifolium) and column
7-8.5 mm long (3.5-5 mm long for S.
rotundifolium).
Conservation status: Data deficient (IUCN
1994).
32. Stylidium fimbriatum Lowrie & Kenneally,
Nuytsia 10: 425 (1996). Type: Western
Australia. Peter Lacy’s camp, 73 km
WNW of Mount Elizabeth homestead,
16° 00’S 125° 20’E, August 1993, M.D.
Barrett230 (holo: PERTH, fide Lowrie &
Kenneally (1996) n.v.\ iso: MEL,w.v.).
Annual, 15-30 cm high. Stem base not
thickened. Stems compressed (with leaves in
basal rosette). Leaves c. 16 per plant,
oblanceolate, 5-20 mm long including petiole,
2-7 mm wide, glabrous; apex obtuse; base
cuneate; margins entire. Scapes c. 2 per plant,
glabrous; sterile bracts absent. Inflorescences
15-30 cm long, determinate, monochasially
cymose; branches glabrous. Bracts linear or
lanceolate, 1-2 mm long, acute or mucronate.
Bracteoles absent. Pedicels absent or
rudimentary. Hypanthium linear, glandular-
hairy at distal end only or glandular-hairy
throughout. Sepals ovate or elliptical, fused into
2 entire lips, 1.5-2 mm long, glandular-hairy,
obtuse. Corolla pink, glandular-hairy on tube
and petals; with sinus on anterior side only.
Paracorolla discontinuous, thick, glabrous, c.
1.5 mm high. Paracorolla lobes or appendages
13-17, all similar, acute, none opposite the
anterior petals, 13-17 opposite the posterior
petals. Paracorolla glands absent. Labellum
attached at base of anterior sinus of corolla
tube, lanceolate, c. 0.6 mm long, thick, glabrous,
acuminate, terminal appendage usually present.
Petals all free,Al+A2+Pl+P2. Anterior petals
c. 3.5 mm long, c. 2 mm wide, bilobed, obtuse.
Posterior petals c. 5.5 mm long, c. 2.5 mm wide,
bilobed, obtuse. Column c. 10 mm long,
conspicuously dilated near distal end forming
a pouch for the stigma and anthers, glabrous;
lateral lobes absent. Corona absent. Capsule
linear, 25-50 mm long excluding sepals, 0.6-0.7
mm wide, without raised longitudinal ribs.
Seeds c. 0.2 mm long, yellow or brown; surface
convex.
Distribution and habitat: Stylidium
fimbriatum is reportedly confined to the
Austrobaileya 5(4): 589-649 (2000)
Bachsten Creek area in the Kimberley region of
We stern Australia (Map 19), where it grows in
seasonally wet herbfields (Lowrie and
Kenneally 1996).
Note: No material of this species was available
for examination. The description above is
based on Lowrie & Kenneally (1996).
E. Stylidium sect. Alsinoida (Mildbr.)A.RBean
comb, et stat. nov.
S. subg. Alsinoida Mildbr. in Engl., Pflanzenr.
35: 40 (1908), as ‘Alsinoides ’ . Type:
S. alsinoides R.Br.
Leaves cauline, alternate. Scapes absent. Bracts
opposite, often leaf-like. Petals laterally fused
(i.e. adjacent posterior and anterior petals
fused), often acute. Paracorolla absent.
Labellum attached at top of corolla tube.
Capsules coherent at apex, shedding medially;
seeds ridged or lacunose, colliculate.
5 species; Malesia, New Guinea, northern
Australia.
33. Stylidium alsinoides RBr., Prodr. 572 (1810);
Candollea alsinoides (R.Br.) F.Muell.,
Syst. census Austral, pi. 86 (1883). Type:
[Queensland. Cook District:] Endeavour
River, [15°2-’S 145° l-’E, June- July 1770],
J. Banks & D. Solander (holo: ?BM,«.v.).
Illustration: R. Erickson, Triggerplants, plates
51,56(1958).
Annual, 18-30 cm high. Glandular hairs 0.05-
0.1 mm long; glands globose or capitate, dark.
Stem base not thickened. Stems elongate,
glabrous. Leaves 20-100 per plant, scattered
along stems, elliptical or obovate, 5.5-14 mm
long including petiole, 3-8 mm wide, glabrous;
apex acute; base cuneate; margins entire.
Petioles 0-2.5 mm long. Scapes absent.
Inflorescences 5-15 cm long, determinate,
monochasially cymose; branches glabrous.
Bracts lanceolate or ovate, 5.5-9 mm long,
glabrous, acute. Bracteoles absent. Pedicels
absent or rudimentary. Hypanthium linear,
glabrous throughout or glandular-hairy at distal
end only. Sepals deltate, with 3 free and 2 fused
for more than half their length, 1.3-2 mm long,
0.3-0.5 mm wide, glabrous, acute. Corolla white,
635
Bean, Stylidium subgen. Andersonia
glabrous, or sparsely glandular-hairy on tube
and petals; tube 0.6-0.9 mm long, without
sinus. Paracorolla absent. Labellum attached
at top of corolla tube, ovate, 0.3-0.5 mm long,
thick, glabrous, obtuse or acuminate, terminal
appendage absent. Petals laterally fused,
(A1 &P 1 )+(A2&P2). Anterior petals 1.7-2.5 mm
long, 0.9-1.2 mm wide, entire, acute or obtuse.
Posterior petals 2-2.5 mm long, 0.8-1 mm wide,
entire, acute or obtuse. Column 2.5-3 mm long,
of uniform width throughout, glabrous; lateral
lobes absent. Corona extending beyond
anthers. Capsule linear, 16-30 mm long
excluding sepals, 0.7-1.2 mm wide, without
raised longitudinal ribs; halves coherent
distally. Seeds ellipsoidal, 0.4-0.5 mm long,
brown; surface lacunose, colliculate.
Selected specimens: Philippines. Philippine Islands,
1906, Loher 6478 (K); Neuva Vizcaya, Luzon, [16°N
121°E], Jan 1913, McGregor 20136 (BM, P); Mt
Irig, Rizal Province, Luzon, [14°N 121°E], Feb 1923,
Ramos 41978 (BM, BRI, L); Mt Marayep, Zambales
province, Luzon, [15°N 120°E], Dec 1924, Ramos &
Edano 44782 (BM, P); Setio, Apulul, Barrio Amungan
Iba, Zambales, [15°N 119°E], Dec 1954, Santos 6105
(L). Sulawesi. Lepo Lepo pr. Kendari, SE Celebes,
[4°S 122°E], Jun 1874, Beccari (L); Lombasang, SW
Celebes, [5°S 119°E], Apr 1921, Bunnemeijer 11091a
(L). Irian Jaya. Merauke, [8°S 140°E], Jul 1923,
Vertenten (BRI, L, P). New Guinea. Wuroi, Oriomo
River, [8°S 143°E, Jan-Mar 1934, Brass 6070 (BRI,
L); Wassi Kussa River, Morehead subdistrict, 8S 141E,
Jul 1968, Henty & Katik NGF38734 (BRI, L); c. 3
miles [5 km] S of Morehead Patrol post, along road
to Tonda, 9S 141E, Sep 1967, Pullen 7242 (L).
Queensland. Cook District: 20.4 km from Peninsula
Development road, towards Iron Range, Jul 1998,
Bean 13606 (BRI, MEL); Yarrabah, Jun 1935, Blake
9642 (BRI); Isabella Falls near Cooktown, May 1970,
Blake 23433 (BRI); Chester River campsite, Jul 1978,
Clarkson 2418 (BRI, K, L, MO, NSW); Scrubby Creek,
N of Silver Plains, Aug 1978, Clarkson 2444 (BRI, K,
NSW); 16.4 km S of aboriginal settlement at Mapoon,
on the road to Weipa, Aug 1983, Clarkson 4943 (BRI,
PERTH, QRS); Finch Bay, Cooktown, May 1993,
Clarkson 10084 & Neldner (BRI, DNA, K, MBA,
PERTH); Endeavour River, Jun-Jul 1819, Cunningham
277 (BM, BRI); west Claudie River falls, 2.5 km NE
of Mt Tozer, May 1992, Fell DF2613 (BRI); Fred’s
Creek, 2.5 km SE of Kennedy Hill, Jul 1991, Forster
PIF8813 (BRI, DNA, MEL); Badu Island, Torres Strait,
Oct 1979, Garnett 220 (BRI); 2 km S of Cooktown,
Jul 1991, Sharpe 5100 8c Levine (BRI, CANB, MEL,
NSW). North Kennedy District: Dunk Island, E of
airstrip, Aug 1959, Adams 20012 (BRI); Travelling
Dairy Creek, 9 km W of Tully, Sep 1992, Bean 4996
(BRI); Hinchinbrook Is., S of Kirkville Hills, Aug 1970,
Everist 9662 (BRI); 50 km NNW of Ingham and 3.5
km E of Bruce Highway, Aug 1976, Lazarides 8125
(BRI, CANB); 3 km W of Cardwell, Sep 1976, Williams
76074 (BRI).
Reconstituted or spirit material examined:
Clarkson 2444 (2 fls); Clarkson 4943 (2 fls);
Clarkson 10084 8cNeldner{ 2 fls);/W/DF2613
(lfl).
Distribution and habitat: Stylidium
alsinoides is widespread, being recorded from
the island of Luzon in the Philippines, southern
parts of Sulawesi, southern New Guinea and
north Queensland (Map 1). In Australia, it
typically grows in sandy soil in swamps
dominated by Melaleuca quinquenervia ,
which are moist even in the dry season. It may
also grow on creekbanks withM leucadendra ,
or in rock crevices where water seepage is
available.
Phenology: Flowers and capsules have been
recorded mostly from April to September for
Australia; throughout the year for Malesia.
Affinities: S. alsinoides is most closely related
to S.fluminense. See notes under that species.
Note: The bracts of S. alsinoides and its
relatives are very large and resemble the leaves.
In fact the transition from stem to inflorescence
can sometimes only be determined by the
phyllotaxis; the leaves are alternate while the
bracts are opposite. The seemingly solitary
axillary inflorescences are in fact cymose and
monochasial as in most other species.
Conservation status: Data deficient (IUCN
1994).
34. Stylidium fluminense F.L.Erickson &
J.H.Willis, Victorian Naturalist 83: 108
(1965). Type: Western Australia. Dales
Gorge, Hamersley Range, [22° 30’S 118°
36’E], 3 July 1958, F.L. Erickson (holo:
MEL; iso: K, MEL, PERTH).
Illustration: J.R. Wheeler (ed.), FI. of Kimb.
Region 876, t. 270D(1992).
Annual, 15-30 cm high. Glandular hairs 0.025-
0.05 mm long; glands globose, dark. Stem base
not thickened. Stems elongate, glabrous.
Leaves 10^10 per plant, scattered along stems,
elliptical, 5-19 mm long including petiole, 1.5-
636
6 mm wide, glabrous; apex acute; base cuneate;
margins entire. Petioles 0-2 mm long. Scapes
absent. Inflorescences 5-11 cm long,
determinate, monochasially cymose; branches
glabrous. Bracts linear or lanceolate, 2.5-10 mm
long, glabrous, acute. Bracteoles absent.
Pedicels absent or rudimentary. Hypanthium
linear, glandular-hairy throughout. Sepals
oblanceolate, with 3 free and 2 fused for more
than half their length, 3-4.1 mm long, 0.4-0.5
mm wide, glabrous, acute. Corolla pink or mauve
or red, glandular-hairy on tube and petals; tube
1.1-1.5 mm long, without sinus. Paracorolla
absent. Labellum attached at top of corolla
tube, ovate, 0.6-0.7 mm long, thick, glandular-
hairy, obtuse or acuminate, terminal appendage
usually present; 0-0.1 mm long. Petals laterally
fused, (A 1&P1)+(A2&P2). Anterior petals 5.2-
5.5 mm long, 1.9-2.3 mm wide, entire, acute.
Posterior petals 5.2-5.5 mm long, 1.9-2.3 mm
wide, entire, acute. Column 5.5-6 mm long, of
uniform width throughout, glabrous; lateral
lobes absent. Corona absent. Capsule linear,
11-16 mm long excluding sepals, 0.7-0.9 mm
wide, without raised longitudinal ribs; halves
coherent distally. Seeds globose or ellipsoidal,
0.25-0.3 mm long, brown; surface convex,
colliculate.
Specimens examined: Western Australia. Dales
Gorge, near Fortescue Falls, Hamersley Range, Aug
1965, Beauglehole 11496 (DNA, MEL); Wittenoom
Gorge, c. 10 km S of Wittenoom P.O., Aug 1974,
Beauglehole 48874 (DNA); Exmouth Gulf, 1895,
Cusack s.n. (MEL); Harding River, 1895, Cusack 140
(MEL); Dale’s Gorge, 40 miles [64 km] E of
Wittenoom Gorge, Jul 1956, Meston s.n. (MEL);
Dale’s Gorge, Hamersley Range, Oct 1989,
Nordenstam & Anderberg 331 (MEL). Northern
Territory. 50 km NNW of Inverway homestead, Jun
1974, Latz 5419 (DNA).
Reconstituted or spirit material examined:
Beauglehole 11496 (2 fls); Beauglehole 48874
(lfl).
Distribution and habitat: Stylidium
fluminense is known from the Hamersley
Ranges in We stern Australia and from one site
in Northern Territory, and there are historical
collections from the Harding River (near
Karratha) and from the Exmouth Gulf (Map 17).
It grows in sheltered sites with permanent
moisture in sandstone gorges.
Austrobaileya 5(4): 589-649 (2000)
Phenology: Flowers and capsules have been
recorded from June to October
Affinities: S.fluminense is most closely related
to S. alsinoides , but differs by the oblanceolate,
acute sepals 3.0-4.1 mm long (debate, obtuse,
1.3-2.0 mm long for S. alsinoides ); corolla tube
1.1-1.5 mm long (0.7-0.8 mm long for S.
alsinoides ); anterior petals 5.2-5.5 mm long
(1.7-2.5 mm for S. alsinoides)., column 5.5-6
mm long (2.5-3 mm for S. alsinoides ); capsule
11-16 mm long (16-30 mm for S. alsinoides)
and seeds 0.25-0.3 mm long (0.4-0.5 mm for S.
alsinoides).
Conservation status: Data deficient (IUCN
1994).
35. Stylidium tenerrimum F.Muell., Fragm. 1:
150 (1859); Candollea tenerrima
(F.Muell.) F.Muell., Syst. census Austral,
pi. 86 (1883). Type: [NorthernTerritory],
between Providence Hill and Macadam’s
Range, [14°—’S129°—’E], October 1855,
F. Mueller (lecto: MEL [MEL 1061526]),
here chosen.
Stylidium mitrasacmoides F.Muell.,
Fragm. 1: 150 (1859). Type: [Northern
Territory], banks of Victoria River, near
Palm Island, [15°—’S 129°—’E, 1855-6],
Flood, n.v. (not located).
Stylidium evolutum Carlquist, Aliso 9: 309
(1978), syn. nov. Type: Northern Territory,
road west from Stuart Highway, opposite
juncture with the Shoal Bay road, [12° 3-
’S 131° O-’E], 25 June 1977, A Carlquist
15190 (holo:RSA).
Annual, 4-30 cm high. Glandular hairs 0.025-
0.05 mm long; glands globose, dark. Stem base
not thickened. Stems elongate, glabrous.
Leaves 5-24 per plant, scattered along stems,
linear or oblanceolate or debate, 1.2-4.8 mm
long, 0.3-0.8 mm wide, glabrous; apex obtuse,
or acute; base truncate; margins entire. Petioles
absent. Scapes absent. Inflorescences 4-11 cm
long, determinate, monochasially cymose;
branches glabrous. Bracts linear, 2-4.5 mm long,
glabrous, acute. Bracteoles absent. Pedicels
absent or rudimentary or present, 0-5 mm long,
glabrous. Hypanthium linear, glabrous
637
Bean, Stylidium subgen. Andersonia
throughout or glandular-hairy throughout.
Sepals oblanceolate or ovate, with 3 free and 2
fused for more than half their length, 2.1-2.8
mm long, 0.3-0.5 mm wide, glabrous, acute.
Corolla white and red, glandular-hairy on tube
and petals or glandular-hairy on petals only;
tube 0.5-0.7 mm long, without sinus.
Paracorolla absent. Labellum attached at top
of corolla tube, ovate, 0.3-0.5 mm long, thick,
glabrous, obtuse or acuminate, terminal
appendage usually present; 0-0.1 mm long.
Petals laterally fused, (A1&P1)+(A2&P2).
Anterior and posterior petals entire, acute or
obtuse. A1 and PI 2-2.6 mm long, 1.2-1.5 mm
wide; A2 and P2 3.3-44 mm long, 1.7-2.3 mm
wide. Column 4-4.5 mm long, of uniform width
throughout, glabrous; lateral lobes absent.
Corona absent. Capsule ellipsoidal or linear, 4.5-
9 mm long excluding sepals, 1-1.6 mm wide,
without raised longitudinal ribs; halves
coherent distally. Seeds ellipsoidal, 0.45-0.5 mm
long, brown; surface lacunose, colliculate.
Specimens examined: Northern Territory. 2 km W
of Stuart Highway on road opposite the juncture of
Shoal Bay road with Stuart Highway, Jun 1978,
Carlquist 15479 (DNA); between Elizabeth River and
RAAF base, 23 miles [37 km] S of Darwin, Jun 1978,
Carlquist 15455 (DNA); Melville Island, McClear
Creek T/O, Jun 1987, Clark 1237 (DNA); Howard
Springs, May 1995, Egan 4984 (DNA); Port Darwin,
1885, Holtze 489 (MEL); Port Darwin, 1885, Holtze
506 (MEL); Port Darwin, 1888, Holtze s.n. (MEL);
near Darwin, undated, Holtze 1170 (MEL); RAAF
Base, swamp off Amy Johnson Drive, Darwin, Apr
1983, King 323 (DNA); 1 mile [1.6 km] SE of
McMinns Lagoon, Aug 1971, Must 758 (DNA);
Ironstone Knob area, behind Holmes Jungle, Jun 1982,
Rankin 2598 (CANB, DNA); Point Stuart, Swim Creek,
Jun 1987, Russell-Smith 5568 & Lucas (DNA); Port
Darwin, undated, Schultz 349 (MEL); North Australia,
1886, Tenison-Woods & Holtze 489 (MEL).
Reconstituted or spirit material examined:
Egan 4984 (3 fls ), Russell-Smith 5568 (3 fls).
Distribution and habitat: Stylidium
tenerrimum is apparently reasonably common
around Darwin, as there are numerous
collections (both old and new) from that area.
However, it has not been recollected from the
type area near the Victoria River (Map 14). It
grows in sandy soils which remain moist after
the wet season, amongst grasses and sedges.
Associated trees include Pandanus spp.,
Melaleuca spp. and Grevillea pteridifolia.
Phenology: Flowers and capsules have been
recorded from April to August
Notes: S. tenerrimum is related to S. alsinoides.
S. tenerrimum is distinctive because of the
asymetrical petals (A1 andA2 of different sizes;
PI and P2 of different sizes), comparatively
broad capsules, and the sometimes pedicellate
flowers and capsules (pedicels up to 4 mm
long). Two specimens of S. tenerrimum
collected by Mueller have been seen, one at
MEL and one at BM. It is uncertain whether
these were both from the same collection. The
MEL specimen is chosen as lectotype, as it is
the better specimen.
The type of S. mitrasacmoides could not
be found, and the application of the name is
somewhat uncertain.
Conservation status: Data deficient (IUCN
1994).
36. Stylidium javanicum Slooten, Bull. Jard.
Bot. Buitenzorg Ser. Ill, 14: 173 (1937).
Types: Java. Plosokerep near halting-
place Terisi, partitions 7,3 and 5 of forest-
section Indramajoe, [6° —* S 108° —’E], 3
May 1936, C.G.G.J. van Steenis 8213 &
D.F. van Slooten (syn: ?BO, n.v., MEL!);
Soemba. East-Soemba, Lea plain, [9°—’S
120°—’E], 28 May 1936, C.AET de Voogd
2512 (syn: ?BO,«.v.).
Illustration: R. Erickson, Triggerplants, plate
56(1958).
Annual, 5-21 cm high. Glandular hairs 0-0.05
mm long; glands globose, dark. Stem base not
thickened. Stems elongate, glabrous. Leaves
10-30 per plant, scattered along stems, elliptical
or obovate, 1.7-4.5 mm long, 0.9-2.1 mm wide,
glabrous; apex obtuse, or acute; base obtuse,
or cuneate; margins entire. Petioles absent.
Scapes absent. Inflorescences 5-9 cm long,
determinate, monochasially cymose; branches
glabrous. Bracts lanceolate or ovate, 2-4 mm
long, glabrous, acute. Bracteoles absent.
Pedicels absent or rudimentary. Hypanthium
linear, glabrous throughout or glandular-hairy
at distal end only. Sepals oblanceolate or
elliptical, with 3 free and 2 fused for more than
half their length, 1.6-2.5 mm long, 0.2-04 mm
wide, glabrous, obtuse or acute. Corolla pink
638
or mauve, glandular-hairy on tube only; tube
0.7-0.8 mm long, without sinus. Paracorolla
absent. Labellum attached at top of corolla
tube, ovate or orbicular, 0.7-0.8 mm long, thin,
glabrous, obtuse or acute, terminal appendage
absent. Petals laterally fused,
(A1 &P1 )+(A2&P2). Anterior petals 2.4-2.8 mm
long, 1-1.4 mm wide, entire, acute. Posterior
petals 2.4-2.8 mm long, 1-1.4 mm wide, entire,
acute. Column 3.5-4 mm long, of uniform width
throughout, glabrous; lateral lobes absent.
Corona absent. Capsule linear, 10-16 mm long
excluding sepals, 0.5-0.8 mm wide, without
raised longitudinal ribs; halves coherent
distally. Seeds ellipsoidal, 0.25-0.3 mm long,
brown; surface lacunose, colliculate.
Specimens examined: Java. Plosokerep near halting-
place Terisi, partitions 7, 3 and 5 of forest-section
Indramajoe, May 1936, van Steenis 8213 & van
Slooten (MEL). New Guinea. Rubulogo Creek c. 18
miles [29 km] N of Port Moresby, [9°S 147°E], Apr
1967, Pullen 6631 (BRI, CANB, L).
Reconstituted or spirit material examined:
Pullen 6631 (2 fls).
Distribution and habitat: Stylidium javanicum
has been found in Java, East Soemba Island
and from eastern New Guinea (Map 2). It is
recorded from open boggy depressions or wet
grass-fields, from 20-500 metres altitude.
Phenology: Flowers and capsules have been
recorded for April and May.
Affinities: S.javanicum is most closely related
to S. cordifolium , but differs by the leaves
being 1-1.7 mm wide (2.9-8 mm wide for S.
cordifolium ); leaf base cuneate (cordate for S.
cordifolium ); sepals 2. l-2.5x0.2-0.3 mm (2.6-
3.5x0.4-07 mm for S. cordifolium ); capsules
0.5-0.8 mm wide (1-1.5 mm for S. cordifolium)
and seeds 0.25-0.3 mm long (0.4-0.5 mm long
for S. cordifolium ).
Conservation status: Not evaluated.
37. Stylidium cordifolium W.Fitzg., J. Proc. Roy.
Soc. Western Australia 3: 217 (1918).
Types: Isdell and King Rivers, 1905-6, W. V.
Fitzgerald (syn, n.vf Messmate Creek,
1905-6, W.V. Fitzgerald '(syn, n.v.);
between Isdell Range, 1905-6, W.V.
Fitzgerald (syn, n.v ).
Austrobaileya 5(4): 589-649 (2000)
Stylidium alsinoides var. cordifolium
Ewart, Jean White & B.Wood, Proc. Roy.
Soc. Victoria ser. 2,23:299 (1911). Types:
Port Darwin, 1890, M. Holtze 1171 (syn:
MEF); Isdell River; Graces Knob;
Messmate Creek in Packhorse range;
between Isdell Range and Mt Bartlett
(syn, n.v.).
Annual, 15—45 cm high. Glandular hairs 0.025-
0.1 mm long; glands globose, dark. Stem base
not thickened. Stems elongate, glabrous.
Leaves 8^10 per plant, scattered along stems,
obovate or orbicular, 3.5-8 mm long, 3.0-8 mm
wide, glabrous; apex acute; base cordate;
margins entire. Petioles absent. Scapes absent.
Inflorescences 4-13 cm long, determinate,
monochasially cymose; branches glabrous.
Bracts lanceolate or ovate, 2-5 mm long,
glabrous, acute. Bracteoles absent. Pedicels
absent or rudimentary. Hypanthium linear,
glabrous throughout or glandular-hairy at distal
end only. Sepals oblanceolate, with 3 free and
2 fused for more than half their length, 2.6-3.5
mm long, 0.4-0.7 mm wide, sparsely glandular-
hairy or glabrous, acute. Corolla pink or red,
glandular-hairy on tube and petals, or on petals
only; tube 0.7-1.3 mm long, without sinus.
Paracorolla absent. Labellum attached at top
of corolla tube, ovate, 0.5-0.7 mm long, thick or
thin, glabrous or glandular-hairy, obtuse or
acuminate, terminal appendage absent. Petals
laterally fused, (A1&P1)+(A2&P2). Anterior
petals 3.3—4.5 mm long, 1-1.8 mm wide, entire,
acute. Posterior petals 3.4-5 mm long, 1-1.8
mm wide, entire, acute. Column 3.5-5 mm long,
of uniform width throughout, glabrous; lateral
lobes absent. Corona absent. Capsule linear,
7.5-18 mm long excluding sepals, 1-1.5 mm
wide, without raised longitudinal ribs; halves
coherent distally. Seeds ellipsoidal, 0.4-0.5 mm
long, brown; surface lacunose, colliculate. Fig.
8A-D.
Specimens examined: Western Australia. Chapman
River, 55 km by road WSW of Karungi Station HS,
May 1976, Beauglehole 51509 (DNA); Barnett
Gorge, c. 250 km SW of Wyndham, Jun 1976,
Beauglehole 52341 (DNA); Gibb River road, 1.5 km
W of Lennard River Gorge turnoff, Jul 1974, Carr
4108 & Beauglehole 47886 (DNA); Vansittart Bay,
north Kimberley, May 1984, Chesterfield 352 &
Forbes (DNA, MEL); c. 12 km W of Mt Hann,
Bean, Stylidium subgen. Andersonia
639
Fig 8. A-D: Stylidium cordifolium. A. habit><0.7; B. anterior view of flower and hypanthiumx5; C. developing
capsulex3; D. seedx20. E-H. Stylidium longissimum. E. habitx0.5; F. anterior view of flowerx5; G. developing
capsulex4; H. seedx40. A-D: Bean 12181; E, G-H: Bean 13671; F: Clarkson 5382.
640
Gardner Plateau, May 1993, Cowie 4313 & Stewart
(DNA); unnamed creek running into Pauline Bay,
north Kimberley, May 1984, Forbes 2169 (MEL).
Northern Territory. Berrimah Lagoon, S of Darwin,
May 1978, Carlquist 15384 (BRI, DNA); Site 55,
Mary River, May 1989, Clark 1753 (DNA); Melville
Island, Soldier Point road camp, Jun 1987, Clark 1209
& Orr (DNA); Berrimah, Darwin, Apr 1976, Dunlop
4140 (DNA, NSW); Kakadu N.P, Mar 1982, Dunlop
6234 (DNA); c. 8 km NNE of Jabiru, Apr 1995, Egan
4846 & Knox (DNA); Howard Springs, May 1995,
Egan 4985 (DNA); Port Darwin, 1890, Holtze 1171
(MEL); Nabarlek, Apr 1979, Rankin 2035 (BRI,
DNA, K); Adelaide River, Arnhem Hwy, Apr 1980,
Rankin 2248 (DNA). Queensland. Cook District:
4.8 km N of Kennedy River crossing on Peninsula
Development road, Jun 1981, Clarkson 3685 (BRI,
K, MO, NSW, PERTH, QRS, RSA); Bulleringa NP, 80
km NW of Mt Surprise, track to Red River past
Donkey Spring, Apr 1998, Forster PIF22595 & Booth
(BRI); Granite Creek road, below Walsh Bluff, Apr
1981, Gray 1948 (BRI, QRS); Dixie station. Cook
shire, Jun 1979, Weaver 33 (BRI). North Kennedy
District: 4 km west of Nymbool, via Mt Garnet, Aug
1997, Bean 12181 (BRI).
Reconstituted or spirit material examined:
Bean 12181 (2 fls); Clarkson 3685 (2 fls );Egan
4985 (3 fls); Gray 1948 (1 fl).
Distribution and habitat: Stylidium
cordifolium is distributed across tropical
Australia (Map 20). It often grows in or on the
margins of Melaleuca viridiflora swamps, in
moist sand. It can sometimes occur on moist
sandy creekbanks with Pandanus spp.,
grasses and sedges.
Phenology: Flowers and capsules have been
recorded from March to August
Affinities: S. cordifolium is closest to S.
javanicum. See notes under that species.
Conservation status: Data deficient (IUCN
1994).
Acknowledgements
I am grateful to the Directors of the following
Herbaria for the loan of specimens: AAU, BM,
CANB, DNA, K, L, MEL, MO, NSW, P and
RSA; to Laurie Jessup for assisting me in
mastering the DELTA program; to Neil Snow
and Wayne Harris for guidance with cladistic
analysis; to Jian Wang for translating
Austrobaileya 5(4): 589-649 (2000)
descriptions of Chinese Stylidium species; to
Keith McDonald for taking me to Cape Melville
to find ‘pink bits’; to Chris Lyons for collecting
the excellent type material of S. confertum\ to
Paul Forster for collecting numerous stylidia
from various parts of Queensland; to Will Smith
for the illustrations and distribution maps; and
to Peter Bostock for the Latin diagnoses.
References
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Bhaskar, V & Kushalappa, C.G. (1992). Stylidium
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species, Modes of speciation on the sandstone
plateau, and comments on floral mimicry. Aliso
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phylogenies: an approach using the bootstrap.
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fimbriatum (Stylidiaceae), a new tropical species
of triggerplant from the Kimberley, Western
Australia. Nuytsia 10(3): 425-7.
Maddison, W.P. & Maddison, D.R. (1992) MacClade
Version 3 - Analysis of Phytogeny and
Character Evolution. Sinauer Associates:
Massachusetts, U.S.A.
Mckee, H.S. (1963). The Bleeser Botanical Collection
from Northern Australia. Contributions from
the New South Wales National Herbarium 3(4):
233-4.
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Engelmann.
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14: 169-74.
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Malesiana Ser. 1, 4: 529-32. Djakarta:
Noordhoff-Kolff N.V.
Swofford, D.L. (1993). PAUP: Phylogenetic Analysis
Using Parsimony. Version 3.1.1 Washington,
D.C.: Smithsonian Institution.
Vallance, T.G. (1990). Jupiter Botanicus in the bush:
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Dassanayake (ed.), A Revised Handbook to the
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642 Austrobaileya 5(4): 589-649 (2000)
Map 2. Distribution of O# Stylidium uliginosum, A A S.javanicum.
Bean, Stylidium subgen. Andersonia
643
110 115 120 125 130 135 140 145 150 155
Map 4. Distribution of O# Stylidium tenerum, A S.divergens.
644
Austrobaileya 5(4): 589-649 (2000)
Map 5. Distribution of O# Stylidium muscicola, A A S.ensatum.
Map 6. Distribution of • Stylidium accedens.
Map 7. Distribution of O# Stylidium capillare.and AS.simulans.
Bean, Stylidium subgen. Andersonia
645
Map 8. Distribution of O# Stylidium lobuliflorum.
Map 9. Distribution of O# Stylidium schizanthum
646
Austrobaileya 5(4): 589-649 (2000)
Map 10. Distribution of O • Stylidium pachyrrhizum, and A S.stenophyllnm.
Map 11. Distribution of O • Stylidium claytonioides, and AS.candelabrum.
Map 12. Distribution of O • Stylidium pedunculatum, and AS.perizostera.
Bean, Stylidium subgen. Andersonia
647
Map 13. Distribution of O# Stylidium ericksoniae, and A S.confertum.
Map 14. Distribution of O • S.tenerrimum, and A A S.trichopodum.
Map 15. Distribution of O# Stylidium oviflorum, and A A S.fissilobum.
648
Austrobaileya 5(4): 589-649 (2000)
Map 16. Distribution of + Stylidium longissimum% S.prophyllum,and AS.aquaticum.
Map 17. Distribution of O • Stylidium diffusion A A S.fluminense,and it S.nomination.
115 120 125 130 135 140 145 150 155
Map 18 Distribution of O • Stylidium rotundifolium.
Bean, Stylidium subgen. Andersonia
649
115 120 125 130 135 140 145 150 155
Map 19. Distribution of • Stylidium dunlopianum. 'kS.fimbriatum.
Map 20. Distribution of • S. cor difolium.
Chromosome numbers of some Acanthaceae from Papua New
Guinea
Thomas F. Daniel
Summary
Daniel, Thomas F. (2000). Chromosome numbers of some Acanthaceae from Papua New Guinea.
Austrobaileya 5(4): 651-659. Meiotic chromosome numbers are reported for nine species
representing eight genera of Acanthaceae from Madang and Morobe provinces in Papua New
Guinea. Chromosome numbers of five species are reported for the first time and two new numbers
are reported for the widely cultivated species Graptophyllum pictum (L.) Griff. Chromosome
numbers obtained in Calycacanthus K.Schum. (n = 16) and Jadunia Lindau (n = ca 16) are the first
reported for these genera. Subfamilial relationships are discussed with respect to the chromosome
numbers now known for these and other Acanthaceae.
Keywords: Acanthaceae, chromosomes, Papua New Guinea, Aphelandra, Calycacanthus,
Graptophyllum, Hypoestes, Jadunia, Lepidagathis, Ruellia,Thunbergia
T.F. Daniel, Department of Botany, California Academy of Sciences, Golden Gate Park, San
Francisco, California 94118, U S A
Introduction
The pantropical family Acanthaceae comprise
more than 4000 species in some 230 genera.
Major concentrations of species occur in the
following regions: Mexico-Central America,
Andean South America, Brazil, tropical Africa,
Madagascar, India, southeastern mainland
Asia, and insular Malesia. Two subfamilial
classifications of Acanthaceae are currently in
use. That of Lindau (1895) includes all genera
of the family recognized up to its publication
but is now largely out of date and contains
many errors. Bremekamp’s 1965 revised
classification made some improvements on
Lindau’s, but did not assign all genera to
suprageneric taxa. An updated subfamilial
classification, based on both morphological
and DNA sequence data, is being formulated
(e.g., McDade et al. 2000, Manktelow et al. in
review, McDade et al. submitted).
Like many large and predominantly tropical
families of flowering plants, the Acanthaceae
remain relatively little-studied cytologically. In
1982, Saggoo and Bir reported that
chromosome numbers had been determined for
only about 219 species in the family, and Daniel
and Chuang (1998) noted that only 62 of the
Accepted for publication 5 January 2000
228 genera of Acanthaceae (i.e., 27%)
recognized in Brummitt (1992) have received
any cytological investigation. Whereas
Acanthaceae occurring in India and Mexico-
Central America have received the most
cytological attention, those in Madagascar and
insular Malesia have received little, if any, such
studies. Barker (1986) noted that no
cytological studies had been carried out on
Australian species either.
In 1992,1 had the opportunity to collect
cytological samples of Acanthaceae in Papua
New Guinea. Chromosome number
determinations based on acanthaceous plants
growing in Papuasia had not previously been
made. In the discussions that follow, Papuasia
refers to New Guinea and the Solomon Islands;
New Guinea refers to the nation of Papua New
Guinea (including the archipelagos of New
Britain and New Ireland) and the province of
Irian Jaya of the nation of Indonesia. Hoft (1992)
recognized 129 species in 30 genera (incorrectly
totaled as 32) of Acanthaceae in Papuasia. At
least 23 of these genera are native and four of
them are endemic there. Unfortunately, there is
no comprehensive systematic treatment of the
Acanthaceae of either Papuasia or the Malesian
region. The recognition of 129 species of the
family in Papuasia by Hoft (1992) is likely a
652
conservative estimate; for example, the genus
of at least one species that I observed as
naturalized in Papua New Guinea ( Blechum
pyramidatum (Lam.) Urb.) was not listed by
Hoft. For comparison, about 60 species of
Acanthaceae have been recognized from
Australia (Barker 1986, 1996), about 160
(including many cultivated ornamentals) were
treated as occurring in Java (Backer and
Bakhuizen van den Brink 1965), and 168 were
recognized earlier this century on the Malay
Peninsula (Ridley 1923).
In this study, meiotic chromosome
numbers are reported for nine species of
Acanthaceae occurring in Papua New Guinea
(Table 1). Six of these species ( Calycacanthus
magnusianus K.Schum .,Graptophyllwn
pictum (L.) Griff., Hypoestesfloribunda R.Br.,
Jadunia biroi (Lindau & K.Schum.) Lindau,
Lepidagathis royenii Bremek., and Ruellia
repens L.) are indigenous to the region, three
(C. magnusianus, G. pictum, and J. biroi ) are
presumed to be endemic there, and three
{Aphelandra sinclairiana Nees, Ruellia
tuberosa L., and Thunbergia grandiflora
Roxb.) are naturalized introductions.
Chromosome counts have been reported
previously for at least 20 acanthaceous species
that occur in Papuasia. None of these counts,
with the probable exception of Graptophyllum
pictum (see discussion below), was based on
plants from New Guinea or the Solomon Islands.
They encompass either widespread species that
occur indigenously in Papuasia (e.g., Acanthus
ilicifolius L.) or species that are native
elsewhere but which have become naturalized
in Papuasia (e.g., Ruellia tuberosa).
Materials and Methods
During July and August of 1992, buds, seeds,
and herbarium vouchers of Acanthaceae were
collected in Madang and Morobe provinces of
northeastern Papua New Guinea. Other
Acanthaceae were grown in a greenhouse in
San Francisco from seed collected in Madang
province. Voucher specimens of the latter were
made from the cultivated plants and the letters
“gh” follow the field-collection numbers for
them. Floral buds for chromosomal studies
were fixed in absolute ethanol:glacial acetic acid
Austrobaileya 5(4): 651- 659 (2000)
(3:1) for 24 hours and subsequently washed
and stored in 70% ethanol until processed.
Anthers were macerated in 1% ferric
acetocarmine and subsequently squashed on
a microscope slide. Chromosomes were studied
under oil immersion using a phase contrast
microscope at a magnification of 1 OOOx. Counts
from at least two cells were made for most
collections and all counts were verified by at
least three persons. Camera lucida drawings
were made of preparations from which counts
were obtained. Voucher specimens are
deposited at CAS and LAE. Camera lucida
drawings are attached to the vouchers at CAS.
Representative drawings for each of the
species native to Papua New Guinea are
illustrated. In the following discussions, all
previously published chromosome counts are
listed as n numbers irrespective of whether they
were originally reported as sporophytic or
gametophytic numbers. Voucher specimens, if
they exist, that document previous counts by
other workers have not been examined.
Results and Discussion
Chromosome numbers obtained from these
studies are summarized in Table 1. The
significance of each count is presented in the
following discussions of the genera studied.
AphelandraRBr.
Aphelandra is a neotropical genus of about
175 species. The genus is represented in New
Guinea by A. sinclairiana Nees, a native of
southern CentralAmerica. This showy species
with orange bracts and large, pink corollas is
sometimes cultivated for ornament and has
become naturalized in Madang Province. Our
count of n = 14 for this species agrees with
previously published counts for it based on
plants from neotropical habitats (McDade
1984). This number is also the most widely
known number in Aphelandra and likely
represents the basic number for the genus
(Daniel et al. 1990). Close relatives of
Aphelandra in Lindau’s (1895) Aphelandreae
(i.e., Holographis Nees and Stenandrium
Nees) both appear to have a basic number of =
13 (Daniel et al. 1984, 1990; Piovano and
Bemardello 1991)
Daniel, Acanthaceae from Papua New Guinea
Table 1. Meiotic chromosome numbers of someAcanthaceae from Papua New Guinea.
653
Species
n
Voucher
Aphelandra sinclairiana
14
Madang: Daniel & Forster 6523
Calycacanthus magnusianus
16
Madang: Daniel & Jebb 6518
Graptophyllum pictum
21
Madang: Daniel et al. 6525
G. pictum
20
Madang: Daniel etal. 6530
G. pictum
ca 20
Madang: Daniel 6611
G. pictum
21
Madang: Daniel 6624
Hypoestes floribunda
15
Madang: Daniel et al. 6551
Jadunia biroi
ca 16
Morobe: Daniel et al. 6603
Lepidagathis royenii
21
Madang: Daniel & Forster 6522
L. royenii
ca 21
Madang: Daniel et al. 6538
L. royenii
21
Madang: Daniel et al. 6607
L. royenii
ca 21
Madang: Daniel et al. 6609
Ruellia repens
12
Madang: Daniel 6610gh
Ruellia tuberosa
17
Madang: Daniel 6626gh
Thunbergia grandiflora
28
Madang: Daniel 6627
Several authors have suggested that x = 7 is
primitive for Acanthaceae (Grant 1955; Raven
1975; Piovano and Bemardello 1991; Daniel and
Chuang 1993). If so, x = 14 likely represents a
tetraploid derivative of this primitive basic
number, and * = 13 has evolved via both
polyploidy and dysploidy.
Calycacanthus K.Schum.
Calycacanthus is a unispecific genus endemic
to Papuasia. The count of n = 16 (Fig. 1) for C.
magnusianus is the first report of a
chromosome number in the genus.
Calycacanthus was included in tribe
Odontonemeae subtribe Odontoneminae by
Lindau (1895) and would be included in
Bremekamp’s (1965) tribe Justicieae subtribe
Odontoneminae. Chromosome numbers
reported for other genera of Lindau’s subtribe
comprise n= 11, 12, 14, 22, 23, and 28 (for
Siphonoglossa Oerst., now treated as
congeneric with Justicia L. of subtribe
Justiciinae); n — 15 (for Rhinacanthus Nggs, a
genus best treated in subtribe Justiciinae
according to Daniel and Chuang, 1998; for
information on a dubious and unconfirmable
report of n= 16 in this genus, see Daniel and
Chuang, 1998); n = 18 (for Streblacanthus
Kuntze and Razisea Oerst., the latter best
treated in subtribe Isoglossinae according to
Daniel, 1999); n = 20 (for Ecbolium Kurz); n =
21 (for Odontonema Nees, Oplonia Raf., and
Psender anthemum Radik.); and n = 42 (for
Mackaya Harvey). Thus, n = 16 is newly
reported for the subtribe as delimited by Lindau
(1895).
Graptophyllum Nees.
Graptophyllum comprises between 10 and 15
species occurring primarily in the southwestern
Pacific region. Barker (1986) noted three
species in New Guinea whereas Hoft (1992)
listed five as occurring in Papuasia. The only
previous reports of chromosome numbers in
the genus are for the widely cultivated species
G. pictum (L.) Griff., which has been reported
to have been probably derived from Papuasian
plants (e.g., Bailey 1949, Barker 1986). My
counts from wild population of this species do
not agree with previous reports for it, all of which
appear to have been based on cultivated plants.
Grant (1955) and Govindarajan and Subramanian
(1983, without citation of voucher) reported/? =
30 and Lakshmi and Bapa Rao (1977, without
citation of voucher) reported n = 18 for G
pictum.
654
Austrobaileya 5(4): 651- 659 (2000)
Fig 1. Camera lucida drawings of meiotic chromosome
preparation. Calycacanthus magnusianus ( Daniel &
Jebb 6518), metaphase I, n = 16.
During my studies, chromosome counts
were obtained from several cells at various
stages of meiosis in each of Daniel 6624 ( n =
21 ), Daniel 6525 (n = 21, Fig. 2A), and Daniel
6530 (;n = 20, Fig. 2B). Similar differences in
meiotic chromosome numbers among different
collections of the same taxon are uncommon
but not without precedent in the Acanthaceae.
For example, both n = 11 and n— 12 have been
reported for Elytraria imbricata (Vahl) Pers.
(Daniel et al. 1990) and Siphonoglossa ramosa
Oerst. (Hilsenbeck 1983), and both n- 22 and
n = 23 have been reported for 5. sessilis (Jacq.)
Oerst. (Hilsenbeck 1983). The single cell with
nearly countable chromosomes in Daniel 6611
can only be estimated to be n = ca 20 because
of dark cytoplasmic staining, overlapping of
the irregularly shaped chromosomes, and the
presence of dark granules.
Lindau (1895) incl uded Graptophyllum in
his tribe Graptophylleae along with a diverse
array of other genera, many of which
subsequently have been shown to be more
closely related to genera in other tribes.
Graptophyllum is similar in numerous
morphological features (e.g., presence of
staminodes, pollen type) to Pseuderanthemum
and its relatives (see Daniel 1995). This latter
assemblage also shares a chromosome number
of n = 21. Based on numbers so far reported
for G. pictum , a basic number of x = 10 is
suggested for Graptophyllum ; however,
I
Fig 2. Camera lucida drawings of meiotic
chromosome preparation. A.Graptophyllum pictum
{Daniel et al. 6525), metaphase I, n = 21. B.
G pictum {Daniel et al. 6530), metaphase I, n = 20.
655
Daniel, Acanthaceae from Papua New Guinea
this number is not currently known in the
genus. Determinations of chromosome
numbers for other species of Graptophyllum
will be necessary in order to confirm this or
establish another number as basic in the genus.
Graptophyllum pictum has long been
assumed to be native, and probably endemic,
to New Guinea (see discussion by Barker 1986).
My collections of the species from forest
habitats in Papua New Guinea differ from
cultivated plants (based on collections from
Hawaii, Panama, Papua New Guinea, and the
West Indies at CAS) by lacking variegated
coloring in the leaves, having narrower (less
than 1 mm wide vs. usually more than 1 mm
wide) and more attenuate calyx lobes, and
having fruits present. The cytological and
morphological differences between wild and
cultivated plants suggest either significant
alteration of plants through domestication or
the provenance of cultivated plants from
another region. Indeed, Fosberg et al. (1993)
suggest that G. pictum may be native to the
Moluccas rather than New Guinea.
HypoestesSol. exR.Br.
This genus consists of about 70 species
occurring in the tropics and subtropics of the
Old World. Two species of Hypoestes are
known from New Guinea. The count of n = 15
(Fig. 3) for H. floribunda R.Br., native to
Australia and New Guinea, is the first report of
a chromosome number for this species. It
agrees with most previously reported counts
for other species of Hypoestes (Daniel and
Chuang 1993,1998).
Using the key to varieties of H.
floribunda provided by Barker (198 6), Daniel
et al. 6551 would appear to be affiliated with
var. varia R.M.Barker. This variety was not
reported from New Guinea by Barker (1986).
Hypoestes floribunda var. neoguineensis
R.M.Barker was reported by Barker (1986) to
occur in the same general region of
northeastern Papua New Guinea where Daniel
et al. 6551 was collected. The pubescent
filaments and glandular corolla with a tube 8
mm in length and lobes 11 mm in length readily
distinguishDawz'e/etal. 6551 from that taxon.
Fig 3. Camera lucida drawing of meiotic chromosome preparation. Hypoestes floribunda ( Daniel et al
6551), late telophase II, n = 15.
656
Austrobaileya 5(4): 651- 659 (2000)
The recent report by Daniel and Chuang
(1998) of n = 30 for the African species
H aristata R.Br. suggests a basic number of x
= 15 for the genus. Meiotic complements of n
= 15 are also known in both Dicliptera Juss.
and Peristrophe Nees (Daniel and Chuang
1993, 1998), close relatives of Hypoestes in
Lindau’s (1895) tribe Odontonemeae subtribe
Diclipterinae.
JaduniaLindau.
This genus of two species is endemic to New
Guinea. Chromosome numbers have not been
reported for either of them. Few buds were
available for study and only an approximate
count of n = ca 16 (Fig. 4A) could be obtained
based on a single cell from J. biroi. In the
preparations from this species, the cytoplasm
stained darkly and the chromosomes were not
as clearly defined as illustrated.
Jadunia was treated by Lindau (1895)
in his subtribe Odontoneminae and he noted
affinities with Calycacanthus. A chromosome
count of n = 16 is noted above for the latter
genus.
LepidagathisWilld.
This genus of 100 or fewer species is mostly
paleotropical in distribution. Five species were
noted by Hoft (1992) as occurring in Papuasia.
Previous chromosome counts of n = 9-12, 21,
22, or 42 have been reported for nine species
(Daniel et al. 1990, see under Teliostachya Nees)
of the genus. My counts of n = 21 (Fig. 4B)
and n = ca 21 for L. royenii , a species known
from New Guinea and Queensland, Australia
(see Barker 1986), represent the first reports of
chromosome numbers for this species. A
chromosome number of n = 21 has also been
reported for L. formosensis C. B. Cl ark e exHayata
(Chuang et al. 1963), a species native to Taiwan
and the Ryukyu Islands.
Only approximate counts could be
obtained for two other collections of
L. royenii. Because of folded or possibly
overlapping chromosomes in the single cells of
Daniel et al. 6538 and 6609 with nearly countable
chromosomes, the exact number of chromosomes
(20 vs. 21 or 21 vs. 22 respectively) could not be
resolved. In both instances, however, it is likely
that 21 bivalents were present. Lindau (1895)
V
A
Fig 4. Camera lucida drawings of meiotic
chromosome preparation. A. Jadunia biroi ( Daniel
et al. 6603), diakinesis, n = ca 16. B. Lepidagathis
royenii ( Daniel et al. 6607), metaphase I, n = 21. C.
Ruellia repens ( Daniel 6610gh), metaphase I, n = 12.
Daniel, Acanthaceae from Papua New Guinea
included Lepidagathis in his tribe Barlerieae
and Bremekamp (1965) placed the genus “and
its nearest allies” into his tribe Lepidagathideae.
The only potential relative of Lepidagathis for
which a chromosome number has been
reported is Barleria. Daniel et al. (1990) noted
that n— 12, 15-21 had been reported for that
genus with n = 20 most prevalent. Given the
diversity of chromosome numbers so far
reported for Lepidagathis , the basic number
of the genus is not readily evident.
Morphological variation among
collections of this species was noted by Barker
(1986). Among the collections from which
chromosome counts were determined, Daniel
& Forster 6522 and Daniel et al. 6609 have
bracts and bracteoles 3-5 mm long with the
abaxial surfaces pubescent with two layers of
trichomes (a dense layer of glandular trichomes
and a subtending layer of eglandular
trichomes); Daniel et al. 6607 has bracts and
bracteoles 6-6.5 mm long with the abaxial
surfaces pubescent with three layers of
trichomes (an upper layer of eglandular
trichomes, a middle layer of sparse glandular
trichomes, and a lower layer of eglandular
trichomes); and Daniel et al. 6538 has bracts
and bracteoles 7.5-9 mm long, apically
caudate-awned, and abaxially pubescent with
three layers of trichomes (as in Daniel et al.
6607).
RuelliaL.
In the broad sense in which this genus is often
interpreted, it comprises some 250 species
occurring worldwide. Hoft (1992) noted that
13 species of Ruellia are known from Papuasia.
A meiotic complement of n = 17 is known for
more than 50 species in this morphologically
diverse genus (Daniel and Chuang 1998). My
count of n = 17 for R. tuberosa, native to the
West Indies and northern South America and
naturalized in New Guinea, agrees with the
majority of previous counts for this species
(e.g., Grant 1955; De 1966;VermaandDhillon
1967; Gill 1971; Long 1976; Valsala Deri and
Mathew 1982) and for other species of the
genus (Daniel et al. 1990). However, occasional
reports of n = 16 have been published for R.
tuberosa (e.g., Sugiura 1936, without citation
of voucher; Ellis 1962, without citation of
657
voucher; SubramanianandGovindarajan 1980;
Govindarajan and Subramanian 1983, without
citation of voucher).
The chromosome number of R. repens L.,
native to southeastern mainland Asia and
insular Malesia, is reported here for the first
time as n = 12 (Fig. 4C). This number has not
been reported previously in Ruellia and is the
lowest number known for any species of the
genus. Recently, Daniel and Chuang (1998)
reported n = 24 for the Brazilian species
R. macrantha (Nees) Mart, ex B.D.Jacks, and
the paleotropical species R. prostrata Poir. The
latter species is morphologically similar to
R. repens and both are sometimes treated in
Dipteracanthus Nees (e.g., Bremekamp and
Nannenga-Bremekamp 1948). It is increasingly
apparent that chromosome numbers in Ruellia
are somewhat more diverse than previously
suspected. Based on chromosome numbers
now known for this genus, probable basic
numbers for it includex = 12 andx = 17.
Ruellia was treated by Lindau (1895) in
his tribe Ruellieae and by Bremekamp (1965) in
his subtribe Ruelliinae. Chromosome numbers
reported for other genera included in these taxa
are n = 25 in Lankesteria Lindl. (Mangenot and
Mangenot 1962) and n = 15,17,19,21,22, and
42 (e.g., Grant 1955, Kaur 1970) in Eranthemum
L.
Thunbergia Retz.
Thunbergia comprises about 100 species
native in the Paleotropics. Numerous species
are widely cultivated and some have become
naturalized. At least three, and perhaps five,
species of Thunbergia are known from New
Guinea. All but one (i.e., T. papuana Bremek.,
which according to Barker (1986) might be
synonymous with T. arnhemica F.Muell.)
appear to be naturalized there. My count of n
= 28 for T. grandiflora Roxb. (native to southern
Asia and apparently introduced and naturalized
in New Guinea) agrees with most previous
counts for the species (e.g., Daniel and Chuang
1989, 1998; Grant 1955; Kaur 1970, without
citation of voucher). Older counts of n = 14
(Nanda 1962, without citation of voucher) and
n = ca 14 (Darlington and JanakiAmmal 1945,
without citation of voucher) have also been
reported for the species. If these latter counts
658
are accurate, then the population I sampled in
New Guinea would appear to be tetraploid
within the species. Discussions of chromosome
numbers in Thunbergia were provided by
Daniel and Chuang (1989,1998).
Conclusions
Some of the same chromosomal patterns that
were summarized by Daniel and Chuang (1993,
1998) and Daniel et al. (1984, 1990) were
observed among Acanthaceae occurring in
Papua New Guinea: widely divergent
chromosome numbers within a genus ( Ruellia ),
dysploidy within a species ( Graptophyllum
pictum ), and relatively high (i.e., n— 14 or more)
haploid numbers for most species. Such
chromosomal rearrangements among
Acanthaceae have probably led to some of
the proliferation in numbers of taxa in this large
family.
Four of the genera studied here
(Calycacanthus, Graptophyllum, Hypoestes,
and Jadunia) would be included within
Bremekamp’s (1965) tribe Justicieae subtribe
Odontoneminae. The diversity of chromosome
numbers encountered among them reflects that
reported for other genera of the subtribe from
other geographic regions (Daniel and Chuang
1993). Daniel and Chuang (1993) indicated that
some of these numbers correlate with other
characters and should be useful in recognizing
natural groupings within the Odontoneminae.
The importance of knowledge of chromosome
numbers for discerning systematic
relationships among taxa of Acanthaceae has
been demonstrated previously (Daniel and
Chuang 1993,1998; Daniel etal. 1984,1990).
Further determinations of chromosome
numbers among Papuasian and Australian
Acanthaceae should assist in the elucidation
of their taxonomy and phylogeny.
Acknowledgements
My fieldwork in Papua New Guinea was made
possible by a fellowship from the Christensen
Research Institute. I am most thankful to that
institute and its then director, Matthew Jebb,
for facilitating my studies and for providing a
hospitable research environment. I am grateful
to M. Jebb, P. Forster, and D. Liddle for
Austrobaileya 5(4): 651- 659 (2000)
assistance in the field; to D. Liddle for arranging
for shipment of bud samples; and to
T.I. Chuang, F.M. Chuang, and F. Almeda for
assistance with chromosome preparations and
interpretations.
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Wahlenbergia celata (Campanulaceae), a new species from
central Queensland
Paul I. Forster
Summary
Forster, Paul I. (2000). Wahlenbergia celata (Campanulaceae), a new species from central
Queensland. Austrobaileya 5(4): 661-665. Anew species Wahlenbergia celata P.I.Forst. is described
and illustrated. It belongs to the group of fleshy-rooted, lithophytic Wahlenbergia species that is
endemic in Queensland and the extreme north-east of New South Wales. Four species are now
recognised in this group with new distributional data presented for the previously described species.
An identification key to the species in this group is provided.
Keywords: Campanulaceae, Wahlenbergia- Australia; Wahlenbergia celata , Wahlenbergia glabra ,
Wahlenbergia islensis , Wahlenbergia scopulicola
Paul I. Forster, Queensland Herbarium, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong,
Queensland 4066, Australia
Introduction
A comprehensive revision of the genus
Wahlenbergia in Australia was provided by
Smith (1992) wherein some twenty-six species
were recognised, including several new taxa.
Smith (1992) did not formalise an infrageneric
classification for Wahlenbergia but did
recognise a number of informal groups based
on shared character states.
The first of these groups he called the
“W. scopulicola Group” and defined it by the
shared characters of being “Densely tufted
perennials with much-branched stems and
short inflorescences, typically with only
solitary flowers..."with “deeply campanulate
corollas and hemispherical capsules”. Three
species were included in this group, namely W.
glabra P.J.Sm., W. islensis P.J.Sm. and W.
scopulicola Carolin ex P.J.Sm., all endemic to
southern Queensland, although W. scopulicola
occurs in New South Wales by about 50 m at
Mt Lindesay. An additional shared character
state for these three species is the fleshy,
tuberous tap-root that anchors the plant into
crevices in the rocky substrates inhabited by
all of them. Unlike the remaining, generally wide-
ranging species of Australian Wahlenbergia ,
the species of this group are restricted
endemics in cliffline microsites.
Ongoing fieldwork in Queensland has
revealed a number of additional populations of
Wahlenbergia that can be assigned to Smith’s
Group 1. One undescribed species is present
in this material and is described in this paper.
There are now a number of additional
collections of the previously described species
from new localities not seen by Smith (1992)
and data is also presented on these.
Taxonomy
Materials and methods
This revision is primarily based on collections
held at BRI (as at early 1999) and CANB
(examined prior to 1993). All species have been
examined in the field. Description format is
derived from that of Smith (1992).
l.Wahlenbergia celata P.I.Forst., sp. nov. W.
scopulicola Carolin ex P.J.Sm. afFmis, sed
sepalis lineari-lanceolatis 2.5^1 mm longis
(adversum sepala angusti-triangularia
1.5-2.2 mm longa), corollae
Accepted for publication 5 January 2000
662
lobis multo brevioribus angustioribusque,
2-3 mmx2-2.2 mm (adversum lobos
ellipticos ad ovatos et 5-9 mmx2.5-3.5
mm) et lobis stigmaticis longioribus ab
eadifferens. Typus: Queensland.
Leichhardt District: Bluebell Rock, State
Forest 34, Ruined Castle Creek catchment,
7 Nov 1998, P.I.Forster PIF23930 &
R.Booth & R.Crane (holo: BRI; iso: BRI,
K, MEL).
Perennial herb with a thickened taproot, tufted,
many-stemmed. Stems 5-25 cm long,
decumbent or pendent, crowded, much-
branched, sparsely hirsute; hairs to 0.5 mm
long. Leaves alternate, rarely subopposite,
elliptic, obovate or oblanceolate, becoming
predominantly oblanceolate towards the top
of the stems, 4-30 mm long, 0.8-7 mmm wide,
apex obtuse to acute, base cuneate, petiolate
for up to 2 mm, sparsely hirsute; margins entire
or irregularly toothed, slightly recurved.
Flowers solitary; pedicels 10-50 mm long,
hirsute in lower half, + glabrous in upper half,
without bracteoles. Hypanthium hemispherical,
1.5-2.2 mm long, 1.5-1.8 mm diameter, glabrous.
Sepals 5, erect to slightly spreading,
linear-lanceolate, 2.5^1 mm long, 0.4-1 mm wide
at base, with scattered indumentum. Corolla
deeply campanulate, blue; puberulous inside
at base, otherwise glabrous; tube 4-5 mm long,
3-4 mm diameter, longer than the sepals; lobes
5, triangular, the apex acute, 2-3 mm long, 2-
2.2 mm wide at the base. Stamens 5, filaments
1-1.2 mm long, white; anthers 2-2.5 mm long.
Austrobaileya 5(4): 661-665 (2000)
Ovary 3-locular. Style 4-5 mm long, 3-fid,
indistinctly constricted 1/2 to 2/3 below from
the stigmatic lobes and covered with pollen
presenting hairs above the constriction; 0 or 1
gland below each stigmatic cleft; stigmatic
lobes 1.2-1.5 mm long. Capsule hemispherical,
3-5 mm long, 3-4 mm diameter, glabrous. Seeds
oblong, 0.4-0.5 mm long, c. 0.2 mm diameter,
tan-brown. Fig. 1.
Additional specimens examined: Queensland.
Leichhardt District: [all NW of Taroom] Palmgrove
N.P., Nov 1998, Forster PIF23734 & Booth (BRI,
MEL, NSW); S.F.35 [now S.F.50], upper reaches of
Sandy Creek, Nov 1998, Forster PIF23900 & Booth
(BRI); Ralphs Big Rock, S.F.34 [now S.F.50], Ruined
Castle Creek catchment, Nov 1998, Forster
PIF23934 et al. (AD, BRI, K, MEL, NSW);
Glenhaughton Gorge, Expedition N.P., Sep 1999,
Forster PIF24792 et al. (AD, BRI, MEL);
Glenhaughton Gorge, Expedition N.P., Sep 1999,
Forster PIF24829 et al. (BRI, MEL).
Distribution and habitat: Wahlenbergia
celata has been found to date at five localities
with four in close proximity in the Bigge Range
north-west of Taroom. Plants grow on heavily
weathered sandstone clifflines on large isolated
monoliths or in deeply incised gorges. The
surrounding eucalypt dominated woodland
comprises admixtures of Corymbia citriodora
(Hook.) KD.Hill & L.A.S Johnson,C. watsoniana
(F.Muell.) K.D.Hill & L. A. S. Johnson, Eucalyptus
apothalassica L.A.S.Johnson & K.D.Hill, E.
fibrosaF.Mudl , E. tenuipes (Maiden & Blakely)
Blakely & C.T.White and Lysicarpus
angustifolius (Hook.) Druce. Species in close
Key to the fleshy-rooted, lithophytic species of Wahlenbergia in Queensland and New South
Wales
1. Leaves hairy, petiolate .2
Leaves glabrous, sessile.3
2. Sepals linear-lanceolate, 2.5^1 mm long; corolla lobes triangular with acute tip,
2-3 mm long, 2-2.2 mm wide at base; stigmatic lobes 1.2-1.5 mm long.W. celata
Sepals narrow-triangular, 1.5-2.2 mm long; corolla lobes elliptic to ovate
with acute tip, 5-9 mm long, 2.5-3.5 mm wide at base; stigmatic lobes
0.5-1.3 mm long. W. scopulicola
3. Corolla lobes 5.5-9 mm long; leaves subopposite, narrowly elliptic,
2.5^1 mm wide. W. glabra
Corolla lobes 3.5-5.5 mm long; leaves all alternate, linear, < 1.5 mm wide. W. islensis
Forster, Wahlenbergia celata
association on the rockfaces include
Conospermum sphacelatum Hook.,
Leptospermum sericatum Lindl., Logania
cordifolia Hook, and Mitrasacme oasena
Dunlop. At one locality W. celata is closely
sympatric with W. islensis but seems to be
restricted to moister microsites.
Notes: Wahlenbergia celata is perhaps most
closely allied to W. scopulicola that is endemic
to the Scenic Rim of south-east Queensland
(Forster 1994). The two species are disjunct by
c. 500 km and differ primarily in floral characters.
W. celata has sepals that are linear-lanceolate
and 2.5-4 mm long; corolla lobes that are
triangular with an acute tip, 2-3 mm long and
2-2.2 mm wide at the base; and stigmatic lobes
1.2-1.5 mm long. By comparison W. scopulicola
has sepals that are narrow-triangular and 1.5-
2.2 mm long; corolla lobes that are elliptic to
ovate with an acute tip, 5-9 mm long and 2.5-
3.5 mm wide at the base; and stigmatic lobes
0.5-1.3 mm long. Smith (1992) stated that W.
scopulicola has sepals up to 3 mm long, but I
have not seen this extreme on the available
material. Wahlenbergia celata also has
considerably less hairy foliage than plants of
W. scopulicola from the type locality (Mt
Lindesay) and an additional site at Mt Cougal.
A population of W. scopulicola at Bushrangers
Cave near Numinbah has only sparsely hairy
foliage and was initially segregated as a
separate entity (Henderson 1997), but does not
differ in other characters from the type material.
Hence the density of foliage indumentum does
not appear to be a useful character to differentiate
between W. celata and W. scopulicola.
Etymology: The specific epithet is derived from
the Latin word celatus (concealed) and refers
to the primary habitat of this species in incised
sandstone gorges.
Conservation status: The known localities are
contained within two National Parks and one
State Forest at pristine sites unlikely to be
disturbed in the forseeable future. On present
knowledge the species is restricted in
occurrence but locally abundant. The area is
poorly explored and contains many hectares
of such habitat and it is highly likely that the
663
species will be found to be quite common in
the area. No conservation coding is
recommended.
2. Wahlenbergia glabraP.J.Sm.,Telopea 5:113
(1992). Type: Queensland Moreton
District: near the summit of Mt Cordeaux,
Dec 1973 ,PJ. Smith 55 (holo: NSWn.v.).
Recent (post 1990) specimens examined:
Queensland. Darling Downs District: The Steamers,
E of Emu Vale, May 1990, Bean 1539 (BRI, NSW);
Mt Huntley, western slopes, Oct 1992, Forster
PIF11841 et at. (BRI, MEL); Condamine Gorge, near
Paddys Knob, Mar 1993, Sparshott KS41 & Sparshott
(BRI). Moreton District: Mt Mitchell, Cunningham’s
Gap, Aug 1992, Forster PIF11097 & Reilly (BRI,
MEL, NSW); Mt Cordeaux, Aug 1992, Forster
PIF11181 & Reilly (BRI); Wilson’s Peak, Main Range
N.P., Oct 1992, Halford Q1537 (BRI); Southern
summit of Mt Doubletop, Main Range S of
Cunninghams Gap, Aug 1994, Leiper AQ632132
(BRI).
Notes: W. glabra has been found at a number
of additional localities within the range as
previously described by Smith (1992). It is
endemic to the “Scenic Rim” of south-east
Queensland (Forster 1994). Wahlenbergia
glabra does not co-occur with W. scopulicola
and is found in more westerly parts of the
Scenic Rim on trachyte substrates.
3. Wahlenbergia islensis P.J.Sm., Telopea 5:
114 (1992). Type: Queensland. Leichardt
District: near Isla Gorge, 13 Sep 1974,
P.J. Smith 132 (holo: NSWn.v.).
Recent (post 1990) specimens examined:
Queensland. Leichardt District: Robinson Gorge N.P
[now Expedition N.P], Sep 1992, Forster PIF11290
& Sharpe (AD, BRI, MEL, NSW); Robinson Gorge,
Expedition N.P, Sep 1995, Forster PIF17787 & Figg
(BRI); Bluebell Rock, S.F.34 [now S.F.50], Ruined
Castle Creek catchment, NW of Taroom, Nov 1998,
Forster PIF23927 et al. (AD, BRI, MEL, NE, NSW);
Bat Cave Gorge, Palmgrove N.P., Sep 1999, Forster
PIF24728 & Booth (BRI, MEL); Turpentine Gorge,
Palmgrove N.P., Sep 1999, Forster PIF24770 &
Booth (BRI, MEL).
Notes: W. islensis is now known from several
more easterly localities in the Expedition and
Bigge Ranges than known to Smith (1992). This
species has been found only on sandstone
clifflines. At Bluebell Rock the species is
closely sympatric with W. celata but occurs in
drier microsites.
664
Austrobaileya 5(4): 661-665 (2000)
Fig. 1 . Wahlenbergia celata. A. habit of flowering and fruiting plant. x0.5. B. single leaf showing indumentum. x5.
C. flower.x5. D. expanded corolla.x5. E. fruit.x5. F. style.xlO. A,B,E from Forster PIF23930 et al.(BRI); C,D,F
from Forster PIF23934 et al. (BRI). Del. W. Smith.
Forster, Wahlenbergia celata
4. Wahlenbergia scopulicola Carolin ex P. J. Sm.,
Telopea 5: 111 (1992). Type: Queensland/
New South Wales. Moreton/North
Coast: Mt Lindesay, 31 May 1959, R.C.
Carolin 964 (holo: NSWn.v.).
Wahlenbergia sp. (Numinbah P.I.Forster
+PIF13868) (Henderson 1997).
Recent (post 1990) specimens examined:
Queensland. Moreton District: The Cougals,
Springbrook N.P., Mt Cougal section, Oct 1994,
Forster PIF15854 & Leiper (BRI); Above the
“Bushrangers” cave, lower slopes of Mt Wagawn, Nov
1992, Leiper AQ548270 (BRI); Bushrangers Cave, 1
km W of Numinbah Border gate, Sepl993, Forster
PIF13868 & Leiper (BRI). New South Wales. S.
face of Mt Lindesay, Nov 1990, Halford 369 (BRI,
MEL, NSW).
Notes: W. scopulicola is endemic to the Scenic
Rim of south-east Queensland (Forster 1994).
This species was known to Smith (1992) only
from Mt Lindesay, but has now been found at
two more easterly localities in the Scenic Rim.
Wahlenbergia scopulicola occurs on rhyolite
substrates and is more easterly in its distribution
than W. glabra.
665
Acknowledgements
This paper is dedicated to the late Ralph Crane
who was present when this species was first
discovered.
Thanks to W. Smith (BRI) for the
illustrations and PBostock (BRI) for translation
of the diagnosis into Latin and comments on
the manuscript. Fieldwork in the “Scenic Rim”
of Queensland in 1992 was facilitated with a
National Estate grant to study montane
heathland. Assistance with fieldwork on which
materials pertinent to this paper were collected,
was provided by R.Booth (BRI), F.Carter
(QPWS - Taroom), R.Crane, S.Figg (formerly
BRI), GLeiper, R.Reilly (formerly BRI) and
PR Sharpe.
References
Forster, P.I. (1994). Bill Tullock Memorial Lecture
1994. “Flora of the Scenic Rim”. SGAP Qld.
Region Bulletin 33: 29-36.
Henderson, R.J.F. (1997). Campanulaceae. In
Henderson, R.J.F. (ed ), Queensland Plants
Names and Distribution , pp. 41-42. Brisbane:
Department of Environment.
Smith, P.J. (1992). A revision of the genus
Wahlenbergia (Campanulaceae) in Australia.
Telopea 5: 91-175.
The Identity of Flindersia pimenteliana and F. oppositifolia
(Rutaceae): Evidence from DNA sequences
Kirsten D. Scott, Wayne K. Harris and Julia Play ford.
Summary
Scott, Kirsten D., Harris, Wayne K. & Playford, Julia. (2000). The Identity of Flindersia
pimenteliana and F. oppositifolia (Rutaceae): Evidence from DNA sequences. Austrobaileya 5(4):
667-669. Nucleotide sequencing of two independent genomic regions has shown that two Flindersia
species, F. oppositifolia F.Muell and F. pimenteliana F.Muell were genetically indistinguishable.
This coupled with supporting morphological observations by Hartley (1969) and Whiffin (1982)
has lead us to reassess the taxonomic status of the two species. The new recombination is
Flindersia pimenteliana F.Muell. forma oppositifolia (F.Muell.) K. D. Scott, W. K. Harris &
J.Playford, comb. & stat. nov.
Key words: Rutaceae, Flindersia pimenteliana, Flindersia oppositifolia DNA sequences, systematics,
Australia
Kirsten D.Scott: Centre for Plant Conservation Genetics, Military Rd, PO Box 157, Fismore
2480.
Corresponding author; email: kscott@zen.uq.edu.au
Wayne K.Harris: Queensland Herbarium, Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha
Road,Toowong QFD 4066
Julia Playford: Department of Botany and the Co-operative Research Centre for Tropical Rainforest
Ecology and Management, The University of Queensland, St. Fucia, Queensland 4072
Introduction
The genus Flindersia R.Br. is a predominantly
Australian group with one species in the
Moluccas, one in New Caledonia and four in
New Guinea (Hartley 1969). The genus has 17
species in total, and they are found from
rainforest through to semi-arid habitats. During
a study on the molecular phylogeny of the
genus (Scott et al. , in press) it became apparent
that F. oppositifolia and F. pimenteliana were
anomalous, in that their nucleotide sequences
in two different genomic regions were identical.
This coupled with the observations and
conclusions of Hartley (1969) that the two
species were very closely related andWhiffin’s
(1982) conclusion that F. oppositifolia
(F.Muell.) was a ‘highly derived montane form’
of F. pimenteliana F.Muell. has lead us to
reassess the taxonomic status of the two
species.
Accepted for publication 7 April 2000
Methods
DNA was extracted from 3 individuals of F.
oppositifolia and F. pimenteliana from both
fresh and herbarium material, using the
extraction protocol of Scott and Playford (1996).
PCR of both the chloroplast and the nuclear
DNA fragments was in a 25pi volume
containing: 1.5 mM MgCl 0 ,10 mM KC1,20 mM
Tris-HCl (pH8.7), 10 mM (NH 4 ), S0 4 , 5 pi of
Qmix (Qiagen, Clifton Hill), 0.2 mM each dNTP,
0.25 pM each primer (chloroplast primers e and
f; Taberlet et al. 1991: nuclear ITS-1 primers;
GN1 - Scott and Playford 1996, and C1-5’ TAC
GTT CTT CAT CGA TGC GA 3’ G.Graham
personal communication), 1.25 U Taq
polymerase (Qiagen, Clifton Hill) and 20 ng
DNA. Thermal cycling was in a FTS-1 Thermal
Sequencer (Corbett Research, Mortlake) on a
program of 94°C for 20 s, 55°C for 20 s, and
72°C for 90 s, for 35 cycles. 3 pi of each reaction
were run on a 1% TBE agarose gel to confirm
amplification. The remaining PCR product was
purified by the addition of an equal volume of
668
PEG buffer (30% Polyethylene Glycol 8000,30
mM MgClQ, followed by a 10 min centrifugation
at 15 000 g. The pellet was rinsed with 70%
ethanol, dried and resuspended in 10 pi of sterile
MQ water for sequencing. Sequencing was in
a 20 pi volume containing 8 pi of ABI dye
terminator chemistry (Perkin Elmer, Melbourne),
0.05 pM of primer and 50 ng of PCR product.
The sequencing reaction was in a FTS-1
Thermal sequencer (Corbett Research,
Mortlake) with a program of 96°C for 10 s, 50°C
for 15 s and 60°C for 4 min, for 25 cycles.
Sequences were run on an ABI 373A DNA
sequencer. Sequences were aligned using
Sequence Navigator (Applied Biosystems Inc,
Verl 1994).
Results
Two gene fragments were sequenced in both
directions for the construction of a molecular
phylogeny of Flindersia (Scott et al. , in press).
The two fragments were the ITS-1 spacer
(including 68 bp of 18S rRNA gene and 20 bp
of 5.8S rRNA) which is nuclear, and the
intergenic spacer between trnL-trnE of the
chloroplast. The ITS-1 being nuclear is
biparentally inherited, while the chloroplast
trnL-trnE would be maternally inherited, as
chloroplasts are maternally inherited in most
plants. Both DNA fragments were able to
differentiate every species within the genus,
with the exception of F. pimenteliana and
F. oppositifolia (Scott et al. in press). The ITS-1
fragment in A pimenteliana and F. oppositifolia
was 311 base pairs long. Sequence divergence
for ITS-1 ranged from 1.2-13.4% between
species pairs, with the exception of
F. pimenteliana and F. oppositifolia that were
identical. Similar studies using ITS-1 have
reported sequence divergences of 5-48.9%
between species in the genus Gentiana (Yuan
et al. 1996), and 0.7-21 % between species pairs
in Fraxinus (Jeandroz et al. 1997). Given that
the resolution of the ITS sequence data has in
this case defined all other species within the
genus, in addition to describing intraspecific
variation in five of the 17 species of Flindersia,
it would support the assertion that
F. pimenteliana and F. oppositifolia do not
constitute discrete taxa.
Austrobaileya 5(4): 667- 669(2000)
The second fragment that was
sequenced, was from the chloroplast, and was
372 base pairs long. The trn fragment, like the
ITS fragment was able to distinguish all other
species within Flindersia , with the exception
of differentiating F. pimenteliana from
F. oppositifolia. The chloroplast fragment was
less variable than the nuclear fragment with
0.25-4.2% sequence divergence between
Flindersia species pairs. Sequence divergence
between species pairs of Alnus have ranged
from 0.87-1.52% with divergence values in
Fraxinus ranging from 0.65-1.14% (Gielly and
Taberlet 1994).
The genetic evidence provided from two
independent gene fragments, would suggest
that F. pimenteliana and F. oppositifolia do
not show a level of genetic diversity which
would be indicative of species recognition
within Flindersia.
Systematics
Whilst different in habit and adult leaf form,
both Hartley (1969) and Whiffin (1982) pointed
out the similarities between F. pimenteliana
and F. oppositifolia in the characters of the
fruit, seed and seedlings and regarded both as
being closely similar and Whiffin (op. cit .)
suggested that F. oppositifolia (as unifoliata )
was a ‘highly derived montain form of
F. pimenteliana ’ and his figure four showing
the phylogeny of the genus places the two
species in the one clade. Based on the
observations of these authors and the
molecular evidence presented above, we
believe that the status of F. oppositifolia
should be reassessed and make the following
recombination.
Flindersia pimenteliana F.Muell. forma
oppositifolia (F.Muell.)K. D. Scott,W. K.
Harris & J.Playford, comb. & stat. nov.;
Hypsophila oppositifolia F.Muell., Viet.
Nat. 9:11 (1892). Type: [Queensland, Cook
District]: Mt Bartle Frere, 1892, Johnson
(lecto: MEL fide Hartley & Jessup (1982))
n.v. Flindersia oppositifolia (F.Muell.)
Hartley & Jessup, Brunonia 5:109 (1982).
Flindersia unifoliata Hartley, J. Arnold Arb.
50:498 (1969); Type: [Queensland, Cook
District]: Mt Bellenden Ker, Sayer 136,
(holo: MELw.v.).
669
Scott, Harris, Playford, Flindersia pimenteliana and F. oppositifolia
Acknowledgements
This research project was supported by a grant
from the Co-operative Research Centre for
Tropical Rainforest Ecology and Management.
We would like to thank the Queensland
Herbarium, Australian National Botanic
Gardens, Brisbane Botanic Gardens, and Trevor
Whiffin for providing plant material.
References
Gielly, L., and Taberlet, P. (1994). Chloroplast DNA
polymorphism at the intrageneric level and
plant phylogenies. C.R. Academy of Science
Paris, Sciences de la vie 317: 685-692.
Hartley, T. G. (1969). A revision of the genus
Flindersia (Rutaceae). Journal of the Arnold
Arboretum 50: 481-523.
Jeandroz, S., Roy, A. and Bousquet, J. (1997). Phylogeny
and phylogeography of the circumpolar genus
Fraxinus (Oleaceae) based on Internal
Transcribed Spacer sequences of nuclear
ribosomal DNA. Molecular Phylogenetics and
Evolution 7: 241-251.
Scott, K.D., McIntyre, C. L., and Playford, J. (in press)
Molecular analyses suggest a need for a
significant rearrangement of Rutaceae
subfamilies and a minor reassessment of species
relationships within Flindersia. Plant
Systematics and Evolution
Scott, K., and Playford, J. (1996). A DNA extraction
technique for PCR in rainforest plant species.
Biotechniques 20: 974-978.
Taberlet, P, Gielly, L., Pautou, G, and Bouvet, J. (1991).
Universal primers for amplification of three
non-coding regions of chloroplast DNA. Plant
Molecular Biology 17: 1105-1109.
Whiffin, T. (1982). Variation and evolution in the
genus Flindersia (Rutaceae). 1 review of the
genus. Australian Journal of Botany 30: 635-
643.
Yuan, Y-M., KUpfer, P., and Doyle, J. J. (1996).
Infrageneric phylogeny of the genus Gentiana
(Gentianaceae) inferred from nucleotide
sequences of the internal transcribed spacers
(ITS) of nuclear ribosomal DNA. American
Journal of Botany 83: 641-652.
Table 1: Genbank and herbarium accession numbers for ITS-1 smdtrnL-tmF
Taxon
Herbarium Accessions
ITS-1 Genbank
trnL-trnF Genbank
or sample origin
Accessions
Accessions
Flindersia oppositifolia
BRIAQ 522053
BRIAQ 484238
BRI AQ 459477
AF025500*
AF026021*
Flindersia pimenteliana
BRIAQ 522064
BRIAQ 522058
Australian National
Botanic Gardens
AF025501*
AF026022*
*
Note that there is only a single genbank submission for the herbarium accessions, as the
sequences for each species were identical.
Cereus uruguayanus (Cactaceae) and its naturalised occurrence
in Queensland, Australia
Paul I. Forster & Miriam Schmeider
Summary
Forster, P.I. & Schmeider, M. (2000). Cereus uruguayanus (Cactaceae) and its naturalised
occurrence in Queensland, Australia. Austrobaileya 5(4):671-677. The first records of Cereus
uruguayanus Ritt. ex Kiesl. as a naturalised weed in Australia are reported. Several populations
occur in western Queensland on heavy clay soils in natural and disturbed woodland of brigalow
(Acacia harpophylla F.Muell. ex Benth.) and belah ( Casuarina cristata Miq.). Endozoochorial
dispersal is thought to be responsible for the spread of this species in natural vegetation and
eradication is recommended. It is estimated that at least 3240 individuals occur at one locality
near Glenmorgan. Size class structure of this population is described which shows a preponderance
of seedling juveniles and large mature plants. The stand is also notable for the high proportion of
fasciated (10.9%)and monstrous (20.2%) individuals that occur. This represents the first numerical
data on fasciation and monstrousity in a population, albeit naturalised, of Cactaceae.
Keywords: Cereus uruguayanus, fasciation, monstrousity, naturalised weeds
P.I. Forster & M. Schmeider, Queensland Herbarium, Brisbane Botanic Gardens Mt Coot-tha, Mt
Coot-tha Road,Toowong QLD 4066
Introduction
Species of Cactaceae have been remarkably
successful in colonising the Australian
continent with thirty species currently
considered as being naturalised (Forster 1996).
The majority of naturalised species are from
the genus Opuntia, although taxa from the
genera Acanthocereus , Epiphyllum , Harrisia
(syn. Eriocereus ), Hylocereus, Nyctocereus,
Pereskia and Selenicereus are also present
(Telford 1984; Hosking etal. 1988; Forster 1996).
The majority of these naturalised species
are shrubby to arborescent, spiny succulents
with Opuntia tomentosa Salm-Dyck attaining
a height of 7 m in some situations. To date,
cacti with globular (eg. Echinopsis or
Mammillaria ) or candelabra habits (eg. Cereus
and related genera) have been largely absent
from the Australian naturalised cactus flora.
Naturalisations of species such as Echinopsis
multiplex (Pfeiff) Zucc. have been localised
adventives and easily contained (Mann 1970;
Hosking et al. 1988) and were excluded from
recent listings of naturalised cacti (eg. Telford
1984; Forster 1996). In this paper we document
an extensive naturalisation of Cereus
Accepted for publication 5 January 2000
uruguayanus Ritt. ex Kiesl. encountered during
a visit in February 1997 to the property “Myall
Park” near Glenmorgan. An additional
naturalisation has also been found near Tara
and another reported from the gemfields at
Anakie (J.Higgins pers. comm. 1999).
Cereus uruguayanus is native to
Argentina, Brazil and Uruguay but has been
widely cultivated around the world since the
early 1800’s, usually under the name
C.peruvianus Mill. Kiesling (1982) established
that the name C. peruvianus was misapplied
and renamed the species as C. uruguayanus
Ritt. ex Kiesl. Hunt (1992) has referred without
justification, both the names C. peruvianus
auct. and C. uruguayanus to the synonymy of
C. hildmannianus and Taylor (1998) has
recently recombined C. uruguayanus as a
subspecies of C. hildmannianus. This latter
combination was made in a privately published
journal series that specialises in automatic
tranfers of names with often no justification to
support them. In the case of Taylor’s new
combination there is no explanation offered and
until such time as a comprehensive revision of
the genus is provided it is more appropriate to
follow the nomenclatural lead of Kiesling (1982),
a recognised authority on Argentinian cacti.
672
In his classic work on the cultivation of
cacti, Borg (1937) stated that the species [as
C. peruvianus ] was “Long known in
cultivation”. Despite its ubiquity in cultivation,
there is little ecological or taxonomic
information available about this species. Britton
& Rose (1920) in their monograph of Cactaceae
provide a brief account of both
C. hildmannianus and C. peruvianus, stating
that the former occurs in Brazil and the latter in
south-eastern South America. Benson (1982)
commented that the species (as C. peruvianus)
was commonly naturalised on Kauai in Hawaii
where it was sometimes a pest of pastures.
Most contemporary books on cacti omit
mention of the species (eg. Barthlott 1979;
Andersohn 1983) and accurately identified
illustrations are scarce and generally incomplete
lacking flowers and fruit (Taylor 1968; Hunter
1988a; Glass & Foster 1989; Innes & Glass 1991;
Silva & Sazima 1995). At a locality in south¬
eastern Brazil, C. uruguayanus is stated to
occur on rocky outcrops in both forested and
deforested areas (Silva & Sazima 1995, as
C. peruvianus ). These authors found that this
cactus was predominantly pollinated by
hawkmoths and that seasonal flowering
coincided with an activity peak for these
insects.
Fasciation and monstrosity of the stem
is common in naturalised Australian
populations ofC. uruguayanus , hence we also
report on its numerical occurrence at one
locality. Fasciation in cacti occurs when the
apical meristem divides in an abnormal manner
forming unusual fan-shaped stems (Synder &
Weber 1966; Boke & Ross 1978; Gibson &
Nobel 1986) and is thought to be due to several
factors, such as external stimuli, disease or
heredity (Synder & Weber 1966). Monstrosity
in cacti occurs where each shoot loses its
vegetative point after producing a few areoles
with new growth points produced in an irregular
manner. Such fasciated and monstrose clones
of cacti are often popular as ornamentals. To
the best of our knowledge, there are no detailed
studies of fasciation and monstrosity in natural
populations of cacti, although the occurrence
of isolated individuals in the wild is
occasionally reported as a curiosity (eg. Graham
1962; Lindsay 1962; Foster 1965; Synder &
Austrobaileya 5(4): 671 - 677 (2000)
Weber 1966; Hunter 1984; Sauleda & Sauleda
1984) and often formally named (eg. Borg 1937;
Lindsay 1963; Backeberg 1976). We believe our
study to be the first that details the numerical
occurrence of fasciation and monstrosity, albeit
in a naturalised population.
Materials and Methods
Site Description & History. The study site is
situated on the property “Myall Park” (27°12’S,
149° 39’E), near Glenmorgan some 330 km west
of Brisbane. “Myall Park” is the site of a private
botanic garden “Myall Park Botanic Garden
Ltd.” that is primarily devoted to Western
Australian species and was established in the
1940’s by the redoubtable David Gordon
(McKenzie 1995). Up until the introduction of
the moth Cactoblastis cactorum (Berg) in 1933,
the area around Glenmorgan was severely
infested with 'prickly pear’ ( Opuntia stricta
(Haw.) Haw.) and would appear to be suitable
for the persistence of different sorts of cacti.
Several other species of cacti were believed to
have been cultivated at the “Myall Park”
homestead by the late 1940’s (N.Lester, pers.
comm. 1997) and it is assumed that the
population of C. uruguayanus originates from
this time. C. uruguayanus was semi-
commercially available from nurseries by 1936
(Swinbourne 1982) and by the late 1950’s was
commonly cultivated inAustralia (Fuaux 1957;
Hayes 1958 [all as C. peruvianus]).
At “Myall Park” individuals of
C. uruguayanus (Voucher: Forster 20334 &
Watson: BRI) are concentrated (27° 12' 16 M S,
149° 39’ 35"E) in c. 6 ha of disturbed, but
reasonably intact woodland dominated by
Casuarina cristata Miq. (belah) and Acacia
harpophylla F.Muell. exBenth. (brigalow) on
heavy clay alluvium. Brigalow often occurs as
‘clumps’ as a result of the formation of‘gilgai’,
which are depressions in the soil that hold
water after heavy rain (Johnson, 1980). The
cacti are predominantly concentrated in these
clumps of brigalow. Scattered individuals occur
outside of this area, nearly always in clumps of
natural vegetation, and the furthest individual
observed was about 1 km away from the main
naturalisation (27°12'40"S, 149 o 40'14"E). A
similar distribution of individuals was also
Forster, Schmeider, Cereus uruguayanus in Queensland
673
Fig. 1 . Cereus uruguayanus. A. budding ‘normal’ stem. B. ‘monstrous’ stem. C. ‘cristate’ stem. D. fruit (whole).
E. fruit cross-section. F. seedling. All from Forster 20334 & Watson (BRI). Del. W. Smith.
674
observed in the population nearTara (Voucher:
Forster PIF24959 & Booth, BRI), except that
the brigalow community is more disturbed
being mainly regrowth.
Methodology : Ten quadrats of 10 x 5 m were
placed deliberately within the main
concentration of cacti. All individuals of cacti
in a quadrat were scored for several features of
size class and stem form.
Four size (age) classes of plants were
designated -
0-50 cm (classified as juveniles) (Fig. IE).
50-100 cm (classified as mature as they are
capable of flowering and several showed
evidence of this).
1 -2 m (generally unbranched if with normal stem
morphology).
> 2m (generally branched and with a candelabra
habit).
Three classes of stem organisation were
recognised -
‘normal’ stems (Fig. 1 A) where the 5-7 ribs are
not sinuately indented between areoles
and are more or less straight.
‘monstrous’ stems (Fig. IB) where 7 or more
ribs are present with marked indention
between areoles and the ribs are rarely
straight.
‘cristate’ stems (Fig. 1C) where it is not possible
to accurately ascertain the rib number due
to the form of apical cell division where
many areoles are densely concentrated
and the ribs are never straight.
Spiral stems as illustrated by Hunter (1988a)
were not observed nor were cristate flowers as
described and illustrated by Muller (1988).
Results: Two hundred and sixty-seven
individuals of C. uruguayanus were recorded
from the 10 quadrats with an average of twenty-
seven individuals per plot. Based on this
average it is estimated that the total population
could be in excess of 3240 individuals.
Collectively there was a preponderance of
immature seedlings and large (> 1 m) individuals
(Fig. 2).
Austrobaileya 5(4): 671 - 677 (2000)
‘NormaF individuals make up the bulk of the
total population surveyed (68.9%), followed by
those with cristate stems (20.2%) and
monstrous stems (10.9%) (Fig. 3). ‘Cristate’
individuals are more frequent in the smaller size
classes and for plants over 2 m in height make
up only 9.3% of the population (Fig. 2).
Discussion
Natural History
There are few detailed studies available of size
class structure in cacti and none for natural
populations of Cereus uruguayanus. As
indicated in the materials and methods, the
cacti were noticeably concentrated in natural
vegetation of brigalow clumps. In South Africa,
Taylor & Walker (1984) found that the closely
related C. jamacaru DC. [as C.peruvianus but
see Glen 1997 for correct nomenclature] could
only establish on fine-textured soils with a high
density of shade trees. The requirement of
“prey refugia” and “nurse” plants that create a
suitable microclimate for establishment of
succulent plants is now well known
(Steenbergh & Lowe 1969; Nobel 1988;
McAuliffe 1984). The clumped distribution of
C. uruguayanus at “Myall Park” indicates that
a similar process is occurring, but it is likely to
be mainly due to “nurse” plant availability
rather than “prey refugia”. Most of the
seedlings observed were not hidden in dense
natural vegetation, and predation was
noticeably absent on individuals that were
otherwise readily accessible. Brigalow clumps
may act as a “nurse” plant for C. uruguayanus
by providing microclimatic conditions suitable
for seedling establishment and by acting as
foci for seed dispersal.
This cactus is dependent on cross¬
pollination between different individuals for
fruit to be produced (Silva & Sazima 1995). The
resultant fleshy fruit with numerous seeds (Fig.
ID) appears suited for endozoochorial dispersal
by birds (Bregman 1988). Most of the cacti that
are serious pests in Australia are thought to
have fruit (and hence seed) that are eaten and
dispersed by birds and mammals (Hosking et
al. 1988). If this is the case forC. uruguayanus ,
it would be worth observing birds that utilise
675
Forster, Schmeider, Cereus uruguayanus in Queensland
Size classes
Fig. 2. Total size class distribution for 267 individuals of C. uruguayanus in 10 quadrats at “Myall Park”.
brigalow clumps for roosting to determine those
that feed on these fruit and whether they
disperse seed over any distance.
As yet this naturalisation is relatively localised,
but isolated plants up to a kilometre away would
indicate successful endozoochorial dispersal
is occurring. The brigalow belt, although
widespread in eastern Australia, is now
endangered due to clearing for intensive
agriculture and cattle grazing and has about
2.2% of its original occurrence conserved in
reserves (Young et al. 1999). Given the scale of
the naturalisation of C. jamacaru in South
Africa by 1984 (c. 3000 ha), and the success of
other cacti as agricultural and environmental
weeds in Australia (Hoskingef al. 1988; Forster
1996), particularly in brigalow communities
(McFadyen 1984), it is important that it be
successfully controlled.
The size class distribution of individuals at
“Myall Park” is different to that found by Taylor
& Walker (1984) for C. jamacaru as there is a
greater preponderance of seedlings and large
mature individuals in relation to intermediate
sized plants. Such a ‘bell’ shaped distribution
was implied by Taylor & Walker (1984) to
indicate unstable populations where stand
structure had not yet stabilised and
competition between individuals was not
restricting seedling establishment. At “Myall
Park”, seedlings were generally well scattered,
although in instances where they were closely
situated competition for resources would had
to have been a factor. Once established, growth
of seedlings of C. uruguayanus is rapid and
maturity is reached within 3 or 4 years (pers.
obs. 1978-1997 on cultivated plants at Didcot).
Prior to 1996 the area near Glenmorgan had
experienced over 5 years of periodic drought
and this may have been responsible for a lack
of intermediate sized plants that would have
established in that period. The only way to
determine these sorts of trends would be to
establish permanent plots with tagged
individuals; however, in the current situation it
would be better if the population was
eradicated.
monstrous
cristate 11%
Fig. 3. Total percentage of ‘normal’, ‘cristate’ and
‘monstrous’ individuals of 267 C. uruguayanus
plants in 10 quadrats at “Myall Park”.
676
Fasciation
Fasciation and monstrosity in C. uruguayanus
was first documented by de Candolle in 1800
for cultivated material and various infraspecific
taxa have been named to encompass these
forms (Britton & Rose 1922, Borg 1937; Kiesling
1982). These infraspecific taxa have been
mainly described under C. peruvianus and
none have been transferred to C. uruguayanus
to date and most discussion of the species still
persists under the misapplied name (eg. Hunter
1988a,b; Muller 1988, Machado et al. 2000).
This current study demonstrates that fasciation
and monstrosity may arise continuously within
a population indicating a genetic tendency for
this to occur. It has been speculated that
somatic cross-overs are a possible mechanism
for inducing this variation (Machado et al.
2000). Some ‘normal’ individuals of
C. uruguayanus were also noted as having the
occasional ‘monstrous’ or ‘cristate’ branch,
hence there is no justification for recognition
of such teratological forms as infraspecific taxa
as undertaken by Britton & Rose (1922) or
Backeberg (1976). Rather, if such forms have to
be provided with a name, then selected clones
should be designated as cultivars. Given the
confused history and doubtful typification of
the infraspecific taxa for fasciated individuals
described under C. peruvianus (Kiesling 1982),
it would be wise to arrive at a totally new set of
names for such forms of C. uruguayanus if so
required.
Fasciated individuals of cacti occur rarely in
nature (eg. Graham 1962; Lindsay 1962,1963;
Synder & Weber 1966) or cultivation, but seem
to be very commonly recorded for
C. uruguayanus^ Kiesling 1982; Hunter 1988a;
Muller 1988; Glass & Foster 1989, Machado et
al. 2000). In the case of the monstrous forms of
Lophocereus schottii (Englem.) Britt. & Rose
described by Lindsay (1963), both were
thought to be clones that reproduced
vegetatively. Both Graham (1962) and Lindsay
(1962) stated or inferred that some populations
of cacti tended to have a greater tendency for
fasciation than others, but apart from
mentioning some localities, did not document
numerical occurrence in the wild. This
Austrobaileya 5(4): 671 - 677 (2000)
naturalised population of C. uruguayanus is
probably unusual for the relatively high
percentage of such plants but may be a result
of the founding individual or individuals
carrying genes for this abnormality. Hunter
(1988b) stated that seedlings from fruit of the
‘monstrose’ form of C. uruguayanus will be
nearly 100% true to fonn. In the current example,
juveniles or small adults of both ‘cristate’ and
‘monstrous’ individuals were more common
than large mature individuals over 2 m in height.
Acknowledgements
Thanks are due to W. Smith for the drawings in
Fig. 1, to A.Wood for translation of Kiesling
(1982), to N.Lester and D.Gordon for snippets
of Myall Park history, to C.C.Walker for a copy
of Taylor (1998) and to G.N.Batianoff and
P.D.Bostock for reviewing the manuscript.
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A revision of Eucalyptus normantonensis Maiden & Cambage
(Myrtaceae) and its allies
A.R. Bean
Summary
Bean, A.R. (2000). A revision of the Eucalyptus normantonensis Maiden & Cambage (Myrtaceae)
and its allies. Austrobaileya 5(4): 679-685. Descriptions, distribution maps and an identification
key are provided for the four species comprising the E. normantonensis group, including one new
species (E. provecta) and one new combination (E. tardecidens). All species occur in northern
Queensland, with one extending to the Northern Territory.
Keywords: Eucalyptus, Eucalyptus normantonensis, Eucalyptus provecta , Myrtaceae, Queensland,
taxonomy, new species, key
A.R. Bean, Queensland Herbarium, Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha Road,
Toowong, Queensland, 4066
Introduction
The box-group of eucalypts, i.e. species with
scaly, short-fibred, persistent bark, reniform
cotyledons, petiolate juvenile leaves, adnate
anthers and tenninal, paniculate inflorescences
was distinguished by Blakely (1934) as
Eucalyptus ser. Buxeales Blakely This natural
grouping is still recognised today. Within the
box-group, members of theft, normantonensis
group share the following characteristics:
rough grey bark at least on lower stems; adult
leaves narrow-lanceolate, concolorous, green
to grey-green; juvenile leaves ovate to
lanceolate, grey-green; fruits 3-5.5 mm long,
hemispherical, ovoid or cylindrical, valves
included.
E. normantonensis was named by Maiden
and Cambage in 1919, and was initially known
only from Normanton in north-western
Queensland. Subsequently, the name was
applied to a wide range of trees and mallees
extending from near the east coast of
Queensland to the extreme east of Western
Australia (Hall & Brooker 1974). Johnson &
Hill (1991) recognised the distinctiveness of
some of these populations by naming
E. persistens with two subspecies, from eastern
Queensland.
The closest affinity of E. normantonensis , and
hence of the group as a whole, has never been
Accepted for publication 17 February 2000
clear. Maiden originally considered it to be a
form of E. gracilis F.Muell., a species from
southern Australia, which differs fundamentally
because of its bisected cotyledons and smaller
sub versatile anthers. Blakely (1934) placed ft.
normantonensis close to E. microtheca
F.Muell. and E. rummeryi Maiden. Pryor and
Johnson (1971) placed it next to E. largeana
Blakely and E. lucasii Blakely. Chippendale
(1988) placed it next to E. tectifica F.Muell. and
E. chlorophylla Brooker & Done. A
phylogenetic analysis of the ‘box’-group will
be necessary to determine more precisely, the
relationships between the constituent taxa.
Herbarium material of the
E. normantonensis group is very similar to
E. largiflorens F.Muell., however
E. largiflorens is readily distinguished in the
field by the linear juvenile leaves, completely
rough almost-black bark and alluvial soil
habitat. Furthermore, the distribution of E.
largiflorens does not overlap with any member
of the E. normantonensis group.
Of the species occurring in the same
geographic area, herbarium material of
E. microtheca and E. chlorophylla has been
misidentified as E. normantonensis , and vice-
versa. E. microtheca differs by its smaller and
thinner-walled fruits, while E. chlorophylla
differs by the larger obconical fruits and very
glossy leaves.
680
The individual members of the
E. normantonensis group (E. normantonensis,
E. persistens, E. tardecidens and E. provecta)
often cannot be distinguished in the herbarium
if material is incomplete. Nevertheless the
differences between the taxa are significant and
not merely quantitative (as outlined in the key).
All taxa are allopatric, allowing placement of
incomplete specimens with accompanying
locality data.
Taxonomy
Eucalyptus normantonensis Maiden &
Cambage, J. & Proc. Roy. Soc. New South
Wales 52:489 (1919). Type: Queensland.
Burke District: Normanton, [17°4-‘ S
141° 0-‘ E],August 1913, R.H. Cambage
3930 (holo: NSW; iso: BRI).
E. bicolor var. xanthophylla Blakely, Key
Eucalypts 232 (1934); E. largiflorens var.
xanthophylla (Blakely) Cameron, Victoria
Naturalist 63: 42 (1946). Types:
Camooweal, Qld, 26 June 1922, S.A. White
s.n. (syn: ?, n.v.); Barrow Creek, N.T., 3
May 1922, S.A. White 244 (syn: NSW;
isosyn: AD).
Mallee 2-8 metres high, lignotuberous. Bark
box-type on lower stems, mottled light and dark
grey, closely adhering; smooth, grey to bronze-
coloured above. Juvenile leaves alternate,
petiolate, broadly lanceolate to ovate, up to
8x3 cm, dull (internally glaucous). Adult leaves
narrowly lanceolate, 8-11.5x0.6-1.4 cm,
alternate, leathery, concolorous, dull-yellow to
shiny-green; penninerved, lateral veins at 35-
50° to the midrib; reticulation dense, incomplete,
oil glands intersectional; petioles 0.6-1.0 cm
long. Inflorescences pseudo-terminal,
paniculate, umbellasters 7-flowered; peduncles
thick, more or less terete, 4.5-8 mm long at
anthesis. Mature buds obovoid to ellipsoidal,
3.0-4.0 mm long, 2-2.5 mm in diameter, pedicels
1.5-4 mm long. Hypanthium unribbed;
operculum scar present; outer operculum shed
long before anthesis; inner operculum
hemispherical, with or without umbo, smooth,
thin; stamens white, in about 4 whorls; inner
whorls inflexed, fertile, 0.8-2.0 mm long; outer
whorls irregularly flexed, 2.0-3.0 mm long,
without anthers. Anthers ovoid, adnate,
Austrobaileya 5(4): 679-685 (2000)
basifixed, opening by pores. Style terete, 1.6-
2.0 mm long, stigma blunt. Ovary 3-4-locular,
ovules in 4 longitudinal rows. Fruits
hemispherical, hemispherical or shortly
cylindrical, 3.0-4.0(-4.5) mm long, 3.0-4.0 mm in
diameter, thick-walled, disc annular,
descending, valves obtuse, enclosed. Seeds
ellipsoidal, finely and evenly reticulate, 0.9-1.2
mm long, not toothed, dark brown; chaff
irregular in shape, smaller than seeds, pale
brown.
Selected specimens: Queensland. Burke District:
Normanton, Aug 1936, Blake 12477 (BRI); 27.9 km
S of Croydon turnoff on Cloncurry road, S of
Normanton, Mar 1990, Brooker 10425 (BRI, CANB,
DNA, NSW); 14.9 km from Mt Isa on Camooweal
road. Mar 1990, Brooker 10433 (BRI, CANB, MEL);
40 km W of Mt Isa on Barkly Highway, Aug 1984,
Hill 1041 et al. (BRI, CANB, NSW); 20 miles [32
km] SW of Normanton, Aug 1953, Perry 3944 (BRI,
CANB); 64 miles [102 km] SE of Burketown, Jul
1954, Speck 4774 (BRI) Gregory North District: 7.7
km W of Dajarra, on road to Mt Isa, Jul 1988, Bean
890 (BRI); 122 km W of Winton, W of Cadell Creek,
Mar 1990, Brooker 10435 (BRI, DNA, MEL, NSW);
Bladensburg N.P., S of Winton, Opalton road. Mar
1998, Forster PIF22187 & Booth (AD, BRI, CANB,
DNA, MEL, NSW); on Standish Ranges, 1 km W of
Ibis Bore, 13 km SW of The Monument, Oct 1984,
Neldner 1524 (BRI); 28 km NNW of ‘Pathungra’,
Sep 1977, Purdie 1032 (BRI). Mitchell District: 32
km S of Stonehenge on road to Jundah, Aug 1978,
Dick WQ187 (BRI). Northern Territory. 20 km SW
of Barrow Ck, May 1994, Albrecht 5864 (BRI, CANB,
DNA, NSW, NT); NW of Mt Strezleckii, May 1952,
Bateman 327 (BRI); 16 miles [26 km] E of Coniston
HS„ Feb 1955, Chippendale 1236 (BRI); near Redbank
Gorge, Macdonnell Ranges, Sep 1958, Chippendale
4884 (BRI, DNA); Ormiston Gorge, north ridge, Jun
1972, Dunlop 2627 (BRI, DNA); Central Mt Stuart,
Jun 1969, Maconochie 714 (BRI, DNA); 7 miles [11
km] N of Tennant Creek, Apr 1948, Perry 570 (BRI).
Distribution and habitat: E. normantonensis
is distributed in parts of western Queensland
as far south as Jundah, and in central parts of
the Northern Territory (Map 1). It grows on
ridges (sometimes lateritised) with shallow
gravelly soil.
Phenology: Flowers are recorded mainly from
March to August.
Notes: E. normantonensis has the ability to
produce flowers and fruits while the leaves are
still in an intermediate stage of ontogeny. This
accounts for the small proportion of disparately
broad-leaved specimens in herbaria.
Bean, Eucalyputus norm ant onensis and allies
Buds and fruits of some Northern
Territory material are larger than the
Queensland material, but the difference is not
considered to be sufficient to warrant any
taxonomic recognition. Similarly for difference
in leaf colour; in plants from the type area they
are green and rather glossy, while from inland
areas they are glaucous or yellowish and not
as glossy.
681
throughout, grey, mottled light and dark grey,
closely adhering. Juvenile leaves alternate,
petiolate, lanceolate to broadly lanceolate, up
to 12x3.5 cm, dull (internally glaucous). Adult
leaves lanceolate to narrowly lanceolate, 8-
13x0.8-1.7 cm, alternate, leathery, concolorous,
dull; penninerved, lateral veins at 35-50°to the
midrib; reticulation dense, incomplete, oil
glands intersectional; petioles 1.0-1.7 cm long.
Inflorescences pseudo-terminal, paniculate,
Fig. 1. Eucalyptus provecta. A. mature bud, with operculum scar x 5. B. fruit, showing enclosed valves x 5. C.
longitudinal section of flower, showing inner fertile stamens and outer staminodes x7. D. seed x 20. All from
Bean 12245.
Conservation status: Not considered to be rare
or threatened.
Eucalyptus provecta A R Beansp. nov. afFinis
E. normantonensi a qua habitu arboris,
cortice ubique persistenti, petiolis
longioribus, filamentis staminalibus in
verticillo extimo 4.5-5.5 mm longis differt.
Typus: Queensland. Cook District: 4.8 km
south of ‘Wirra Wirra’, via Forsayth, 18°
38’S 143°43’E, 7August 1997 ,A.R.Bean
12245 (holo: BRI; iso: CANB, MEL, NSW,
QRS).
Tree to 12 metres high, rarely a mallee,
lignotuberous. Bark box-type, persistent
umbellasters 7-flowered; peduncles thick, more
or less terete or somewhat flattened, 4-8 mm
long at anthesis. Mature buds ovoid to
obpyriform, 3.5-5.0 mm long, 2.0-3.0 mm in
diameter, pedicels 3-6 mm long. Hypanthium
unribbed; operculum scar present; outer
operculum shed long before anthesis; inner
operculum hemispherical, with or without
umbo, smooth, thin; stamens white, inner
whorls inflexed, fertile, 1.5-2 mm long, outer
whorls irregularly flexed, 4.5-5.5 mm long,
without anthers. Anthers ovoid, adnate,
basifixed, opening by pores. Style terete, 1.7-2
mm long, stigma tapered. Ovary 3^1-locular,
ovules in 4 longitudinal rows. Fruits ovoid-
682
truncate to obconical, 3.5-5.5 mm long, 3.5-5.0
mm in diameter, thick-walled, disc annular,
valves obtuse, enclosed. Seeds ellipsoidal,
finely reticulate, 0.9-1.5 mm long, not toothed,
dark brown; chaff irregular in shape, smaller
than seeds, pale brown. Fig. 1.
Specimens examined: Queensland. Burke District:
Chudleigh Park station, upper reaches of Stawell
River, May 1995, Forster PIF16503 & Figg (BRI,
CANB, NSW). Cook District: 21.8 km from Forsayth
towards Einasleigh, Jun 1987, Bean 606 (BRI);
Bulleringa N.P., 80 km NW of Mount Surprise, Oct
1992, Bean 5119 (BRI, CANB, MEL); 0.5 km E of
‘Wirra Wirra’ FIS., east of Forsayth, Aug 1997, Bean
12249 (BRI); just S of Beverley Hills HS, SW of
Einasleigh, Aug 1998, Bean 13765 & Fox (BRI,
CANB); crest of Newcastle Range, Jul 1954, Blake
19525 (BRI); Gulf Development road, c. 15 km W of
Mt Surprise, Aug 1979, Clarkson 2534 & Byrnes
(BRI); Mopata H ‘Beverley Hills’, edge of Red plateau,
W of head of Robertson River, Sep 1994, Godwin
EU589C (BRI, CANB, DNA, NSW); c. 20 km S of
Robin Hood station on track to Percy Vale, Aug 1982,
Hill 1062 & Johnson (BRI, CANB, NSW); NW of
Werrington Stn, NW of Hughenden, Oct 1978,
Martens r 1246 (BRI, CANB); 12 miles [19 km] E of
Forsayth township, Jul 1953, Perry 3848 (BRI); 39
miles [62 km] S of Forsayth township, Jul 1953, Perry
3852 (BRI).
Distribution and habitat: E. provecta extends
from Bulleringa National Park (north of Mt
Surprise) to Chudleigh Park station north of
Hughenden (Map 1).
Phenology: Flowers are recorded from May to
August.
Affinities: E. provecta is most closely related
to E. normantonensis, but differs by the tree
habit, bark persistent throughout, petioles 1.0-
1.7 cm long (0.6-1 cm for normantonensis ),
filaments in outer whorl 4.5-5.5 mm long (2-3
mm fox normantonensis) and the mostly larger
fruits. E. provecta differs from E. tardecidens
by the flowers with staminodes, the often
shorter buds, and slightly longer style.
E. provecta differs from E. persistens by the
flowers with staminodes, the style 1.7-2 mm
long (2.5-3 mm long for persistens ) and the
larger fruits.
Conservation status: Not considered to be rare
or threatened.
Austrobaileya 5(4): 679-685 (2000)
Etymology: From the Latin provecta -
advanced, carried forward, extended; in
reference to the rough bark extending
throughout, in contrast to E. normantonensis.
Eucalyptus persistens L A S.Johnson &
K.D.Hill, Telopea 4(2): 336 (1991);
Eucalyptus persistens L. A.S. Johnson &
K.D.Hill subsp .persistens, Telopea 4(2):
336 (1991). Type: Queensland. South
Kennedy District: 4.8 km NE of
Shuttleworth bore, ‘Lou Lou Park’
Station, 22° 15’S 146°09’E, 21 August
1984, K.D. Hill 1182 & LA.S. Johnson
(holo: NSW; iso: BRI, CANB, MEL).
Tree to 12 metres high, or sometimes a mallee,
lignotuberous. Bark box-type, mottled light and
dark grey, closely adhering; persistent
throughout. Juvenile leaves alternate, petiolate,
ovate, up to 10x4 cm, dull. Adult leaves
narrowly lanceolate, 7.5-13x0.9-2 cm, alternate,
leathery, concolorous, somewhat shiny;
penninerved, lateral veins at 35-50° to the
midrib; reticulation dense, incomplete, oil
glands intersectional; petioles 0.8-1.7 cm long.
Inflorescences pseudo-terminal, paniculate,
umbellasters 7-flowered; peduncles thick, more
or less terete, 4-8 mm long at anthesis. Mature
buds ovoid to obovoid, 4-6 mm long, 2-3 mm
in diameter, pedicels (2-)4-6 mm long.
Hypanthium unribbed; operculum scar absent;
inner operculum conical, smooth, thin; stamens
white, inner whorls inflexed, fertile, 1-2 mm
long; outer whorls irregularly flexed, 4-5 mm
long, fertile. Anthers ovoid, adnate, basifixed,
opening by pores. Style terete, 2.5-3 mm long
at anthesis, stigma blunt to tapered. Ovary 3-
4-locular, ovules in 4 longitudinal rows. Fruits
ovoid-truncate to cylindrical, 3—4.5 mm long,
3-3.5 mm in diameter, thick-walled, unribbed;
disc annular; valves obtuse, enclosed. Seeds
ellipsoidal, finely reticulate, 1.2-1.5 mm long,
not toothed, dark brown; chaff irregular in
shape, smaller than seeds, pale brown.
Selected specimens: Queensland. Burke District: W
of “Warang”, White Mountains N.P., Jun 1992, Bean
4624 (BRI, MEL). North Kennedy District: “Upsan
Downs”, 2.5 km S of Greenvale, Apr 1990, Batianoff
900428 (BRI, CANB, NSW); near Bogie River at
“Etonvale”, WSW of Bowen, Mar 1992, Bean 4245
(BRI, NSW); Flinders Highway, 18.4 km W of Charters
Towers, Aug 1997, Bean 12291 (BRI, CANB); Valley
Bean, Eucalyputus normantonensis and allies
of Lagoons, 57 km SW of Mt Garnet, May 1989,
Brooker 10186 (BRI, CANB, DNA, MEL, NSW);
between Wairuna and Minnamoolka, Sep 1996,
Cumming 15124 (BRI); 2 miles S of Middle Ck, Mt
Garnet, Jun 1971, Hyland 5073 (BRI, QRS); Charters
Towers-Clermont road, 45 miles [72 km] from
Charters Towers, May 1960, Johnson 1855 (BRI); 1
mile [1.6 km] S of Niall Station, Jul 1954, Lazarides
4632 (BRI, CANB); 6 miles ESE of Conjuboy Station,
Jul 1953, Perry 3745 (BRI). South Kennedy District:
19 miles [30 km] WNW of “Scartwater” station. May
1964, Adams 977 (BRI, CANB); 1.5 miles [2.4 km]
W of Carmichael Station, Jul 1964, Adams 1175 (BRI,
CANB); “Cairo”, c. 35 miles [56 km] N of Clermont,
Feb 1962, Besset E322 (BRI); NNW of Clermont,
between Miclere and Brown Creeks, Apr 1945, Blake
15688 & Webb (BRI); Mt Coolon-Collinsville road,
0.7 km SW of Caves Ck, Jan 1996, Champion 1302
& Pollock (BRI, CANB, NSW); near the homestead
on Moonoomoo Station, Oct 1983, Henderson
H2788 et al. (BRI); 12.4 km W of Eungella Dam, Aug
1976, Kleinig DK297 (BRI, CANB); 3 miles [5 km] S
of Cape River on Charters Towers-Clermont road,
Jun 1966, Pedley 2128 (BRI); Mitchell District: 137
km S of Torrens Creek on road to Aramac, May 1994,
Brooker 11918 (BRI, CANB); Berricania, Apr 1919,
White s.n. (BRI); S of Mannya N.P., Aug 1994,
Fensham 1796 (BRI).
Distribution and habitat: E. persistens
extends from the White Mountains area to just
north of Clermont, and east to Woodstock and
Collinsville (Map 2). It often grows on the
slopes of lateritic ridges, on pale infertile soils.
It may also grow in deep sands or loams in
undulating terrain.
Phenology: Flowers are recorded mainly from
May to October.
Affinities: It differs from the rest of the group
by the persistent outer operculum.
Conservation status: Not considered to be rare
or threatened.
Eucalyptus tardecidens (L A S. Johnson &
K. D.Hill) A.R.Bean comb, et stat. nov.
Eucalyptus persistens subsp.
tardecidens L.A.S.Johnson & K.D.Hill,
Telopea 4(2): 337 (1991). Type:
Queensland. Cook District: 5.0 km S of
Mt Carbine on Mareeba road, 16°33’ S 145°
09’E, 12 August 1984, K.D. Hill 1066,
L. A.S.Johnson & D.F. Blaxell (holo:
NSW; iso: BRI, CANB, MEL).
Mallee or tree to 7 metres high, lignotuberous.
Bark box-type, mottled light and dark grey,
683
closely adhering, persistent throughout.
Juvenile leaves alternate, petiolate, broadly
lanceolate, c. 8.5x3.0 cm, dull. Adult leaves
lanceolate or narrowly lanceolate, 7.5-15x0.8-
2.1 cm, alternate, leathery, concolorous, green,
dull to shiny; penninerved, lateral veins at 35-
50° to the midrib; reticulation dense, incomplete,
oil glands intersectional; petioles 1.0-1.5 cm
long. Inflorescences pseudo-terminal,
paniculate, umbellasters 7-flowered; peduncles
thick, more or less terete or somewhat flattened,
4-9 mm long at anthesis. Mature buds obovoid
to obpyriform, 4.5-6 mm long, 2.5-3 mm in
diameter, pedicels 3.5-6 mm long. Hypanthium
ribbed; operculum scar present; outer
operculum shed long before anthesis; inner
operculum conical to almost hemispherical,
smooth, thin; stamens white, inner whorls
inflexed, 1-2 mm long, fertile; outer whorls
irregularly flexed, 4.5-5 mm long, fertile. Anthers
ovoid, adnate, basifixed, opening by pores.
Style terete, 2-2.5 mm long at anthesis, stigma
blunt. Ovary 3-4-locular, ovules in 4
longitudinal rows. Fruits ovoid-truncate to
cylindrical, 4.5-5.5 mm long, 4-4.5 mm in
diameter, thick-walled, faintly longitudinally
ribbed; disc annular; valves obtuse, enclosed.
Seeds ellipsoidal, finely reticulate, 0.9-1.2 mm
long, not toothed, dark brown; chaff irregular
in shape, smaller than seeds, pale brown.
Selected specimens: Queensland. Cook District: 9.4
miles [15 km] by road N of Palmer River towards
Cooktown, Aug 1973, Brooker 4023 (BRI, CANB);
25.3 km from Mt Molloy towards Mt Carbine, Nov
1992, Brooker 11330 (BRI, CANB, NSW); 11.7 km
SE of Mt Janet, on survey road along the Dividing
Range, Sep 1984, Clarkson 5508 (BRI, CANB, NSW,
QRS); 7.2 km E of the Peninsula Development road
on a track to the West Normanby River, Sep 1984,
Clarkson 5532 (BRI, CANB, DNA, MEL, NSW); 14.7
km N of Lakeland Downs on track to Bob’s Hut, Oct
1993, Clarkson 10151 & Neldner (BRI, CANB, NSW);
2 km S of Mt Carbine on Mareeba road, Nov 1995,
Forster PIF18126 & Spokes (BRI, QRS); Campbell
Creek on Curraghmore Holding, Nov 1971, Hyland
5694 (BRI, QRS); 16 km S of Lakeland Downs, Nov
1989, Jobson 986 & Lum (BRI, DNA, CANB, MEL);
Mt Carbine, Oct 1976, Knowlton 58 (BRI, QRS); 16
miles WNW of Mt Carbine, Jun 1968, Pedley 2601
(BRI); near Kelly St George River, c. 58 miles [93
km] NW of Mareeba, Oct 1962, Smith 12053 (BRI);
Desailly Range, Jun 1971, Stocker 745 (BRI, QRS).
Distribution and habitat: E. tardecidens has
a restricted distribution in northern
684
Austrobaileya 5(4): 679-685 (2000)
Map 1. Distribution of Eucalyptus provecta A, E. normantonensis •
Map 2. Distribution of Eucalyptus tardecidens A, £. persistens •
Bean, Eucalyputus normantonensis and allies
Queensland, extending from Lakeland Downs
to south of Mt Carbine (Map 2).
Phenology: Flowers are recorded mainly from
October to February.
Affinities: E. tardecidens has the largest fruits
of the group. It differs from E. persistens by its
ribbed hypanthium, shorter style, longer and
broader fruits and presence of an operculum
scar.
685
Conservation status: Although the
geographical extent of this species is small, it
is very common within its area of occurrence.
No conservation coding is recommended.
Key to the species in the Eucalyptus normantonensis group
1. Stamens all fertile. 2
Outer whorl of stamens without anthers.3
2. Outer operculum shed very early; fruits 4.5-5.5x4-4.5 mm. E. tardecidens
Outer operculum retained until anthesis; fruits 3-4.5x3-3.5 mm. E. persistens
3. Mallees; bark rough at base, smooth above; outer filaments 2-3 mm
long; petioles 0.6-1.0 cm long. E. normantonensis
Trees; bark rough throughout; outer filaments 4.5-5.5 mm long;
petioles 1.0-1.7 cm long. E. provecta
Acknowledgements
I am grateful to Will Smith for the illustrations,
to Les Pedley for the Latin diagnosis, and to
the Director of NSW for access to that
Herbarium.
References
Blakely, W.F. (1934). Sydney: A Key to the Eucalypts.
1st edition. Sydney: The Worker Trustees.
Chippendale, G.M. (1988). Myrtaceae - Eucalyptus,
Angophora in Flora of Australia , Volume 19.
Canberra: Australian Government Publishing
Service.
Hall, N. & Brooker, M.I.H. (1974). Forest Tree Series
Leaflet No. 160. Canberra: Australian
Government Publishing Service.
Johnson, L.A.S. & Hill, K.D. (1991). Systematic studies
in the eucalypts - 4. New taxa in Eucalyptus
(Myrtaceae), Telopea 4(2): 321-49.
Pryor, L.D. & Johnson, L.A.S. (1971). A Classification
of the Eucalypts. Canberra: Australian National
University.
Homoranthus coracinus (Myrtaceae), a new species from
Queensland
A.R. Bean
Summary
Bean, A.R. (2000). Homoranthus coracinus (Myrtaceae), a new species from Queensland.
Austrobaileya 5(4): 687-689. A new and rare species of Homoranthus with black petals is described
and illustrated, and compared to related species.
Key words: Homoranthus, Homoranthus coracinus , taxonomy, Queensland flora, Myrtaceae
A.R. Bean, Queensland Herbarium, Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha Road,
Toowong, Qld, 4066
Introduction
Byrnes (1981) named a number of new taxa in
Darwinia Rudge and Homoranthus A.Cunn.
ex Schauer, distinguishing Homoranthus
mainly by the presence of laciniate sepals. The
circumscription of Homoranthus was changed
by Craven and Jones (1991), who placed an
emphasis on placentation and ovule number.
The species described here conforms to their
concept of Homoranthus. Because of the
laciniate sepals, it also fits into Homoranthus
sensu Byrnes.
The genus contains a high proportion of
rare species of restricted distribution, on acidic
substrates with little or no soil development.
Two such species were described by Hunter
(1998) from New South Wales.
Homoranthus coracinus A.R.Bean sp. nov.
afifinis H. darwinioidi autemconflorescentia
3-6 flora, foliis applanatis, petalis atris,
stylo infra stigma pilis tantum aliquot
praedito differt. Typus: Queensland.
Leichhardt District: Mt Mooloolong, Ka
Ka Mundi National Park, via Springsure,
7 May 1999,A.R. Bean 14843 (holo: BRI
(1 sheet + spirit); iso: CANB, MEL, NSW,
distribuendi).
Prostrate to semi-prostrate shrub, to 0.3 m high
and to 1.2 m diameter. Bark grey, fibrous, slightly
furrowed. Leaves narrowly-obovate to
Accepted for publication 11 January 2000
obovate (as seen when viewed perpendicular
to branchlet axis), 3.0-5.5 mm long, c. 0.4 mm
wide, and 1.0-1.6 mm thick, mucronate, green
or grey-green, margins entire; oil glands
scattered, rather obscure. Petioles c. 0.5 mm
long. Conflorescence terminal, anauxotelic, 3-
6-flowered. Bracteoles cymbiform, 6-7 mm long,
2.4-2.7 mm wide, gland-dotted, apex obtuse,
persistent at least until anthesis, enclosing
hypanthia; pedicels (anthopodia) absent;
peduncles thick, 2-3 mm long. Hypanthium
cylindrical, 5.3-6 mm long, 2.8-3.3 mm diameter;
puncticulate, yellow, with 4 indistinct
longitudinal ribs basally; smooth, pink to red
and unribbed distally. Sepals 5, laciniate, 3.2-
4.3 mm long, 1.2-1.5 mm wide, erect, white, with
2-5 acute linear lobes, each 1.0-1.9 mm long.
Petals broadly obovate, 2.7-3.0x2.2-2.7 mm,
black, margin entire. Stamens 10, alternating
with staminodes. Staminodes scarcely adnate
to the adjacent stamen. Filaments c. 0.8 mm
long, terete, tapering. Anthers globose, white,
basifixed, dehiscing by small pores. Style 13-
16 mm long, glabrous except for a few spreading
hairs below the stigmatic area; stigma
unexpanded, papillose. Ovules 7-8, collateral
in two vertical rows. Fruit indehiscent, not
enlarging after anthesis. Fig. 1.
Additional specimen examined: Queensland.
Leichhardt District: Ka Ka Mundi section of Carnarvon
N P., Apr. 1999, Neill, Bouchard & Neill s.n. (BRI).
Distribution and habitat: Known only from
the north-western slopes of Mt Mooloolong
Austrobaileya 5(4): 687-689 (2000)
Fig. 1. Homoranthus coracinus. A. flowering branchxl. B. exterior view of flower at anthesis with one
bracteole removedx4. C. longitudinal section of flowerx8. all from Bean 14843.
in Ka Ka Mundi National Park, SW of
Springsure. It grows on a sloping shelf of
quartzose sandstone with skeletal soil,
supporting only scattered shrubs and stunted
trees. The associated species include
Eucalyptus trachyphloia F.Muell.,
Leptospermum lamellatum Joy Thomps.,
Leucopogon flexifolius R.Br. and Homoranthus
zeteticorum Craven & S.R. Jones.
Phenology: Flowers have been recorded in
April and May.
Affinities: H. coracinus differs from all other
Homoranthus spp. (and perhaps from all other
Bean, Homoranthus coracinus
Myrtaceae) by its petals, which are quite black
in both fresh and dried material. It is most
closely related to H. darwinioides and H.
porteri , both of which also have large
persistent bracteoles and laciniate sepals. H.
coracinus differs from H. darwinioides by its
3-6-flowered conflorescence, flattened leaves,
black petals and style with only a few hairs
below the stigma. H. coracinus differs from H
porteri by the much shorter leaves, the fewer
sepal laciniae, the black petals and the longer
hypanthia.
689
Conservation status: Less than 100 plants are
known from the type locality. It is possible that
more populations will be found, but its habitat
(extensive treeless areas on sandstone slopes)
is not common. Applying the guidelines of the
IUCN (Anon. 1994), a status of‘endangered’ is
recommended (Criterion D).
Etymology: From the Latin ‘ coracinus ’
meaning raven-black, in reference to the petals.
To accommodate H. coracinus , the key given
in Craven & Jones (1991) should be adjusted
as follows:
4. Leaves 3-5.5 mm long
Conflorescence 3-6-flowered; petals black; sepals with 2-5 laciniae.H. coracinus
Conflorescence 2-flowered; petals white; sepals with 6-10 laciniae.H. darwinioides
4. Leaves 4.5-17 mm long (if less than 6 mm long then sepals with fewer than 5 laciniae)
Acknowledgements
I am grateful for the assistance of Bob Campbell,
Queensland Parks and Wildlife Service, to Will
Smith for the illustration and to Les Pedley for
the Latin diagnosis. Thanks also to Greg Neill
and David Bouchard for directions to the site.
References
Anonymous (1994). IUCN Red List Categories.
International Union for Conservation of
Nature and Natural Resources: Switzerland.
Byrnes, N.B. (1981). Notes on the genus Homoranthus
(Myrtaceae) in Australia. Austrobaileya 1(4):
372-5.
Craven, L.A. & Jones, S.R. (1991). ATaxonomic review
of Homoranthus and two new species of
Darwinia (both Myrtaceae, Chamelaucieae).
Australian Systematic Botany 4: 513-33.
Hunter, J.T. (1998). Two new rare species of
Homoranthus (Myrtaceae: Chamelaucieae)
from the Northern Tablelands of New South
Wales. Telopea 8(1): 35-40.
Two new species of Apatophyllum McGillivray (Celastraceae)
from Queensland
A.R. Bean and L.W. Jessup
Summary
Bean A.R. & Jessup L.W. (2000). Two new species of Apatophyllum McGillivray (Celastraceae)
from Queensland. Austrobaileya 5(4): 691-697. Apatophyllum teretifolium and A. flavovirens,
both endemic to Queensland, are newly described and illustrated. The distribution, habitat and
conservation status of each is discussed. Generic distinctions between Apatophyllum and Maytenus
are discussed, including a comparison of the stomatal types possessed by the two genera. A key to
all known species of Apatophyllum is presented.
Keywords: Apatophyllum, Maytenus , Gymnosporia , Apatophyllum teretifolium , Apatophyllum
flavovirens, Celastraceae, key, taxonomy, stomata, new species, Australian flora, Queensland.
A.R. Bean, Queensland Herbarium, Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha road,
Toowong, Queensland 4066, Australia
L.W. Jessup, Queensland Herbarium, Brisbane
Toowong, Queensland 4066, Australia
Introduction
The genus Apatophyllum was described by
McGillivray (1971), comprising two species,
both rare, from easternAustralia. Since then, a
further species has been named from Western
Australia (Cranfield & Lander, 1992).
Apatophyllum is closely related to both
Psammomoya Diels & Loes. and Maytenus
Molina.
Apatophyllum has been adequately
compared and contrasted with Psammomoya
(McGillivray loc. cit.; Cranfield & Lander, loc.
cit.). However, no previous comparisons have
been made between Apatophyllum and
Maytenus Molina, except McGillivray’s report
of an examination of wood structure by C.R.
Metcalfe (loc. cit.) where he states that “a
comparison [of Apatophyllum ] with two
sclerophyllous Australian species of Maytenus
did not reveal any ... significant similarities”.
Apatophyllum has been distinguished
from Maytenus (Jessup 1984) by its solitary
inflorescence. The discovery of a species with
cymose 1-3 flowered inflorescence has
therefore somewhat blurred the generic
boundary between these genera. Maytenus
Accepted for publication 7 April 2000
Botanic Gardens Mt Coot-tha, Mt Coot-tha road.
undoubtedly contains a rather heterogeneous
assemblage of species, and this is evidenced
by the recent reinstatement of Gymnosporia
(Wight &Arn.) Hook.f. by Jordaan & vanWyk
(1999). A review of the whole of Maytenus and
its allies is needed to establish the phylogenetic
relationships of the group, but we feel that the
maintenance of Apatophyllum as a genus is
justified, because of differences in
inflorescence structure, leaf venation and
stomatal type.
C.R. Metcalfe (in McGillivray loc. cit.)
reported paracytic stomata for Apatophyllum
and Psammomoya, which was the first recorded
occurrence of that type of stoma for the family
Subsequently Den Hartog & Baas (1978)
reported paracytic stomata from several
Celastraceae genera, but from only two
Australian species ( Hedraianthera
porphyropetala F.Muell. and Euonymus
globularis Ding Hou). We have examined the
stomata of several species of Apatophyllum
and Maytenus s.L. The results are reported
below.
Apatophyllum differs from Australian
Maytenus by its inflorescences formed strictly
in the axils of true leaves, usually solitary; the
well developed peduncles and the greatly
692
Austrobaileya 5(4): 691 - 697 (2000)
reduced primary inflorescence axis; paracytic
stomata; and the parallel leaf venation and
pungent leaf apices. Maytenus, by contrast,
often has inflorescences borne in the axils of
small bracts or prophylls; the primary axis is
well-developed, giving rise to a pseudo-
racemose inflorescence (or, in the case of
M fasciculiflora Jessup, both peduncle and
primary axis are greatly reduced), usually
several-flowered; stomata laterocytic or
cyclocytic; and the leaf venation reticulate,
even in the narrowest leaves, and leaf apices
never pungent.
To accommodate the two new species
described below, the generic description for
Apatophyllum given in Jessup (1984) needs to
be amended as follows: leaves opposite,
subopposite or alternate, inflorescences 1-3
flowered, cymose, axillary, bisexual; sepals 4
or 5, petals 4 or 5, stamens 4 or 5; ovary +
immersed in disc, 2 or 3-locular.
Jordaan and van Wyk (1999) reinstated
the genus Gymnosporia (Wight & Arn.)
Hook.f. We accept this change and agree that
the names Maytenus emarginata (Willd.)
DingHou, Gymnosporia emarginata (Willd.)
Theu. andG senegalensis(Lam.) Loss, should
not be applied to Malesian and Australian
specimens sensu Hou (1962) and Jessup (1984).
We consider that these specimens (including
Brass 6229 which was cited by Den Hartog &
Baas (1978)), should rightly be called
Gymnosporia inermis Merr. & Perry.
Stomata
Microscope slides of cleared cuticle were
prepared for 4 species of Apatophyllum
(A. teretifolium, A. flavovirens, A. constablei
McGillivray, A. olsenii McGillivray) and 3
species of Maytenus (M silvestris Lander &
L.A.S.Johnson, M. cunninghamii (Hook.)
Loes., M. disperma (F.Muell.) Loes. and
for G. inermis.
Stomatal types were classified using the
terminology given in Dilcher (1974), Den Hartog
& Baas (1978) and Inamdar et al. (1986). All of
the Apatophyllum species examined were
found to possess paracytic stomata,
characterised by the consistent presence of
two subsidiary cells adjacent to the guard cells
and with the same orientation (Fig. 3A-C).
Maytenus silvestris and Gymnosporia inermis
were found to possess laterocytic stomata,
according to the terminology of Den Hartog
& Baas (1978), where three or more subsidiary
cells border onto the lateral sides of the
guard cell pair. In Maytenus disperma and
M cunninghamii the stomata are apparently
cyclocytic, where the subsidiary cells are not
all arranged laterally to the guard cells. In both
the laterocytic and cyclocytic types examined,
the subsidiary cells are partially submerged,
so that they are not plainly visible when the
rest of the epidermis is in sharp focus.
Species
Stomatal type
Voucher
Maytenus silvestris
laterocytic
Halford Q1551 (BRI)
Maytenus cunninghamii
cyclocytic
GrimshawPG473 (BRI)
Maytenus disperma
cyclocytic
Forster 3235 et al. (BRI)
Gymnosporia inermis
laterocytic
Brass 6229 (BRI)
Apatophyllum olsenii
paracytic
Forster 16341 & Thompson (BRI)
Apatophyllum teretifolium
paracytic
Bean 1401 (BRI)
Apatophyllum flavovirens
paracytic
Bean 2225 (BRI)
Apatophyllum constablei
paracytic
Constable & McGillivray 3061
(BRI)
Bean & Jessup, Apatophyllum in Queensland
Taxonomy
Apatophyllum teretifolium A.R.Bean & Jessup
sp. nov. affinis A. constablei autem
stipulis brevioribus, fructibus
longioribus, floribus pro parte maxima 4-
meris, petalis brevioribus differt. Typus:
Queensland. Leichhardt District:
Lonesome National Park, NNE of Injune,
6 May 1999, A.R.Bean 14832 (holo: BRI;
iso: AD, K, MEL, MO, NSW, PERTH,
distribuendi)
Apatophyllum sp. (Expedition Range E.J.
Thompson AQ440723) in Henderson
(1997).
Densely branched rounded shrub 25-40 cm
high, glabrous. Leaves sessile, opposite to sub¬
opposite or occasionally alternate, linear, (6-)
8-12 x 0.3-0.5 mm, elliptical in cross-section;
venation obscure, longitudinal; apex acute to
acuminate, pungent-pointed. Stipules linear,
0.9-1.5 mm long, brown, persistent, mostly
broad-based and with 2 or 3 short lateral lobes;
apex acute to acuminate. Inflorescence axillary,
reduced to a single flower, with 1 or sometimes
2 pairs of empty pherophylls shortly distal to
the base of the axis. Anthopodia 1.7-2.5 mm
long. Pherophylls cymbiform, 0.9-1.5 mm long,
apex acute, margins denticulate. Perianth 4 or
5-merous. Sepals debate, 0.6-0.7 mm long,
persistent on mature capsule; apex acuminate,
margins denticulate. Petals debate, 0.9-1.2 mm
long, 0.7-0.9 mm wide at base, white, somewhat
persistent beyond anthesis; apex acute, margins
entire. Stamens 4 or 5, equal in number to petals,
inserted on margin of disc; filaments tapering,
0.4-0.5 mm long, somewhat persistent beyond
anthesis; anthers basifixed, 0.3-0.4 mm long
and c. 0.4 mm across. Disc fleshy, shallowly
cupular. Ovary ovoid, 2-locular, almost
completely immersed in disc; ovules 2 per
loculus. Style 0.6-0.7 mm long. Capsule
compressed obovoid, 2-valved, 5.2-6 x 2.0-
2.8 mm, surface smooth or minutely papillose.
Seeds ellipsoidal, c. 3.2 mm long, c. 1.4 mm in
diameter; testa dark brown, smooth but with
fine transverse markings; aril white, clasping
base of seed. Fig. 1, 3C.
693
Specimens examined: Queensland. Leichhardt
District: Lonesome N.P., north-east of Injune, Mar
1990, Bean 1401 (BRI); Expedition Range, c. 30 km
WSW of Bauhinia Downs, Mar 1984, Thompson s.n.
(BRI). Burnett District: near Panda Lane, Waaje, NW
corner of Barakula S.F., via Chinchilla, Mar 1998,
Bean 13138 (BRI, CANB, NSW, MEL).
Distribution and habitat: A. teretifolium is
known from 3 localities; the Expedition Range,
Lonesome National Park, and the Barakula State
Forest near Chinchilla (Map 1). It grows in
Eucalyptus dominated woodland to low open
woodland with a heathy understorey, on
shallow sandy soils.
Phenology: Flowers and fruits are recorded for
March and May.
Notes: A large proportion of seeds are evidently
destroyed by caterpillars. Insect predation is
likely to be a limiting factor for regeneration of
this species.
A. teretifolium differs from A.
constablei by the shorter stipules, longer fruits,
the predominantly 4-merous flowers and the
shorter petals.
Conservation status: Three populations of A.
teretifolium have been recorded. Applying the
IUCN guidelines (Anon. 1994), a category of
‘vulnerable’ is proposed (Criterion D2).
Etymology: From the Latin teretifolius ,
meaning ‘withterete leaves’.
Apatophyllum flavovirens A.R.Bean & Jessup
sp. nov. affinis A. olsenii autem foliis
latioribus, stipulis longioribus,
inflorescentiis cymosis, pedunculis
anthopodiis longioribus, ovario 3-
loculari, petalis longioribus differt. Typus:
Queensland. Leichhardt District: Bull
Creek Gorge, 15 km W of ‘Castlevale’, W
of Springsure, 4 September 1990, A.R.
Bean 2225 (holo: BRI; iso: AD, DNA, MEL,
NSW, PERTH).
Apatophyllum sp. (Bull Creek A.R.Bean
2225) in Henderson (1997).
694
Austrobaileya 5(4): 691 - 697 (2000)
Fig. 1. Apatophyllum teretifolium. A. fertile branchlet x 2. B. flower (with one petal removed) and prophylls
20. C. fruit x 10. D. seed x 10. A-C, Bean 14832; D, Bean 1401.
Bean & Jessup, Apatophyllum in Queensland
695
Fig. 2. Apatophyllum flavovirens. A. fertile branchlet x 0.8. B. oblique view of flower x 8. C. fruit x 5. D. seed
x 10. all from Bean 2225.
696
Branched rounded shrub 100 cm high,
glabrous. Petioles 0.5-1 mm long. Leaves
alternate, narrowly lanceolate, 9-21 x 1.6-3.0
mm, flat; venation parallel, longitudinal; apex
acute to acuminate, pungent-pointed. Stipules
linear, 0.3-0.7 mm long, white or yellowish,
persistent, entire to obscurely lobed; apex
acute. Inflorescence axillary, dichasially cymose
with up to 3 flowers or reduced to a single
flower, with 1-several pairs of pherophylls.
Anthopodia 2.0-4.0 mm long at anthesis.
Pherophylls cymbiform, 0.8-1.3 mm long, apex
acute, margins entire. Peduncles 2-3.5 mm long.
Perianth 5-merous. Sepals deltate, 0.6-0.9 mm
long, persistent; apex acute, margins entire.
Petals deltate, 1.7-2.0 mm long, 1.1-1.2 mm wide
at base, yellow, persistent; apex acute, margins
entire. Stamens 5, inserted on margin of disc;
filaments tapering, c. 0.4 mm long; anthers
dorsifixed, c. 0.3 mm long and c. 0.3 mm across.
Disc fleshy, flat. Ovary ovoid, 3-locular, partially
immersed in disc; ovules 2 per loculus. Style
0.5-0.6 mm long. Capsule obovoid, 3-valved,
8-9.5 x 6-7 mm, surface smooth. Seeds
cylindrical, c. 4.0 mm long, c. 1.7 mm in diameter;
testa brown, longitudinally striate; aril white,
clasping base of seed. Fig. 2, 3B.
Additional specimen examined: Queensland.
Leichhardt District: Bull Creek Gorge, S of Springsure-
Tambo road, Sep 1999, Bean 15370 (BRI, K, MO,
NSW).
Austrobaileya 5(4): 691 - 697 (2000)
Distribution and habitat: A. flavovirens is
known only from the type locality (Map 1),
where it grows on a skeletal sandstone slope
(west facing) in open woodland with
Eucalyptus decorticans, Corymbia
hendersonii and Acacia shirleyi. Associated
understorey species include Micromyrtus
leptocalyx, Triodia mitchellii, Grevillea
longistyla and Acacia spp.
Phenology: Flowers and fruits are recorded for
September.
Affinities: A. flavovirens differs from A. olsenii
by the leaves 1.6-3 mm wide (0.7-1.5 mm for A.
olsenii ), stipules longer, 0.3-0.7 mm long (vs.
0.2-0.3 mm long for A. olsenii), cymose
inflorescence, longer peduncles and pedicels,
3-locular ovary (2-locular for A. olsenii ) and
the longer and wider petals.
Conservation status: A. flavovirens is known
from only 19 individuals at the type locality.
No further plants were discovered during a three
day field trip to the area in September 1999.
Applying the IUCN guidelines (Anon. 1994), a
category of‘critically endangered’ is proposed
(Criterion D).
Etymology: From the Latin flavovirens ,
meaning yellowish green, in reference to the
leaf colour.
Key to the species of Apatophyllum
1. Leaves linear, to 0.6 mm wide, terete or almost so .2
Leaves linear-lanceolate to lanceolate, 0.7-3 mm wide, flat.4
2. Perianth 5-merous; petals 1.2-3.5 mm long.3
Perianth predominantly 4-merous; petals 0.9-1.2 mm long. A. teretifolium
3. Petals 1.2-1.5 mm long, margins entire.A. constablei
Petals 2-3.5 mm long, margins fimbriate. A. macgillivrayi
4. Leaves 1.6-3 mm wide; inflorescence cymose; ovary 3-locular. A. flavovirens
Leaves 0.7-1.5 mm wide; inflorescence solitary; ovary 2-locular. A. olsenii
Acknowledgements
We are grateful to Will Smith for the illustrations
and map, to Wayne Harris for assistance in
preparing the stomatal slides and for
photographing them, to Phil Sharpe for a
German translation, to Keith McDonald for
assistance in the field, and to Les Pedley for
the Latin diagnoses.
Bean & Jessup, Apatophyllum in Queensland
697
Fig. 3. Stomatal patterns in Apatophyllum. A. Apatophyllum olsenii x 500. B. Apatophyllum flavovirens x 500.
C. Apatophyllum teretifolium x 300. A, Forster 16341 & Thompson ; B, Retm 2225; C, iterw 1401 (all BRI).
References
Anonymous (1994). IUCN Red List Categories.
International Union for Conservation of
Nature and Natural Resources: Switzerland.
Cranfield, R.J. & Lander, N.S. (1992). Apatophyllum
macgillivrayi (Celastraceae), a new species from
south-west Western Australia. Nuytsia 8(2):
191-4.
Den Hartog, R.M. & Baas, P (1978). Epidermal
Characters of the Celastraceae sensu lato. Acta
Botanica Neerlandica 27: 355-88.
Dilcher, D.L. (1974). Approaches to the Identification
of Angiosperm leaf remains. The Botanical
Review 40: 1-157.
Henderson, R.J.F. (ed.) (1997). Queensland Plants,
Names and Distribution. Brisbane: Department
of Environment.
Inamdar, J.A., Mohan, J.S.S. & Subramanian, R.B.
(1986). Stomatal Classifications - a review.
Feddes Repertorium 97: 147-60.
Jessup, L.W. (1984). Celastraceae, in Flora of Australia,
Volume 22. Canberra: Australian Government
Publishing Service.
Jordaan, M. & Wyk, A.E. van (1999). Systematic
studies in subfamily Celastroideae (Celastraceae)
in southern Africa: reinstatement of the genus
Gymnosporia. South African Journal of Botany
65: 177-81.
McGillivray, D.J. (1971). Apatophyllum'. An interesting
new Australian genus in the family Celastraceae.
Kew Bulletin 25: 401-6.
Hou, D. (1962). Celastraceae I, in Flora Malesiana
ser. 1, 6: 227-91.
Jasminum domatiigerum subsp. australis (01eaceae),a new
subspecies from north Queensland
Wayne K. Harris & William J. McDonald
Summary
Harris, Wayne K. & McDonald, William J (2000). Jasminum domatiigerum subsp. australis
(01eaceae),a new subspecies from north Queensland. Austrobaileya 5(4):699-703. Jasminum
domatiigerum subsp. australis from north Queensland is described as new and is illustrated and
compared with other related species from Australia. Previous records of this species are from New
Guinea where it occurs in similar habitats. This is the first reported occurrence from Australia.
Notes are provided on its distribution, habitat and conservation status. A key to the Australian and
New Guinea trifoliolate species of Jasminum is provided.
Key words: Oleaceae, Jasminum domatiigerum, Jasminum domatiigerum subsp. australis ,
Queensland, New Guinea, systematics, ecology, distribution.
Wayne K.Harris & William J.McDonald: Queensland Herbarium, Mt Coot-tha Botanic Gardens,
Mt Coot-tha Road, Toowong 4066
Introduction
The genus Jasminum L. in Australia consists
of ten species and a number of subspecies and
was most recently revised by Green (1984).The
genus is widely distributed throughout eastern
and northern Australia and two species extend
further westwards. J. calcareum F.Muell.
extends from central Australia through to
Western Australia. J. didymum subsp .lineare
(R.Br.) P.S. Green is the most widespread
extending west from the Great Divide in eastern
Australia through central Australia and into the
northern parts of Western Australia south of
the Kimberley. Jasminum Section Trifoliolata
DC. is represented by two species and three
subspecies. Lingelsheim (1927) described nine
species from Papua New Guinea of which two
belong to section Trifoliolata and one of these
is recorded from Australia. Jasminum
domatiigerum Lingelsh. has a limited
distribution in northern Queensland where it
occurs as a scrambling climber in the wetter
regions in complex notophyll and mesophyll
vine forests.
Jasminum domatiigerum Lingelsh. subsp.
australis W.K. Harris & W. J.McDonald
subsp. nov. Subspecies Jasminii
domatiigeri a typo foliolis petiolulisque
majoribus, pedicellis longioribus et
Accepted for publication 5 January 2000
floribus majoribus (8-15 mm long.)
differens; ad North Queensland restricta
est. Typus: Queensland. Cook District:
Forrester Road Malanda, 17°2LS 145°
34’E; 20 Jul. 1998,7?. Jensen 0 0 8 8 8:
(holo: BRI).
Description: Evergreen scrambling or twining
shrub to about 4 m; stem pubescent to
tomentose with orange-brown simple hairs.
Leaves opposite, trifoliolate; petioles 8-20 mm
long channelled on the upper surface,
pubescent; lamina coriaceous, ovate to broadly
ovate, glabrous to sparsely pubescent on the
adaxial surface, glossy, sparsely pubescent on
the abaxial surface, slightly discolorous,
terminal leaflet (5.5—)6.6—8.0(—8.5) cm long by
(3.6-)4.0-5.0(-5.5) cm broad,petiolule(1.3-)
1.4-1.9(—2.2) cm long, lateral leaflets (2.6-)3.4-
4.0(-4.5) cm long and (1.7—) 1.8—2.6(—3.6) cm
broad, petiolules (2-)3.0-5.0(-6.0) cm long;
margins entire not thickened, slightly recurved;
apex acute to acuminate; base cordate; venation
simple, pinnate, on upper surface depressed,
on lower surface prominently raised with 3 or 4
secondary veins each side of the midrib.
Inflorescences to 90 mm long, on axillary side
shoots, sub-umbellate, (1—) 3 (-7)-flowered,
flowers perfumed, bracts linear subulate,
puberulent 3-6 mm long; pedicels 2-9 mm long,
pubescent. Calyx pubescent, tube 2-7 mm long
700
and 2-2.5 mm in diameter, with 4-5 linear to
triangular teeth, 0.8-2.0 mm long. Corolla
hypocrateriform, white to cream on the outside,
white to cream internally, tube 8-15 mm long
with 4-6 lanceolate or narrow lanceolate acute
lobes, 5-7 mm long. Stamens 2; anthers 3.5^1
mm long, on filaments 1.5-2 mm long, attached
about midway along the corolla tube, not
exerted. Ovary 0.8-1.0 mm long; style about 10
mm long, not exerted, heterostyly not observed;
stigma slightly bilobed about 1.5 mm long. Fruit
spherical to slightly ovoid, often paired (or
single by abortion), 8-10 mm long by 6-8 mm
broad, purple-black. Fig 1,2.
Phenology: Flowering occurs from May
through to July, fruits appearing September to
November. Flowers are sweetly fragrant.
Specimens Examined : Queensland. Cook District:
Daintree R headwaters. Black Mountain area, Daintree
NP, [16°23’S 145°12’E], May 1998, P.I. Forster +
PIF 22968 , (BRI); Herberton Range, SF 194, Mt
Baldy, [17°18’S 145°24’E], Oct 1997, P.I. Forster
PIF 21854 (BRI). North Kennedy District: Arthur
Bailey Road S of Ravenshoe, [17°40’S 143°31’E],
Jun 1995, P.I. Forster PIF 16749 , (BRI, QRS, MEL
,K); Mt Fox, [18°49’S 145°51’E], Oct 1949, F. Mull,
s.n. [AQ 073283] (BRI); South Kennedy District:
Broken River Track, Eungella NP, Nov 1998, W.J.
McDonald + 4484 (BRI); Rd along Clarke Range, SF
679 Teemburra c. 8km S of Crediton township, Jun
1994, W.J. McDonald + 5970 (BRI); Diggings Road
Eungella NP, 20°5-’S 148°3-’E, without date, S.
Pearson SP465 (BRI); Broken River, Eungella NP,
21°1-’S 148°3-’E, May 1985, M.D. Pearson s.n. [AQ
398226] (BRI).
Austrobaileya 5(4): 699-703 (2000)
Distribution and habitat : J. domatiigerum
subsp. australis has been recorded from the northern
regions of Queensland from the Daintree River
to the Eungella National Park in the south (Map
1). The species is associated with complex
mesophyll and notophyll vine forests where it
is a slender scrambling climber. Soil types are
red and usually derived from granite.
Diagnostic Features: J. domatiigerum
subspecies australis is readily distinguished
from the type of the species by its larger flowers
(corolla tub 9-12 mm long), longer pedicels (up
to 9 mm long), larger leaflets (up to 8 mm long)
and longer petiolules (lateral petiolule up to 5
mm long).
Affinities : J. domatiigerum has close affinities
with J. didymum and J. dallachii. It differs from
the former in having densely pubescent to
tomentose branches, petioles and calyces and
hairy domatia. J. dallachii has smaller leaflets,
fewer flowers on a shorter inflorescence and
domatia which are formed at the vein
intersections by a flap of tissue between the
primary and the secondary veins.
Conservation Status: By IUCN (1994) criteria
the taxon is Data Deficient (DD).
Etymology : Latin meaning ‘of the south’ in
reference to its occurrence in Australia.
Key to Jasminum sect. Trifoliolata species in Australia and New Guinea
1. Stems, petioles and leaves pubescent-tomentose.2
Stems, petioles and leaves glabrous or minutely puberulent.5
2. Terminal leaflets mostly less than 6cm long, inflorescences
short, less than 3 cm long mostly 1-5- flowered. J. dallachii
Terminal leaflets greater than 6 cm long, inflorescence greater
than 3 cm long, mostly 5-7- flowered.3
3. Leaves without domatia. J. gilgianum
Leaves with domatia.4
4. Corolla tube 6-8 mm long .
Corolla tube 9-12 mm long
J. domatiigerum subsp. domatiigerum
. . . J. domatiigerum subsp. australis
Harris, McDonald, Jasminum domatiigerum subsp. australis 701
5. Terminal leaflet lanceolate to ovate.6
Terminal leaflet linear to narrow lanceolate. J. didymum subsp. lineare
6. Terminal leaflet more than 4 cm long
Terminal leaflet less than 4cm long .
. J. didymum subsp. didymum
J. didymum subsp. racemosum
Acknowledgements
We thank Dr. G.P. Guymer for provision of
facilities at BRI. Peter Bostock provided the
Latin diagnosis.
References
Green, P.S. (1984). A revision of Jasminum in
Australia. Allertonia 3(6): 403-438.
Lingelsheim, A. (1927). 111. Die Oleaceen Papuasiens.
in Lauterbach L., Beitrage zur Flora von
Papuasien. ser. xiv: 1-22.
Anon (1994). IUCN Red List Categories. International
Union for Conservation and Nature, Gland,
Switzerland. 2 lp.
Map 1 . Distribution of J. domatiigerum subsp.
australis in North Queensland
Fig 1 . Adaxial leaf surface showing prominent domatia
along major veins and at vein intersections. Leaf is c.
3.5 cm broad. PI. Forster PIF 16749 (BRI)
702
Austrobaileya 5(4): 699-703 (2000)
QUEENSLAND HERBARIUM (BRI)
Brisbane Australia
537154
AUSTRALIAN NATIONAL HERBARIUM
ATHERTON (QRS) csmo d™ 0 m a „ t
Jasminum dallachii
NOT
. fragrant.
— • Queensland Herbarium (BRI)
CC^ct ^
^5,
do^ah
SSyt. £XLtS A-a.l/f
nalc(-n
iTKi>
Fig 2. Jasminum domatiigerum subsp. australis W.K. Harris & W.J. McDonald. Photograph of holotype at
BRI. (Approx, half natural size.)
Harris, McDonald, Jasminum domatiigerum subsp.- australis
703
PLANTAE PAPUANAE
Botanical Collections of the Department of Forests
Papua and New Guinea
J.S. Weiners ley .Q.F.9M+-9 April 1957
Jasminum
HERBARIUM OF THE UNIVERSITY OF
NEW ENGLAND (HE)
Jasmin-urn domatiigerum Lingelsh.
Jet. R.Kiew 9.8.81
Mt. Kum, near Mt. Hagen Western Highlands, T.N.G.
Alt. c. 7,000 ft. Lat.5 SO 5. Long. 144 IS E.
OE&^CEril
JASMINUM
Climber over low shrub3 in montane
forest. Flowero: white, scented.
Collected Mt, Kuni, near Mt,Hagen,
T.N.G. 6500'.
J.S. WOMEESliEY. 9.5.57.
ENGLAND
Hj; i; [i A p | UM
272147
Fr is a a nF
BRt LOAN
;
Fig 3. Jasminum domatiigerum subsp. domatiigerum Lingelsh. J.S. Womersly N.G.F. 9449, Mt Kum, PNBG.
(BRI). (Approx, half natural size)
A new species and lectotypification in Campanulaceae:
Lobelioideae
DavidAlbrecht
Summary
Albrecht, D.E. (2000). A new species and lectotypification in Campanulaceae: Lobelioideae.
Austrobaileya 5(4):705-709. Pratia puberula Benth. is lectotypified and Lobelia leucotos Albr.
described and illustrated, with notes on distribution, habitat and relationships with other species of
Lobelioideae.
Keywords: Pratia puberula ; Lobelia leucotos ; Campanulaceae; Lobelioideae; Queensland.
David Albrecht, Northern Territory Herbarium, Parks & Wildlife Commission of the Northern
Territory, PO Box 1046, Alice Springs, N.T., Australia 0871
Introduction
A taxonomic treatment of Australian
Lobelioideae by the author is progressing.
However, it may be some time before
completion due to the unresolved problem of
generic limits and to species complexes
requiring further detailed study. Nevertheless,
some undescribed taxa within this subfamily
present few problems, and it seems sensible to
expedite publication of names for them so that
they are available for use. This paper validates
the name of a new species from Queensland.
A specimen of this taxon, apparently collected
at Cooper’s Creek, South Australia was cited
by Bentham (1868) as a syntype of Pratia
puberula Benth. In order to extricate this
specimen from Pratia puberula sensu stricto
so that the entity that it represents can be
described without confusion, Pratia puberula
is here lectotypified.
Floral measurements are based on fresh, spirit
or rehydrated material.
Typification of Pratia puberula
Pratia puberula Benth., FI. Austral. 4: 133
(1868). Type: Victoria, marshes at Cobra
[Cobberas Mountains],A Mueller (lecto:
K, here selected); probable isolecto:
Cobra, Jan 1854, F. Mueller (MEL [MEL
1592621]).
Accepted for publication 10 March 2000
In the protologue of Pratia puberula , Bentham
(1868) cited the following three collections:
New South Wales, Glendon (Leichhardt);
Victoria, moist, grassy, and marshy places at
Cobra and Mount Barkly (F. Mueller); and
South Australia, Cooper’s Creek (Bowman).
Mueller’s cited collection from Victoria actually
consist of three specimens, two from the
Cobberas Mountains (K and MEL [MEL
1592621 ]) and one from Mount Barkly Ranges
(MEL [MEL 1592622]).
Bentham’s protologue includes a description
of the fruit and seeds of this species, both
critically diagnostic in the Lobelioideae, and
as Mueller’s specimens from the Cobberas
Mountains are the only syntypes with fruit
and seed, albeit rather immature, one of them
has been selected here as lectotype. Bentham
does not appear to have examined Mueller’s
specimen housed at MEL and must have based
his fruit and seed description on the K specimen
which is, therefore, here selected as lectotype.
There have been various applications of the
name Pratia puberula and much confusion
between it and P. pedunculata (R.Br.) Benth.
has resulted. The lectotypification of Pratia
puberula as undertaken here maintains the
prevailing concept of P. puberula in South
Australia (Toelken, 1986) and in Victoria, where
P. puberula is regarded as a synonym of P.
pedunculata (Willis 1973, Albrecht 1999).
Further study of the P. pedunculata - P.
puberula complex is required to determine
706
whether or not more than a single taxon warrants
recognition within it. Clarification of the
typification of P. pedunculata is also required.
Whilst all of Mueller’s specimens cited by
Bentham represent the one entity here defined
as Pratiapuberula, Bentham’s other syntypes
of this name are not of this species. Leichhardt’s
specimen from Glendon (MEL 1592620) is
Pratia purpurascens (R.Br.) E.Wimm., while
Bowman’s specimen is of the species described
here as Lobelia leucotos. There is some
uncertainty about the origin of Bowman’s
collection as the specimen has two labels. The
smaller of the two labels gives the location
“Cooper Creek” and is numbered 228, while the
larger label gives the location “Capes River” in
Mueller’s hand with a further annotation by
J.H. Willis indicating the collector as ‘Bowman’.
It is highly unlikely that Bentham’s citation
“South Australia, Cooper’s Creek” is correct
for this specimen as Lobelia leucotos is not
known to occur in this part of Australia. There
is a Cooper Creek, west-south-west of Mackay,
Queensland, which is closer to, but still slightly
to the south of, the known range ofZ. leucotos.
The most likely location of collection of
Bowman’s specimen is Cape River, Queensland.
This location is to the south of Charters Towers
and within the known range of Lobelia leucotos
(Cumming9589 (BRI) is from near this location).
Bowman collected specimens of other species
(e.g. Eremophila debilis (Andrews) Chinnock
and Rhynchospora pterochaeta F. Muel 1.) from
the Cape River area of Queensland (J. Clarkson
pers. comm.).
Taxonomy
Lobelia leucotos Albr., sp. nov. Lobeliae
stenophyllae Benth. affinis sed floribus
unisexualibus, lobis corollae in latitudine
subequalibus, tubo corollae fisso
profundiore, hypanthio puberulo dense,
alveolis seminibus elongata plerumque
differt. Typus: Queensland. Cook District:
Mareeba on the property of J. Clarkson
on La Spina Road, 17°01’S, 145°24’E, 9
March 1993, J.R. Clarkson 9788 (holo
(functionally male): BRI [AQ 580063]; iso:
DNA, K, L, MBA, MEL; para (functionally
female): ditto, J.R. Clarkson 9787, BRI
Austrobaileya 5(4): 705-710 (2000)
[AQ 580062]; iso para: DNA, K, L, MBA,
MEL, MO, NSW, PERTH).
Herbaceous dioecious perennial. Stems
prostrate or decumbent, puberulous, rarely
almost glabrous, rooting at the nodes. Leaves
distichous, sometimes purple-tinged below,
lanceolate or elliptic to ovate or obovate, 4^14
mm long, 2-15 mm wide, puberulous, with
margins toothed, apex acute to obtuse, and
base cuneate or occasionally obtuse; petiole
ill-defined or rarely to 3mm long. Flowers
solitary, axillary. Bracteoles 0.1-0.6 mm long.
Pedicels 3-35 mm long, puberulous or rarely
almost glabrous. Hypanthium obconic,
ellipsoid or obovoid and 1-2.5 mm long in
functionally male flowers, ovoid to obovoid and
2-4.5 mm long in functionally female flowers,
puberulous externally. Calyx lobes erect,
triangular, 1-2.2 mm long, puberulous, at least
some toothed towards base. Corolla
zygomorphic, 2-lipped, 4.5-8 mm long, generally
slightly smaller on female plants, white
throughout or occasionally with a slight green
or purplish tinge externally, + puberulous
externally; lobes lanceolate-triangular to
narrowly elliptic, glabrous on inner surface,
acute; upper lobes two, 3.5-5 mm long, 0.6-1.2
mm wide, slightly more deeply cut than the
lower lobes, erect; lower lobes three, 2.5-4 mm
long, 0.6-1.3 mm wide, spreading or recurved,
with a green zone at the base; tube 2-4 mm
long, split to within 0.8-1.5 mm of the base,
puberulous internally. Stamens with filaments
2.5-4 mm long, adnate to the corolla tube for
0.5-1.5 mm above the distal edge of the
hypanthium, distally connate for up to c. 0.5
mm, the dorsal three evenly tapered, green
becoming purplish distally, glabrous or
puberulous on inner surface, the ventral two
distally broad, thickened and white tinged
purple (these features less pronounced in
functionally female flowers), tapering rather
abruptly to a narrower green lower part,
puberulous on inner surface. Anther tube 1.4-
2 mm long in functionally male flowers, 0.8-1.4
mm long in functionally female flowers,
glabrous on the exterior surface, two ventral
anthers each with a seta 0.15-0.3 mm long,
dorsal anthers lacking apical setae. Style
glabrous to puberulous; stigma protruding from
the orifice of sterile anther tube in mature
Albrecht, Lobelioideae
functionally female flowers. Capsule ovoid to
ellipsoid, slightly compressed, 4-9.5 mm long,
3-5 mm diameter, puberulous; apical valves
raised 0.5-1.5 mm above base of calyx lobes.
Seeds brown, ellipsoid to broadly ellipsoid,
sometimes asymmetrical, slightly compressed,
0.4-0.6 mm long, 0.3-0.4 mm wide; testa
reticulate, alveolae all elongate or elongate and
+ isodiametric. Fig. 1A-G
Specimens examined: Queensland. Burke District
16.4 km N of Hughenden, Sep 1988, Ingleby QHW
38 (NSW). Cook District: c. 50 m N of Barron Falls
railway station, Jan 1993, Albrecht 5210 & Jobson
(MEL); E Side of Kennedy Hwy, c. 1.5 km S of
Mareeba, Jan 1993, Albrecht 5211 & Jobson (MEL);
Black Rock (Lynd), Apr 1988, Horsup 89 (BRI);
Mareeba, Apr 1962, McKee 9066 (BRI, CANB, NSW);
Goldmine Creek, 19 km from Mareeba towards
Kuranda along Hwy, May 1972, Wrigley & Telford NQ
272 (CBG). North Kennedy District: Cape River, no
date. Bowman [MEL 1592619](MEL); Mt Fox, Dec
1949, Clemens [AQ 417048] (BRI); 22 km N of Burra
microwave tower, towards Poison Valley, W of
Pentland, Apr 1990, Cumming 9589 (BRI);
Herberton, Jan 1936, Flecker 1317 (AD); Hellhole
Creek, near Taravale Homestead, 15km along
Taravale road from Ewan road turnoff. May 1994,
Jobson 2990 & Dixon (MEL); Herberton, Jan 1912,
Kenny [AQ 27373] (BRI); Nigger Creek, Herberton,
no date, Kenny [AQ 27374] (BRI); c. 45 miles SE of
Mt Garnet, Jan 1968, Morain 282 (BRI); Herberton,
Jun 1905, Ringrose [AQ 27380] (BRI); “Lansdown”,
c. 25 miles S of Townsville, Mar 1971, Robertson
T211 (BRI); Townsville, no date, Simmons 283 (BRI).
Distribution and habitat: Lobelia leucotos is
endemic in north Queensland. The species is
known from scattered populations in the Cook
and North Kennedy Pastoral Districts and from
a single collection in the Burke Pastoral District.
The known latitudinal range of the species is
from approximately 16 °S to 21°S. Most
collections have been made in grassy
woodlands on heavy basalt-derived soils, with
fewer collections on sandy or skeletal
substrates. Some populations are likely to
experience regular burning (J. Clarkson pers.
comm.).
Phenology: Flowering specimens have been
collected between January and June. Fruiting
specimens have been collected in March and
April.
Notes: Without supplementary water plants die
back during dry periods and resprout after
707
rains. Plants are known to occur in gardens
and under such conditions will remain leafy
throughout the year (J. Clarkson pers. comm.).
Lobelia leucotos and L. stenophylla Benth.
(s. str., which occurs from southern
Queensland to northern New South Wales)
share two diagnostic character states that are
absent or extremely rare in other Australian
Lobelioideae. In both species, the two ventral
staminal filaments are broad and thickened
distally, and narrow and attenuate proximally
(Fig. 1C). InL. leucotos these features are more
accentuated in functionally male flowers than
in functionally female flowers. The two ventral
anthers in both species also lack penicillate
hairs at the apex, though each have a single
apical seta (Fig. ID). This feature is otherwise
known only in Hypsela tridens E.Wimm.
Lobelia leucotos differs from L. stenophylla in
a number of important floral features set out in
the following synopsis:
L. leucotos'. Flowers functionally unisexual;
corolla lobes white on inner surface, subequal
in width or occasionally the upper pair slightly
narrower; corolla tube split to within 0.8-1.5
mm from its base; hypanthium densely
puberulous externally; seed alveolae
predominantly elongate.
L. stenophylla'. Flowers bisexual; corolla lobes
white, blue or mauvish on inner surface, the
upper pair less than half the width of the lower
3 lobes; corolla tube split to within 2^1 mm
from its base; hypanthium glabrous or rarely
sparsely puberulous externally; seed alveolae
predominantly + isodiametric.
Lobelia leucotos occurs in coastal and inland
locations between the latitudes of c. 16°S to c.
21°S, whilst L. stenophylla occurs
predominantly in near-coastal areas and has a
more southerly distribution between c. 20°S and
c. 32°S.
Specimens of Lobelia leucotos have previously
been determined as Lobelia quadrangularis
R.Br., Pratiapurpurascens (R.Br.) E.Wimm. or
P. puberula Benth. All those three species differ
from Lobelia leucotos in having penicillate hairs
subtending the seta at the apex of the ventral
two anthers, and in having all staminal filaments
708
Austrobaileya 5(4): 705-710 (2000)
Fig. 1. Lobelia leucotos: A. Habit, x 1. B. Functionally male flower, x 8.C. Androecium of functionally male
flower showing anther tube, two ventral staminal filaments and one of three dorsal staminal filaments, x 10. D.
Anther tube orifice of functionally male flower, x 40. E. Fruit, x 6. F. Seed, x 50.G. Seed ornamentation showing
elongate alveolae, x c. 400 A-G - drawn from cultivated material originating from type locality, J. Clarkson s.n.
Albrecht, Lobelioideae
evenly textured and tapering more or less
gradually from apex to base. L. quadrangnlaris
also differs from L. leucotos in having bisexual
flowers with blue to mauve corollas and anther
tubes with short fine hairs on the dorsal exterior
surface. P. purpurascens also differs from L.
leucotos in having glabrous or almost glabrous
hypanthia, the upper corolla lobes markedly
narrower than the lower lobes, and larger seeds
>0.7 mm rather than< 0.6 mm long. P. puberula
also differs from L. leucotos in having more
deeply split corolla tubes split to <0.7 mm rather
than 0.8—1.5 mm from the base and indehiscent
rather than dehiscent fruit.
Specimens of L. leucotos show considerable
variation in leaf size, shape and prominence of
marginal toothing. Corolla size and plant
indumentum, on the other hand, exhibit low
variability within the species. Very few ripe fruit
of this species have been preserved in
herbarium specimens. In those studied, the
narrow summit of the fruit expands to 0.5-1.5
mm above the distal edge of the hypanthium
and separates into two valves when fully
mature. The calyx lobes persist and remain
erect, often obscuring the summit of the fruit
(Fig. IE). This cryptic dehiscence is similar to
that found in L. darlingensis (E.Wimm.)Albr.
(Albrecht, 1994).
Conservation status: This taxon is not
considered rare or threatened.
Etymology: The specific epithet is derived from
Greek leucon-, white-, and Otion, auricle or little
ear, in reference to the upper corolla lobes in
this species, which bear some resemblance to
erect rabbit ears.
709
Acknowledgements
I am most grateful to John Clarkson for
collecting live material of Lobelia leucotos and
for providing information on habitat and
collecting localities; to Peter Jobson for his
generous help with fieldwork; to the nursery
staff of the Royal Botanic Gardens Melbourne,
and Alice Springs Desert Park for their attentive
care of cultivated plants; to Neville Walsh for
the Latin diagnosis; to Philip Short and Don
Foreman for checking types at K and BM for
me; to Clyde Dunlop for comments on the
manuscript; to the Directors of BRI, NSW, SYD
and MEL for loans and access to their
collections; and to Sally Mumford for preparing
the illustration.
References
Albrecht, D.E. (1994). A new combination in Lobelia
L. (Campanulaceae: Lobelioideae). Telopea 5:
791-792.
Albrecht, D.E. (1999). Campanulaceae: Lobelioideae.
In N.G. Walsh & T.J. Entwisle (eds). Flora of
Victoria 4: 563-578. Port Melbourne: Inkata
Press.
Bentham, G. (1868). (Pratia in Campanulaceae, Flora
Australiensis: a Description of the Plants of the
Australian Territory ) 4: 133. London: L. Reeve
& Co.
Toelken, H.R. (1986). Campanulaceae - Pratia. In
J.P Jessop & H.R.Toelken (eds). Flora of South
Australia III: 1374-1376. Adelaide: South
Australian Government Printing Division.
Willis, J.H. (1973). A Handbook to Plants in Victoria
2: 630. Carlton: Melbourne University Press.
Chamaesyce ophiolitica (Euphorbiaceae), a new and endangered
species endemic to serpentine vegetation in central Queensland
Paul I. Forster
Summary
Forster, Paul .1. Chamaesyce ophiolitica P.I.Forst. (Euphorbiaceae), a new and endangered species
endemic to serpentine vegetation in central Queensland. Austrobaileya 5(4):711—714 (2000).
The new species Chamaesyce ophiolitica is described and illustrated. Information is provided on
its distribution, habitat and conservation status. Its affinities are with the C. drummondii complex.
C. ophiolitica is restricted to serpentine soils north of Rockhampton in central Queensland. The
species is known from three extant localities and fulfils the criteria to be listed as an Endangered
species.
Keywords: Chamaesyce ophiolitica, Euphorbiaceae, serpentine.
Paul I. Forster, Queensland Herbarium, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong,
Queensland 4066, Australia.
Introduction
Revisions of the Australian species of
Chamaesyce S.F.Gray and Euphorbia L. were
undertaken by Hassall (1977a), although only
the latter was formally published (Hassall
1977b). New combinations in Chamaesyce were
published by Hassall (1976) and for some
additional Queensland species by Forster &
Henderson (1995) to enable use of the generic
name in the most recent census of Queensland
plants (Forster & Henderson 1997).
Unfortunately no overall revision of the
Australian species of Chamaesyce has been
published.
The species that is the subject of this
short paper was first collected in 1920 by Bill
Francis at the Warren State Farm north of
Rockhampton and his specimen was annotated
by Hassall as being of affinity to Chamaesyce
petala (Ewart & L.R.Kerr) P.I.Forst. &
R.J.F.Hend., a species that occurs in northern
Australia in the Northern Territory. Apart from
a collection in 1960, this plant was only really
brought to attention with a number of
collections in the late 1980’s that coincided with
an increase in attention to the floristics and
vegetation communities that occur on
serpentine soils and rocks in central
Queensland (Batianoff & Specht 1992;
Batianoff etal. 1990,1991,1997,2000).
At least eighteen species of plants are
endemic to the serpentine vegetation of central
Queensland with a number of these being of
significance for conservation and listed as rare
or threatened taxa (Batianoff et al. 2000). The
species described here has been rarely collected
and is directly threatened by changes to the
serpentine landscape due to mining and
agriculture. It is formally named in this paper to
expedite efforts to ensure its conservation and
to draw attention to an otherwise obscure
existence.
Taxonomy
Chamaesyce ophiolitica P.I.Forst., sp. nov., a
Chamaesyce petala (Ewart & L.R.Kerr)
P.I.Forst. & R.J.F.Hend. folii lamina
cordato-elliptica usque obovata
(adversum laminam oblongam), cyathii
bracteis truncatis usque oblongo-
truncatis (adversum bracteas
spathulatas), stipulis infirme evolutis et
sparse ramosis (adversum stipulas bene
evolutas et valde fimbriatas) et seminibus
multo majoribus (1.7-1.8 mm longis x 1-
1.1 mm latis x 1 - 1.1 mm crassis contra
circa 1 x 0.7 x 0.7 mm) differt. Typus:
Queensland. Port Curtis District: west
of Canoona, 1 March 1994, P.I.Forster
PIF15042 ScA.RBean (holo: BRI; iso: AD,
DNA, MEL).
Accepted for publication 31 May 2000
712
Herbaceous annual, monoecious, of prostrate
habit and up to 5 cm high and 20 cm across,
with white latex. Stems branching divaricately,
lower stems up to 2 mm diameter, upper leaf¬
bearing stems up to 1 mm diameter, glabrous or
with sparse to dense erect trichomes to 0.3 mm
long; interpetiolar stipules deeply divaricate to
subulate, 0.4-0.8 mm long, glabrous. Leaves
discolorous; petioles 0.8-1 mm long, 0.3-0.4
mm wide, channelled on top, glabrous or with
sparse trichomes; lamina cordate-elliptic to
obovate, often unequal at base, 2-12 mm long,
1.3-11 mm wide, glabrous, or with scattered to
sparse trichomes on upper surface and sparse
to dense trichomes on the lower surface,
glaucous blue-green above, paler below;
margins entire or very weakly serrulate; apex
acute to rounded; base unequal, cordate to
lobate. Cyathia solitary or occasionally paired;
peduncles 0.2-1.5 mm long, 0.5-0.7 mm
diameter, glabrous or with scattered to dense
trichomes. Cyathium 2-2.5 mm diameter; bracts
4 or 5, truncate to oblong-truncate, irregularly
fimbriate, 0.4-0.8 mm long, 0.9-1 mm wide,
white; glands 4 or 5, elliptic to oblong-reniform,
0.4-0.5 mm long, 0.7-1 mm wide, green;
trichomes sparse to dense and up to 0.5 mm
long. Male flowers: filament 1.5-1.7 mm long,
c. 0.3 mm wide, flattened, anther reniform, c. 0.4
mm long and 0.3 mm wide. Female flower: ovary
trilobed, up to 1 mm long and 1.2 mm diameter,
with dense trichomes; styles 3,0.5-1 mm long,
bilobed for 0.2-0.5 mm with the tips recurved,
with sparse trichomes. Fruit trilobed, c. 3 mm
long and 3 mm diameter, with dense trichomes.
Seeds obconical-trigonous, 1.7-1.8 mm long,
1-1.1 mm wide and 1-1.1 mm thick, fissure 1.5-
1.6 mm long, pale brown. Fig. 1.
Other specimens examined. Queensland. Port Curtis
District South Percy Island, 50 km NE of Arthur
Point, Shoalwater Bay, Oct 1989, Batianoff 11422 et
al. (BRI); W of Canoona, Jan 1988, Forster PIF3393
(BRI); Warren State Farm, Mar 1920, Francis
AQ202944 (BRI); On Rockhampton - Marlborough
road. May 1960, Johnson 1720 (BRI); Mt Wheeler,
Rockhampton, Jan 1989, Specht 3 & Reeves (BRI).
Notes: Chamaesyce ophiolitica is compared
to C. petala which appears to be the most
similar species on morphological characters.
C. ophiolitica differs from C. petala in the
leaves being cordate-elliptic to obovate (versus
oblong), the cyathial bracts more truncate to
Austrobaileya 5(4): 711 - 714 (2000)
oblong-truncate (versus spathulate), the
poorly developed stipules are little-branched
(versus well developed stipules that are
fimbriate) and much larger seeds (1.7-1.8x1-
1.1 x l-l.l mm versus c. 1 x 0.7 x 0.7 mm).
A comparison may also be made to the
widespread C. drummondii (Boiss.)
D.C.Hassall which differs in the oblong leaves,
well developed stipules (0.8-2 mm long), poorly
developed stipules with an uneven margin and
smaller seeds (c. 1 x 0.7 x 0.7 mm).
The whole species complex which
includes Chamaesyce drummondii is in need
of a critical biosystematic revision. This was
undertaken in part by Hassall (1977a) but is
now in need of updating. The distribution of
C. ophiolitica is allopatric to that of C. petala
and it is perhaps more likely that the new
species is derived from the widespread
C. drummondii.
There is some interesting variation in
the limited material to hand of C. ophiolitica.
Within the same population it is possible to
obtain individuals that are either glabrous or
with dense coverage of trichomes on the foliage
and some floral parts. These individuals appear
to grow in close proximity to one another and
further study is required to ascertain whether
this is a simple case of a single character switch
or is correlated with more fundamental
differences. Serpentine soils present a relatively
severe environment for plants and it has been
demonstrated elsewhere that morphologically
distinct races of the same species can coexist
and maintain their distinctive nature over time
(Rajakaruna & Bohm 1999).
Distribution: Chamaesyce ophiolitica has
been collected from four localities in central
Queensland north of Rockhampton. However,
only three of these collections occurred within
the last 40 years.
Habitat: This new species is restricted to upper
slopes and sides of low ridges on soils derived
from serpentine rocks and occurs in stony
situations in open woodland dominated by tree
species such as Eucalyptus fibrosa subsp.
(Glen Geddes M.I.Brooker 10230) and
Corymbia xanthope (A.R.Bean & Brooker)
K.D.Hill & L.A.S.Johnson . These serpentine
Forster, Chamaesyce ophiolitica
713
Fig. 1. Chamaesyce ophiolitica. A. habit of flowering shoot, x 2. B. undersurface of glabrous leaf. *4. C.
undersurface of pubescent leaf. x8. D. cyathium from above.x 16. E. cyathium and fruit from side. x8. F. female
flower from side. xl6. G. cyathium with cyathial bracts and glands removed showing male flowers. xi6. H. face
view of fruit. x8. I. side view of seed. xl6. J. ventral view of seed. xl6. All from Forster 15042 (BRI). Del. W.
Smith.
714
Austrobaileya 5(4): 711 - 714 (2000)
landscapes cover c. 1000 km 2 in central
Queensland (Batianoff et al. 2000).
Conservation Status: Chamaesyce ophiolitica
is currently known from only three localities,
one (South Percy Island) that is National Park.
Using the IUCN Red List categories this
species can be classified as Endangered on the
criteria - B. Extent of occurrence estimated to
be less than 5000 km 2 or area of occupancy
estimated to be less than 500 km 2 , and estimates
indicating any two of the following:
1. Severely fragmented or known to exist at
no more than five locations.
2. Continuing decline, inferred, observed or
projected, in any of the following: (a) extent
of occurrence, (b) area of occupancy, (c)
area, extent and/or quality of habitat, (d)
number of locations or subpopulations.
C. Population estimated to number less than
2500 mature individuals and either: 1. An
estimated continuing decline of at least 20%
within 5 years or 2 generations, whichever is
longer.
C. ophiolitica is a very insignificant
annual herb and it is likely that further
populations may be found. As yet, the effects
of fire and other disturbances on its ecology
are unknown. It should be noted that the
vegetation on serpentine soils has now been
intensively studied (work of Batianoff and
collaborators) and that this species has been
shown to be sparsely distributed in the area. A
relevant comparison may be made with the
recently described Bursaria reevesii L.Cayzer
& M.D.Crisp, a small shrub that is only known
from a handful of localities on the serpentine
(Cayzeretal. 1999).
Etymology: The specific epithet refers to the
occurrence of this species on soils derived from
serpentine.
Acknowledgements
Thanks to Will Smith for the illustrations, to
George Batianoff for the latest information on
the serpentine vegetation of central
Queensland and to Peter Bostock for
translation of the diagnosis into Latin and for
comments on the manuscript.
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Stackhousia tryonii Bailey. A nickle-
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central Queensland. Australian Journal of
Botany 38: 121-130.
Batianoff, G.N., Specht, R.L. & Reeves, R.D. (1991).
The serpentinite flora of the humid subtropics
of eastern Australia. Proceedings of the Royal
Society of Queensland 101: 137-157.
Batianoff, G.N., Reeves, R.D. & Specht, R.L. (1997).
The effects of serpentine on vegetation
structure, species diversity and endemism in
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Ecology of Ultramafic and Metalliferous Areas.
“Proceedings of the Second International
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New Caledonia, July 31-August 5, 1995.
Noumea: Center ORSTOM, BP A5, 98848.
Batianoff, G.N., Neldner, V.J. & Singh, S. (2000).
Vascular plant census and floristic analysis of
serpentine landscapes in central Queensland.
Proceedings of the Royal Society of Queensland
109:1-30.
Cayzer, L.W., Crisp, M.D. & Telford, I.R.H. (1999).
Bursaria (Pittosporaceae): a morphometric
analysis and revision. Australian Systematic
Botany 12: 117-143.
Forster, P.I. & Henderson, R.J.F. (1995). New
combinations in Chamaesyce (Euphorbiaceae)
from Queensland, Australia. Novon 5: 323-324.
Forster, P.I. & Henderson, R.J.F. (1997).
Euphorbiaceae. In R.J.F. Henderson (ed.)
Queensland Plants Names & Distribution , pp.
69-76. Brisbane: Department of Environment.
Hassall, D.C. (1976). Numerical and cytotaxonomic
evidence for generic delimitation in Australian
Euphorbieae. Australian Journal of Botany 24:
633-640.
Hassall, D C. (1977a). Systematic studies in Australian
Euphorbieae (Euphorbiaceae). Ph D thesis,
University of Queensland.
Hassall, D.C. (1977b). The genus Euphorbia in
Australia. Australian Journal of Botany 25:
429-453.
Rajakaruna, N. & Bohm, B.A. (1999). The edaphic
factor and patterns of variation in Lasthenia
californica (Asteraceae). American Journal of
Botany 86: 1576-1596.
Clausena smyrelliana (Rutaceae: Aurantioideae), a new and
critically endangered species from south-east Queensland
Paul I. Forster
Summary
Forster, Paul.I. (2000). Clausena smyrelliana (Rutaceae: Aurantioideae), a new and critically
endangered species from south-east Queensland. Austrobaileya 5(4): 715-720. Clausena
smyrelliana is described and illustrated. Information is provided on its distribution, habitat,
phenology and conservation status. The species is currently known from one extant individual
in the wild and a conservation status of Endangered is recommended.
Keywords: Clausena, Clausena brevistyla, Clausena smyrelliana, Rutaceae
P I. Forster, Queensland Herbarium, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong,
Queensland 4066, Australia
Introduction
In 1992 Greg Smyrell, then of Maryborough,
brought to my attention a single individual of a
small Rutaceous tree that he had encountered
in a remnant patch of littoral microphyll
vineforest near Dundowran north of Hervey
Bay in south-east Queensland. Subsequent
collections of flowers and fruit led to the
conclusion that this individual could be
classified in the genus Clausena and the
material was allocated a code name, segregated
in the holdings at BRI in November 1995 and
listed in the census of plants for Queensland
(Forster 1997). All of this material was sent on
loan to NSW in 1996 for examination by
D.J.Mabberley who has prepared an account
of this group of Rutaceae for the Flora of
Australia. On return of this loaned material in
1997, some of the specimens were redetermined
as Micromelum minutum (G.Forst.) Wight &
Arn. and one as Anacardiaceae indeterminate.
Admittedly the Clausena species does
resemble Euroschinus falcata Hook.f.
superficially, and examination of the BRI
holdings of this species did reveal a further
collection of Clausena. However, if a more
than cursory examination of material is made,
then the Rutaceous nature of the material
(obvious oil glands in the foliage) is evident.
A monograph of the genus Clausena
Burm.f. has been recently published wherein
some fifteen species were recognised with
C. brevistyla Oliver native to Australia (Molino
Accepted for publication 7 April 2000
1994). Only C. brevistyla var. brevistyla was
considered to occur in Queensland, with both
C. brevistyla var. brevistyla and C. brevistyla
\M.papuana (Lauterb.) J.F.Molino considered
to occur in New Guinea and parts of Indonesia.
The Queensland distribution of C. brevistyla
var. brevistyla is from the tip of Cape York south
to the Goodnight Scrub National Park west of
Booyal. The other species of Clausena grow
in Africa, mainland Asia and other parts of
Malesia. Clausena brevistyla was included in
Clausena section Clausena together with
C. excavata Burm.f. from Malesia and Asia,
C. kanpurensis J.F.Molino from India,
C. harmandiana (Pierre) Pierre ex Guillaumin
from south-east Asia, C. lansium (Lour.) Skeels
from southern China and C. poilanei
J.F.Molino from Vietnam.
Several species of Clausena have
economic significance. Clausena lansium
(“Wampi”) from southern China is commonly
cultivated for the large edible faiit. Some species
such as C. anisata (Willd.) Hook.f. ex Benth.
from Africa and C. heptaphylla (Roxb.) Wight
& Arn. ex Steudel from India have aromatic
foliage rich in essential oils (Lockwood 1984;
Okunade & Olaifa 1987). The foliage and roots
of C. anisata are also used for a range of folk
remedies (Coates Palgrave 1977; Beentje 1994).
Other species such as C. sanki (Perrottet)
J.F.Molino are also occasionally cultivated (as
C. anisum-olens : Merrill 1912; Madulid 1995).
In Queensland there are records of cultivated
plants of C. lansium , but otherwise no
additional species have been recorded.
716
Examination of the essential oils of the
leaves of the Clausena species from
Dundowran revealed similarities to the oils
found in Clausena brevistyla as found in
Queensland, and total dissimilarity to that
found in Micromelum minutum (J.J.Brophy
unpubl.). The leaf essential oils of both
Clausena brevistyla and the species from
Dundowran are predominately monoterpenoid
in content with less than 10% of sequiterpenes,
whereas that of Micromelum minutum is
predominately sequiterpenoid in content with
less than 10% of monoterpenes. The Clausena
species from Dundowran (Voucher: P.I.Forster
PIF17928) has a leaf essential oil mainly
composed of alpha-pinenene (73%), beta-
caryophyllene (9%) and caryophyllene oxide
(4%). Clausena brevistyla (Vouchers:
PI.Forster PIF25985, PIF21850, PIF25006
respectively) has a leaf essential oil composed
mainly of alpha-pinene (67-73%), beta-pinene
(5-13%), beta-caryophyllene (4-7%),
bicyclogermacrene (2-3%), or 3-carene (7-31 %)
and limonene (56-84%), or purely alpha-pinene
(91%).
Seedlings of the Clausena species from
Dundowran are reputedly dissimilar to those
of Clausena brevistyla (G.Smyrell pers. comm.
1995) with the two taxa easily distinguished on
leaf characters. While it was obvious that an
additional species of Clausena was represented,
it was also necessary to determine if more than
a single individual was extant and to collect
adequate material of flowers and fruit to enable
comparison with the taxa enumerated by Molino
(1994).
Visits during 1999 and 2000 to both Mon
Repos Conservation Park (last sighting 1984)
and the Baffle Creek district (last sighting 1920)
were unsuccessful in relocating populations
of the Clausena, however, these historical
records reinforce the existence of this taxon in
natural vegetation at more than a single locality.
Nearly all of the suitable vegetation type
(Regional ecosystem 12.2.2.: Mixed microphy 11/
notophyll rainforest on Quaternary coastal
dunes and beaches) in these areas has been
destroyed for housing or agriculture and is
considered as endangered (Young & Dillewaard
1999). A relatively intensive survey of this
Austrobaileya 5(4): 715-720 (2000)
vegetation type and similar vegetation on basalt
has been undertaken in the Bundaberg area
and around Hervey Bay (mostly unpublished
since the compilation of Forster et al. 1991) and
the Clausena has not been located in any of
the fifteen additional localities examined.
The recent collection of flowering
material of the Clausena at the Dundowran
locality has enabled its status to be assessed.
Based on the keys and sectional descriptions
of Molino (1994), this material can be placed in
Clausena section Clausena. It does not appear
to correspond to any of the six described
species in this section and is formally described
herein.
Clausena smyrelliana PI Forst. sp. nov.;
Clausenae brevistylae affinis, foliolis
superne nitidis marginibus integris
(adversum foliola obscure viridia
marginibus dentatis usque crenatis),
floribus 5-meris (adversum flores 4- rare
5-meros), alabastris oblongis (adversum
alabastra globosa), petalis lanceolatis
(adversum petala ovata usque
lanceolato-ovata) et staminibus 10
(adversum 6 vel 8 rare 10) differt.Typus:
Queensland. Wide Bay District:
Dundowran, 8 Nov 1999, P.I.Forster
PIF25182 (holo: BRI [1 sheet + spirit]; iso:
A, K, F, MEF, QRS).
Shrub or small tree to 4 m high. Bark smooth,
greenish-brown with irregular longitudinal
strips of cream-yellow lenticels; blaze yellowish
with strong citrus-scent; wood yellow-straw.
Feaves alternate, pinnate; rachis up to 170 mm
long; leaflets 6-10, alternate, conspicuously
dotted with oil glands and with a slight citrus-
like scent when crushed, petiolules 4—7 mm long,
yellow to pink; blades elliptic or ovate, often
unequal, 20-100 mm long, 10-68 mm wide,
glossy above, dull and somewhat glaucous
below; lateral veins 7-9 per side of midrib and
visible on both surfaces, interlateral veins more
visible below; apex acute-retuse or acuminate-
retuse, rarely mucronate; base obtuse to
oblique; margin entire, glandular, glabrous.
Inflorescences a terminal panicle, up to 90 mm
long. Flowers 5-merous, c. 10 mm long and 10
mm in diameter; pedicels 3-4 mm long, 0.8-1
Forster, Clausena Smyrelliana
mm in diameter, with conspicuous oil glands
and dense uniseriate trichomes; calyx 5 lobed,
1.1-1.5 mm long, irregularly ciliate; petals 5,
imbricate in bud, lanceolate and strongly
recurved, 7-7.5 mm long, 1.8-2 mm wide,
glabrous, cream. Stamens 10, filaments dilated
basally, 4-6 mm long; anthers c. 1 mm long and
1 mm wide, longitudinally dehiscent; style c. 6
mm long and 1.2 mm in diameter with an obtuse
tip; gynophore well developed, but not
markedly distinct from the ovary, c. 1 mm long
and 1 mm diameter; ovary 4-5 locular, c. 1 mm
long and 1 mm in diameter, ovules 2 per locule.
Fruits baccate, depressed-globose, soft-fleshy,
8-9 mm long, 10-11 mm in diameter, white, with
conspicuous oil-glands. Fig. 1.
Other specimens examined. Queensland. Port Curtis
Creek District: Baffle Creek District, Apr 1920,
C.T. White AQ152118 (BRI). Wide Bay District:
Dundowran, Oct 1995, Forster PIF17928 & Smyrell
(BRI); ditto, Jan 2000, Forster PIF25307 (BRI, MEL,
QRS); ditto. May 1992, Smyrell AQ542962 (BRI);
ditto, Nov 1992, Smyrell AQ563892 (BRI); ditto,
Sep 1991, Telford 11337 (BRI; BISH, CANB, NSW
n.v.[distributed as Euroschinus falcata ]); Mon Repos
Environmental Park [now Conservation Park] s.dat.
[71984], Randall 409 (BRI).
Notes: Clausena smyrelliana is compared to
C. brevistyla , the only other species in this
genus in Australia. The two species are easily
distinguished by the dentate-margined, dull
green foliage of the latter and the entire-
margined glossy green foliage of the former.
Other differences include C. smyrelliana
having yellow to pink petiolules, 5-merous
flowers, oblong buds, lanceolate petals and 10
stamens, whereas C. brevistyla has green
petiolules, 4-merous flowers (although 5 have
been rarely noted by Molino 1994), globose
buds, ovate to lanceolate-ovate petals and (6)
8 (10) stamens.
Clausena smyrelliana may possibly be
confused with Micromelum minutum and this
species is present in two of the localities that
the former has been recorded from.
M. minutum has dull green foliage with
irregularly crenate margins to the leaflets,
yellowish lateral nerves on the underside of
the leaf, linear-subulate filaments and on
maturity has orange-red hard-fleshy, ovoid
fruit, whereas C. smyrelliana has glossy green
foliage with entire margins to the leaflets, green
717
lateral nerves on the underside of the leaf,
basally dilated filaments and on maturity has
white, soft-fleshy, depressed-globose fruit.
C. smyrelliana may also be confused with
Euroschinus falcata Hook.f. (Anacardiaceae);
however, the former has conspicuous oil glands
in the leaf which are lacking in the latter.
The single individual of C. smyrelliana
at Dundowran has flowered and fruited
profusely nearly every year since 1992
(G Smyrell pers. comm. 1999 & pers. obs.). The
fruits contain viable seeds and many hundreds
of seedlings have been propagated from these.
The seedlings produce saplings that are
indistinguishable from the parent tree, although
their own potential viability is as yet unknown.
Presumably this species is bird dispersed and
why it is not more common both at Dundowran
and throughout the species range is a mystery.
Perhaps it is one of these species that requires
an exceptionally wet year for seedlings to
establish, as this also seems to be the case for
Alectryon ramiflorus where recruitment in the
wild is infrequent. Most of the 1990’s were
drought effected in south-east Queensland and
may have influenced the establishment of
seedlings of many vineforest plants.
Distribution: Three localities have been
recorded for Clausena smyrelliana , over a
longitudinal distance of 110 km between Baffle
Creek in the north and Dundowran in the south.
Only the Dundowran locality is currently known
to have an extant population.
Habitat: Clausena smyrelliana grows in
littoral microphyll vineforest within close
proximity to the sea (100-200 m distance). The
canopy of the vineforest at the type locality is
uneven, but more or less closed with no
emergents. Common canopy species include
Alectryon coriaceus, Argyrodendron sp. (Kin
Kin W.D.Francis AQ81198), Cleistanthus
cunninghamii , Diospyros fasciculosa and
Sterculia quadrifida.
Phenology: Clausena smyrelliana has been
recorded in flower from November to January
and ripe fruits have been collected in May.
Conservation Status: Clausena smyrelliana
may be regarded as critically endangered under
the IUCN categories ofA. Population reduction
718
Austrobaileya 5(4): 715-720 (2000)
Fig. 1. Clausena smyrelliana. A. branch with immature fruit. x0.4. B. undersurface of leaf showing lateral and
interlateral venation. x0.8. C. flower from side, x 4. D. bud from side, x 4. E. stamen, x 8. F. upper part of
pedicel, calyx and gynostegium. x8. G cross-section of gynophore and ovary showing ovules, x 8. H. fruit from
side, x 3. I, fruit from above, x 3. J. cross-section of immature fruit showing 5 locules. x 4. A & J from Forster
PIF25307 (BRI); B from Forster 17928 & Smyrell (BRI); C-G from Forster PIF25182 (BRI); H & I from
Smyrell AQ542962 (BRI). Del. W. Smith.
Forster, Clausena Smyrelliana
in the form of either of the following: 1. An
observed, estimated, inferred or suspected
reduction of at least 80% over the last 10 years
or three generations, whichever is the longer,
based on (and specifying any of the following:
(C) a decline in area of occupancy, extent of
occurrence and/or quality of habitat. B. Extent
of occurrence estimated to be less than 100
km 2 or area of occupancy estimated to be less
than 10 km 2 , and estimates indicating any two
of the following: 1. Severely fragmented or
known to exist at only a single location. D.
Population estimated to number less than 50
individuals. It is recommended that the
conservation coding of Endangered is given
to this species and that a Recovery Program is
initiated to conserve the species.
Clausena smyrelliana may be the most
threatened vascular plant species in
Queensland at this time. At least two other
rainforest species in south-east Queensland are
in similar, although less dire straits. Alectryon
ramiflorus S.T.Reynolds from the Childers area
(slightly inland from the localities for
C. smyrelliana) is known from 37 trees in the
wild (Barry & Young 1997) and Acronychia
littoralis T.Hartley & J.B.Williams from littoral
rainforest further south, is known from less than
20 trees in the wild in Queensland
(W.McDonald, pers. comm. Feb 2000). In both
instances, concerted searches in recent years
have added further localities and individuals
to the known populations of both these species.
It would be worthwhile to search for further
individuals of C. smyrelliana throughout the
known range of the species, and perhaps
further to the north around Deepwater and
Eurimbula National Parks where similar
vegetation is still extant.
Etymology: The species is named for Greg
Smyrell, an enthusiastic amateur botanist with
a long standing interest in rainforest
conservation, who brought this species to
attention.
719
Economic Uses: The leaf essential oil of
Clausena smyrelliana is unlikely to have
economic potential (J.J. Brophy pers. comm.
Jan 2000). Clausena smyrelliana is an
attractive shrub or small tree and has potential
as a garden plant, especially because of the
prolonged fruiting period.
Acknowledgements
Thanks to Greg Smyrell for persisting with his
belief in the validity of this species, Maureen
Schmitt for assistance in the surveys around
Bundaberg, Mon Repos and Baffle Creek, Peter
Bostock for comments on the manuscript and
translation of the diagnosis into Latin, Will
Smith for the illustrations, Aileen Wood for
some French translation and Joe Brophy for
unpublished data on the leaf essential oils.
References
Barry, S.J. & Young, P.A.R. (1997). Species Recovery
Plan. Alectryon ramiflorus S.Reynolds
Sapindaceae. Unpublished Report, Queensland
Department of Environment.
Beentje, H.J. (1994). Kenya Trees, Shrubs and Lianas.
Nairobi: National Museums of Kenya.
Coates Palgrave, K. (1977). Trees of Southern Africa.
Cape Town/Johannesburg: C.Struik Publishers.
Forster, PI. (1997). Rutaceae. In R.J.F.Henderson (ed.)
Queensland Plants Names and Distribution , pp.
184-188. Brisbane: Queensland Herbarium,
Department of Environment.
Forster, PI., Bostock, P.D., Bird, L.H. & Bean, A.R.
(1991). Vineforest Plant Atlas for South-east
Queensland. Brisbane: Queensland Herbarium.
Lockwood, G.B. (1984). The essential oil from leaves
of Clausena heptaphylla. Fitoterapia 55: 123—
124.
Madulid, D.A. (1995). A Pictorial Cyclopedia of
Philippine Ornamental Plants. Manila:
Bookmark.
Merrill, E.D. (1912). A Flora of Manila. Manila:
Bureau of Printing.
Austrobaileya 5(4): 715-720 (2000)
720
Molino, J.F. (1994). Revision du genre Clausena
Burm.f. (Rutaceae). Bulletin du Museum
National d’Histoire Naturelle 4 e series. Section
B Adansonia 16: 105-153.
Okunade, A.L. & Olaifa, J.I. (1987). Estragole: an acute
toxic principle from the volatile oil of the leaves
of Clausena anisata. Journal of Natural
Products 50: 990-991.
Young, P.A.R. & Dillewaard, H.A. (1999). Southeast
Queensland. In P.S.Sattler & R.D.Williams
(eds.). The Conservation Status of Queensland’s
Bioregional Ecosystems , Chapter 12. Brisbane:
Environmental Protection Agency.
The first botanical record for Australia
John F. P. Windolf
Summary
Windolf J.F.P. (2000). The first botanical record for Australia. Austrobaileya 5(4): 721-723.
Notes on the first identification and written record of a botanical species, Ximenia americana, in
the Commonwealth of Australia on 21st September 1606, probably at Long Island, Torres Strait,
as well as the historical background of the circumstances of its notation.
Key words: Prado, Ximenia americana , Torres Strait, Queensland, Australia
J.F.P. Windolf, 53 Pandanus Avenue, Coolum Beach, Queensland, 4573, Australia
Introduction
Throughout the botanical world the initial
record of a particular species in any defined
geographical or ecological region is a matter of
some importance, from both the scientific and
historical point of view. When such an event
relates to the first record of any kind for an
entire continent it takes on a much more
significant role. Australia is fortunate in that
many of the log books and diaries of early
European expeditions to this continent have
been preserved, and that their writers were men
of sufficient intellect to take a genuine interest
in the area’s natural science and to record what
they saw.
Historical background
The first proven European sighting of what is
now known as Australia was made by the
Dutchman Willem Jansz and his crew in March
1606 on the west coast of Cape York Peninsula.
Although a map relating to their activities is
still in existence, and we know from secondary
sources that they landed in several places and
explored a considerable section of coastline,
there is no known contemporary written
account of the expedition (Sharp 1963:17 &
Whittaker et. al. 1975:196), and it is necessary
to examine subsequent voyages to determine
who was the first to leave a record of his exploits
in the Australian region.
Some six months after Jansz, in September
1606, the Spanish navigator Luis Vaez de Torres
Accepted for publication 31 May 2000
traversed the strait separating Australia and
New Guinea, now named in his honour. Torres
was originally the second-in-command to
Pedro Fernandez de Quiros in what is generally
referred to as the 1606 Spanish South Seas
Expedition. The prime purpose of the
expedition was to continue the search for the
supposed southern continent then believed to
exist in the South Pacific. After crossing the
Pacific from Callao, Peru, and spending some
time in Espiritu Santo, Vanuatu, the fleet
became separated. Quiros returned to
Acapulco, Mexico, in the San Pedro y San Pablo
while Torres set out for Manila in the Philippines
with the other two ships, the San Pedro and a
smaller vessel called Los Tres Reyes, during
which time he sailed along the south coast of
New Guinea, discovering the strait now named
after him in the process.
There are two extant manuscripts
describing this section of the expedition: a
formal letter thatTorres wrote to King Philip III
of Spain soon after his arrival in Manila on 22
May 1607, and an extensive relation penned
by an entretenido, or “gentleman volunteer”,
named Don Diego de Prado y Tovar (Don Diego
de Prado y Tovar orig. ms.. Stevens 1930 &
Windolf in prep.) who was on board the San
Pedro during this period, and it is in this
account that we find the first record of an
identifiable botanical species from what is now
Australia. Prado had no known officially
designated capacity on board, but he was an
astute observer of many aspects of the natural
world, and made numerous notes on his
observations.
722
Dating and location of the observation
The expedition had entered what is now
Australian territory on or about 11 September
1606 (Gregorian dating). They spent the next
three weeks finding a way through the maze of
islands, reefs and sandbars that fill Torres Strait
before exiting into theArafura Sea on 4 October
1606. This dating is based on the date in Peru
carried westward without any correction for
crossing the International Date Line, and as
such is one day behind Australian Eastern
Standard Time (AEST). It can be independently
verified due to an eclipse of the moon, which
Prado recorded. The details of this eclipse have
been calculated, and it occurred on 16
September GMT, or 15 September Ship Time
(Kelly 1966:255). It is a relatively simple matter
to assign dates to their day-to-day activities at
this time in relation to this eclipse, and we find
that the relevant botanical discovery took place
in the late afternoon of Thursday 21 September
1606AEST.
Several researchers have attempted to
plot the expedition’s route through the strait,
but due to a variety of reasons no universal
consensus has been arrived at. Prado named
the island concerned Isla de Vulcan (Volcan)
Quemado, because of the amount of pumice
stone that they saw there, the English
translation of the Spanish name being “Island
of Extinct Volcano”. (Stevens 1930:163 &
Windolf in prep.). Unbeknown to them, this
pumice had nothing to do with the geology of
the island itself, having been borne there on
ocean currents. The best known attempt to
identify individual islands is that by Brett Hilder
(Hilder 1980), who thought that Volcan
Quemado was probably Long Island (Hilder
1980:81). However, because of the
topographical similarities between many of the
islands in central Torres Strait, and the often
less than comprehensive description of their
appearance given by Prado, the present author
considers that the exact identification ofVolcan
Quemado remains unresolved. Identification of
the general area, however, presents no problem,
and there is no doubt that the site of the
observation is well within the boundaries of
the State of Queensland.
Austrobaileya 5(4): 721-723 (2000)
The species
The species noted by Prado is identified as
Ximenia americana L. It is widespread in
tropical regions (Willis 1973:1232) and in the
Americas is often referred to as the Nicaraguan
Plum. This is an important point in its
identification when relying on Prado’s own
words describing the plant:
...hallamos...muchos arboles de siruelas que
llaman de nicaragua, son de grande guesco y
poco carne.
(Prado, orig. ms. & Stevens 1930:162).
This is translated as: “...we found...many plum
trees that are named after Nicaragua: they have
large stones and little flesh.” (Windolf in prep.).
Ximenia americana is a scrambly shrub
or small tree up to 5 m tall, the fruit being a
typical plum-like, pyriform or globular drupe,
yellow in colour. On islands, and near the coast,
it tends to grow on sand dunes and in forests
on the landward side of mangroves (George
1984:15-16).Longlsland(10 02’S 142 51’E)
itself is generally low and swampy, but heavily
wooded. The main island lies on the
northwestern end of the Long Island Reefs
complex and there are a number of small
mangrove-covered islets along the northern
and northeastern sides of the reefs (NP15
1973:233). The species has been collected from
three islands in Torres Strait (Dauan, Yorke and
Murray) in modern times as well as from
numerous localities in tropical Queensland and
New Guinea (Pers. comm. Queensland
Herbarium).
The fruit is considered edible, but is
reported as sometimes being purgative. It
contains appreciable quantities of oil rich in
ximenic acid (George 1984:16). Cribb states that
the fat extracted from the seeds is used as a
substitute for ghee in parts of India (Cribb
1982:46). It is not known whether it was used
for this purpose in the Americas, or whether
the Torres Strait Islanders or Australian
Aborigines used it as a food source.
723
Windolf, First Botanical Record for Australia
The genus is named after Francisco Ximenes,
a Spanish naturalist who wrote extensively on
the subject of medicinal plants in the early
sixteen-hundreds.
This species is relatively common in tropical
America and Prado was probably familiar with
it there. Given the confidence of his statement,
the correlation between his description and the
actual appearance of the fruit, and the ecological
and geographical affinity with sites where
Ximenia americana is known to occur, there
seems no reason to question the correctness
of his identification.
Conclusions
It is considered proven that the first recorded
identification of any botanical species in the
Commonwealth of Australia was that of
Ximenia americana L., made by Don Diego de
Prado y Tovar on Thursday 21 September 1606
AEST (Gregorian dating) in Torres Strait,
possibly on Long Island, in the botanical
district of Cook, Queensland.
Acknowledgements
Acknowledgement is due to the Director and
staff of the Queensland Herbarium and to Philip
Sharpe for providing details of Ximenia
americana , and to my dear wife Frances for
her reading of the text and useful suggestions.
References
Admiralty Chart 2321. Torres Strait and Approaches.
Cribb, A.B. & J.W. (1981). Useful Wild Plants in
Australia. Sydney: Collins.
George, Alexander S (Ed.). (1984). Flora of Australia.
Volume 22. Canberra: Aust. Gov. Pub. Serv.
Heeres, J.E. (1899). The Part Borne by the Dutch in
the Discovery of Australia 1606-1765. London.
Hilder, Brett. (1980). The Voyage of Torres. Brisbane:
University of Qld. Press.
Kelly, Celsus O.F.M. (1966). LaAustrialia del Espiritu
Santo. Cambridge: Cambridge Uni. Press.
NP15. (1973). Australia Pilot. Volume III. Sixth
Edition. Hydrographic Department. Taunton.
Prado y Tovar, Don Diego de. Relacion Summaria de
Don Diego de Prado y Tovar. Original
Manuscript, Safe l/73a, Mitchell Library,
Sydney.
Sharp, A. (1963). Discovery of Australia. Oxford Uni.
Press.
Stevens, Henry N. (ed.) and Barwick (trans.). (1930).
New Light on the Discovery of Australia as
revealed by the Journal of Don Diego de Prado
y Tovar. Hakluyt Soc. Ser. II, no. 64. Cambridge:
Cambridge Uni. Press.
Willis, J.C. (1973). (Revised by Airy Shaw). A
Dictionary of the Flowering Plants and Ferns.
Cambridge: Cambridge Uni. Press.
Whittaker, J.L., Gash, N.G., Hookey, J.F., Lacey, R.J.
(1975). Documents and Readings in New
Guinea History. Prehistory to 1889. Brisbane:
The Jacaranda Press.
Windolf, John F. P. (in prep). The Life and Travels of
Don Diego de Prado y Tovar.
Rediscovery of Dischidia torricellensis (Schltr.) P.I.Forst., an
unusual epiphytic asclepiad from New Guinea
Paul I. Forster
Summary
Forster, PI. (2000). Rediscovery of Dischidia torricellensis (Schltr.) PI.Frost., an unusaul
epiphytic asclepiad from New Guinea. Austrobaileya 5(4):725-728. An amplified description
and illustrations are provided for Dischidia torricellensis (Schltr.) P.I.Forst.. The first collection
in over 60 years is reported from a new locality in Chimbu Province in Papua New Guinea.
Keywords: Dischidia torricellensis, Asclepiadaceae, New Guinea.
P I. Forster, Queensland Herbarium, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong,
Queensland 4066, Australia
Introduction
Many of the Asclepiadaceae that occur in
Papuasia are poorly known, having been
collected only once or not more than a few
times. At least 192 species are considered to
naturally occur in the region, with some 164
endemic species (Forster 1996). A great number
of these have yet to be adequately illustrated
and patterns of variation remain to be
rigorously documented for nearly every single
species. One such poorly known species is
Dischidia torricellensis{ Schltr.) P.I.Forst. last
collected in 1939 (Forster 1990).
Much of the early documentation of the
asclepiad flora of New Guinea was undertaken
by the remarkable Rudolf Schlechter (Nicholas
1992). Many of the plants that he found and
later described are still only known from his
original collections or have been rarely
collected since. Schlechter went to the then
German New Guinea to search for sources of
rubber or rubber substitutes, but spent much
of his time collecting plant specimens,
particularly of Orchidaceae, but also of other
plants including Asclepiadaceae. InApril 1902
Schlechter was in the Torricelli Mountains of
northeastern New Guinea (now in Madang
Province of Papua New Guinea). His collection
numbered 14445 represented an unusual
epiphytic subshrub and he subsequently
described it as the new genus and species
Accepted for publication 7 April 2000
Spathidolepis torricellensis Schltr. (Schlechter
1905). Schlechter considered his new plant to
be allied to Dischidia but differing in the small
coronal lobes and thin, leathery leaves.
In more recent times I came to examine
his work and this particular species and made
the conclusion that S. torricellensis could be
adequately accommodated within Dischidia
R.Br. (Forster 1990). There is considerable
variation in Dischidia s.l. and while there are
recent regional accounts for the Malay
Peninsula (Rintz 1980) and Australia (Forster
& Liddle 1996), there is no monograph available
for the group. Despite this, the basic patterns
of morphological variation are reasonably well
known. Perhaps having overlooked the transfer
of Spathidolepis into Dischidia , Johns (1995)
listed Spathidolepis as being one of two
endemic genera of Asclepiadaceae present in
New Guinea.
Dischidia torricellensis is an unusual
species within Dischidia for a number of
reasons. Firstly the leaves are herbaceous
(Fig. 1 A), whereas most species have fleshy to
succulent leaves. Secondly the corolla lobes
are distinctive in the way the edges are strongly
reflexed (Fig. 1C). Thirdly the pollinaria are not
typical for the majority of Dischidia species in
the + unwinged nature of the caudicles (Fig.
IF); however, at least one other species
(D. superba Rintz) also has this feature (Rintz
1979).
726
A recent (1997) excellent collection
(including spirit material) of D. torricellensis
has enabled re-examination of the floral
structure (Fig. 1) and prompted the present
contribution. Data from this collection have
confirmed my previous conclusions as to the
generic placement of this species but have
enabled a few additional observations to be
made. Firstly the staminal corona lobes of
D. torricellensis are not particularly small (as
mentioned by Schlechter 1905), but can be quite
well developed and markedly fleshy structures
(Fig. 1D,E). They are not similar to the staminal
corona lobes found in many species of
Dischidia that are generally membranous and
with incurved upper lobes (e.g. D. bengalensis
Colebr. or D. subulata Warb.). On the other
hand they are somewhat similar, albeit fleshier,
to those illustrated forD. parvifolia Ridl. from
Malaysia (Rintz 1980). Many species of
Dischidia have very poorly developed staminal
coronal lobes (e.g. D. imbricata (Blume)
Steud.), thus demonstrating that there is not
only considerable variation in this feature, but
that it varies from being virtually non-existent
to well developed. Such a gradation is repeated
in other asclepiad groups (e.g. Marsdenia R.Br.
s.l. Forster 1995; or various Stapelieae) and can
be interpreted as being more useful at the
specific level rather than generic. Rintz (1980)
has quite clearly illustrated that there is a
complete gradation from those species where
the staminal coronal lobes are poorly developed
to those where the lobes are prominent and
often incurved, hence Schlechter’s justification
for recognition of Spathidolepis on the basis
of the small coronal lobes is unwarranted.
Dischidia torricellensis also is
unusual within the genus in not being an
obvious climber or twiner. In the previous
description (Forster 1990) it was stated that the
species was a ‘liana’, presumably based on the
“small root climber” label data of Brass 12915.
The collection of Takeuchi 11736 dispels this
notion. This species seems to have a habit
akin to that found in some species of the closely
allied genus Hoya R.Br. where a small number
Austrobaileya 5(4): 725-728 (2000)
of species have either pendent non-twining
stems, or erect, non-twining stems as opposed
to the majority that are twiners (Forster et al.
1998) or in the allied MicholitziaN .E.Br. from
south-east Asia (Goyder & Kent 1994).
The arrangement of the hairs in the
corolla tube ofD. torricellensis fits the pattern
of Group C (corolla throat only pubescent) of
Rintz (1980). Whether or not Rintz’s groups
based on the arrangement of hairs on the corolla
are natural remains to be seen. Certainly in
terms of foliage and pollinaria, D. torricellensis
does not closely resemble the Malayan species
of Rintz’s Group C and its position within the
genus must be viewed as being isolated. These
hairs are antrorse near the corolla mouth but
retrorse just above the staminal column. In
D. torricellensis , these hairs effectively block
access to the staminal column and nectar
source at the base. This arrangement tends to
indicate that a specialised pollination syndrome
is involved and that D. torricellensis is similar
to the majority of Dischidia species in this
respect. Pauw (1998) has recently speculated
that this type of asclepiad flower (a closed tube
with access restricted by hairs) is adapted to
bird pollination, based on this syndrome being
observed in the morphologically similar flowers
of Microloma species from southern Africa.
To date there are no available observations on
the pollinators of any species of Dischidia.
Similar sized, but open mouth campanulate
flowers in Marsdenia cymulosa Benth. are
visited and possibly pollinated by small
chloropid flies (Forster 1992) and perhaps
similar sized insects, rather than birds pollinate
flowers of Dischidia.
Dischidia torricellensis (Schltr.) P.I.Forst.,
Austrobaileya 3: 288 ( 1 990).
Spathidolepis torricellensis Schltr., in
K.Schum. & Lauterb., Nachtrage FI.
Schutzgeb. Stidsee 356 (1905). Type:
Papua New Guinea [Kaiser-Wilhemsland].
Madang Province: Torricelli-
Gebirges, April 1902 , R.Schlechter
14445 (lecto: K (photo atBRI!); isolecto:
BO!), fide Forster (1994: 515).
Forster, Dischidia torricellensis
Epiphytic subshrub, branches up to 50 cm long
and with white latex; indumentum on foliage
comprising simple, uniseriate trichomes. Stems
cylindrical, up to 3 mm diameter, with antrorse
indumentum; internodes up to 90 cm long.
Leaves petiolate, herbaceous, lanceolate-
elliptic, 3-9 cm long, 1-3.2 cm wide; apex
caudate to cuspidate, obtuse at tip; base
cuneate; lateral venation comprising 16 to 18
veins per side of the midrib, largely indistinct,
interlateral venation reticulate and largely
indistinct; upper surface medium green, midrib
sunken, glabrous; lower surface glaucescent,
midrib raised, glabrous or with scattered,
antrorse indumentum on margins and midrib;
colleters 2 at lamina base. Inflorescence
persistent, an umbelliform raceme up to 8 mm
long; bracts triangular, 0.5-1 mm long, 0.5-1
mm wide, with sparse indumentum; peduncle
up to 10 mm long and 2 mm diameter. Flowers
urceolate, (3) 4-5.5 mm long, (2) 4-5 mm
diameter at base and (2.5) 3-4.5 mm diameter at
mouth; pedicels 2-4 mm long, 0.5-1 mm
diameter, with sparse to dense indumentum;
sepals oblong to obtuse-ovate, 0.7-2 mm long,
0.9—1 mm wide, ciliate and with 1 or 2 glands at
base of each sinus. Corolla fleshy, green when
immature, white at anthesis; tube (2) 3.5^1 mm
long, (2) 4-5 mm diameter, externally glabrous
or minutely papillose, internally glabrous or
with a few isolated hairs; lobes erect, triangular-
ovate, fused for two-thirds of length, each
strongly jointed in middle and with margins
strongly reflexed, 1.8-2 mm long, 2-2.2 mm wide,
externally glabrous, internally with dense hairs
to 1 mm long blocking entrance to tube.
Staminal corona (1) 2-2.5 mm long, (2) 3-3.5
mm diameter, attached at bottom of staminal
column and comprising 5 separate lobes; each
lobe spathulate-obovate and recurved or
winged towards base on either side, the entire
lobe (0.75) 2-2.5 mm long, (0.75) 1.5-1.7 mm
wide, the wings (0.3) 0.4—0.6 mm wide. Staminal
column 1.5-2 mm long, 1-2 mm diameter; anther
appendages oblong-obtuse, 0.5-0.8 mm long,
0.2-0.3 mm wide; fissure between anther wings
0.6-0.8 mm long. Style-head oblong-conical,
0.7-0.8 mm long; ovaries 1-1.5 mm long,
glabrous. Pollinaria c. 0.6 mm long and 0.4 mm
wide; polliniaheld erect, oblong, 0.35-0.4 mm
long, 0.1-0.2 mm wide, yellow; corpusculum
727
oblong, 0.2-0.3 mm long, 0.1-0.2 mm wide;
caudicles somewhat winged near pollinia, 0.15-
0.18 mm long, c. 0.1 mm wide. Follicles fusiform
(immature), 110-120 mm long, c. 2 mm diameter,
glabrous. Fig. 1.
Specimens examined: West Papua. Jayapura: 6 km
SW of Bernhard Camp, Idenburg River [3° 28’S, 139°
08’E], Feb 1939, Brass 12915 (BRI; A n.v.); Rouffaer
River [not localised], Sep 1926, Docters van Leeuwen
10275 (BO, L). Papua New Guinea. Chimbu
Province: Crater Mountain Wildlife Management area,
east of Haia Village, 6°43’S, 145° 00’E, Mar 1997,
Takeuchi 11736 (BRI; LAE n.v.).
Typification: Schlechter’s original collection
of this plant would have been deposited at B
although he did not specify this (Schlechter
1905, 1913). This particular specimen is not
extant having been destroyed in the
firebombing of B in World War II. A duplicate
of his number 14445 present at K was selected
as lectotype and a further duplicate at BO as
an isolectotype for the name (Forster 1994).
Further duplicates of this number have not been
located in the herbariaA, CANB, L, MEL, NSW,
SING, WRCL where some New Guinean
Schlechter material is extant.
Distribution: Dischidia torricellensis has now
been recorded from four places. Three of these
collections predate 1940. One of these is
obscure and the type locality is very broadly
defined. The Takeuchi collection confirms the
continued existence of this species in the wild
after an interval of 60 years and extends the
range considerably eastwards. It is not
inconceivable that this species has a broad
distribution in suitable habitats between 700
and 1200 m over much of upland New Guinea.
Habitat: This plant has been collected from
midmontane rainforest at altitudes between 700
and 1200 m. It occurs as an epiphyte on
branches.
Acknowledgements
Thanks to Wayne Takeuchi for the specimens,
Peter Bruyns for the illustrations and Peter
Bostock for comments on the manuscript.
Various herbaria have allowed access to
collections in situ or on loan over an extended
period of time.
728
Austrobaileya 5(4): 725-728 (2000)
References
Forster, P.I. (1990). Notes on Asclepiadaceae, 2.
Austrobaileya 3: 273-289.
Forster, P.I. (1992). Insects associated with the
flowers of Marsdenia cymulosa Benth.
(Asclepiadaceae) and their possible role in
pollination. Australian Entomological
Magazine 19: 45-47.
Goyder, D.J. & Kent, D.H. (1994). Micholitzia
obcordata N.E.Br. (Asclepiadaceae:
Marsdenieae) reinstated. Asklepios 62: 13-19.
Johns, R.J. (1995). Malesia - an introduction. Curtis’s
Botanical Magazine 12: 52-62.
Nicholas, A. (1992). The Asclepiadaceous works of
Rudolf F. Schlechter (1872-1925). Willdenowia
22: 215-264.
Forster, P.I. (1994). Type collections of
Asclepiadaceae at Herbarium Bogoriense (BO).
Australian Systematic Botany 7: 507-519.
Forster, P.I. (1995). Circumscription of Marsdenia
(Asclepiadaceae - Marsdenieae) with a revision
of the genus in Australia and Papuasia.
Australian Systematic Botany 8: 703-933.
Forster, P.I. (1996). A checklist of the Asclepiadaceae
of Papuasia. Science in New Guinea 22: 15-
22 .
Forster, P.I. & Liddle, D.J. (1996). Dischidia and Hoya
(Asclepiadaceae). In A.G.Wilson (ed.). Flora
of Australia 28: 231-240. CSIRO Publishing,
Melbourne.
Forster, PI., Liddle, D.J. & Liddle, I.M. (1998).
Diversity in the genus Hoya (Asclepiadaceae:
Marsdenieae). Aloe 35: 44-48.
Pauw, A. (1998). Pollen transfer on birds’ tongues.
Nature 394: 731-732.
Rintz, R.E. (1979). Three new species of
Asclepiadaceae from Peninsular Malaya.
Blumea 25: 225-231.
Rintz, R.E. (1980). The peninsular Malayan species
of Dischidia (Asclepiadaceae). Blumea 26: 81-
126.
Schlechter, R. (1905). Asclepiadaceae. In K. Schumann
& K. Lauterbach, Nachtrage zilr Flora der
Deutschen Schutzgebiete in der Siidsee.
Leipzig: Gebruder Borntrager.
Schlechter, R. (1913). Die Asclepiadaceen von
Deutsch-Neu-Guinea. Botanische Jahrbucher
fiir Systematik, Pflanzengeschichte und
Pflanzengeographie 50: 81-164.
Fig. 1. Dischidia torricellensis . A, flowering branch. B, bud. C, side view of flower. D, side view of dissected
flower. E, side view of staminal column and staminal corona with one corona-lobe removed. F, pollinarium.
Scale bars: A, 10 mm; B, C, 1 mm (at B); D, 1 mm; E, 0.5 mm (at B); F, 0.25 mm (at D). All from Takeuchi
11736 (BRI). Del. P.V.Bruyns.
Austrobaileya 5(4): 729 (2000)
Note
Cryptolepis papillata P.I.Forst. & Sarcolobus porcatus P.I.Forst.
(Asclepiadaceae), newly recorded from West Papua
Systematic collection of the flora of the Birds
Head Peninsula ofWest Papua on the island of
New Guinea is currently being undertaken by
staff of the National Herbarium of the
Netherlands, Leiden branch and the Herbarium
Bogoriense( Veldkampetal. 1997). Amongst a
number of Asclepiadaceae sent for routine
identification are two collections that represent
the first records for those particular species
from West Papua. As both species have been
rarely collected it is felt worthwhile to formally
document these records.
Cryptolepis papillataP.I.Forst., Austrobaileya
3:277(1990).
West Papua. Birds Head Peninsula, surroundings of
Ayawasi, 1°09’S, 132° 29’E, Sep 1995, Ave 4059 (BRI;
L n.v.).
Previously known from Papua New Guinea in
Morobe Province. This new collection extends
the known range considerably westward.
Three species of Cryptolepis (C. papillata , C.
perakensis (Gamble) P.I.Forst., C. lancifolia
P.I.Forst.) have now been recorded for West
Papua (Forster 1990, 1991a, 1993a, 1996).
Sarcolobus porcatusP.I.Forst., Austrobaileya
3:353(1991).
West Papua. Birds Head Peninsula,surroundings of
Ayawasi, 1°09’S, 132°29’E, Mar 1996, Ridsdale 2321
(BRI; L n.v).
Previously known from Papua New Guinea in
Morobe Province. This new collection extends
the range considerably westward. Four
species of Sarcolobus ( S. porcatus , S. retusus,
K.Schum., S. secamonoides(Sch\tr.)V.l.¥orst.,
S. vittatus P.I.Forst.) have now been recorded
for West Papua (Forster 1990, 1991b, 1993b,
1996).
Acknowledgement
Thanks to M. Polak (L) for sending the material
for identification.
References
Forster, P.I. (1990). Notes on Asclepiadaceae, 2.
Austrobaileya 3: 273-289.
Forster, P.I. (1991a). Cryptolepis lancifolia
(Asclepiadaceae: Periplocoideae), a new species
from West Papua. Blumea 35: 381-383.
Forster, P.I. (1991b). A taxonomic revision of
Sarcolobus R.Br. (Asclepiadaceae:
Marsdenieae) in Australia and Papuasia.
Austrobaileya 3: 335-360.
Forster, PI. (1993a). Conspectus of Cryptolepis R.Br.
(Asclepiadaceae: Periplocoideae) in Malesia.
Austrobaileya 4: 67-73.
Forster, P.I. (1993b). Additional records for some
species of Finlaysonia Wallich, Gymnanthera
R.Br, Heterostemma Wight & Arn. and
Sarcolobus R.Br. (Asclepiadaceae) in Melanesia
and Papuasia. Austrobaileya 4: 129-130.
Forster, P.I. (1996). A checklist of the Asclepiadaceae
of Papuasia. Science in New Guinea 22: 15-
22 .
Veldkamp, J.F., Roos, M.C. & Rifai, M.A. (eds.) (1997).
Expeditions and other fieldwork. Flora Malesia
Bulletin 12(2): 37-39.
P.LForster
Queensland Herbarium, Brisbane Botanic Gardens, Mt Coot-tha Road,Toowong, Queensland
4060
Austrobaileya 5(4): 731 (2000)
Note
A new combination in Morinda L. (Rubiaceae) for Australia
As a result of the publication of Queensland
Plants: Names and Distribution (Henderson
1997), it has come to our attention that there is
a nomenclatural problem regarding the eastern
and northern Australian species recorded
therein under Rubiaceae as Morinda acutifolia
F.Muell.
George Bentham (1867) published the name
Coprosma acutifolia for material collected by
Goodwin and Dallachy at Durandoo, New
South Wales, and sent to him in 1856 by
Ferdinand Mueller in Melbourne. This material
had been labelled Coprosma acutifolia by
Mueller.
Unfortunately, by 1867 the name
Coprosma acutifolia had already been
published by Joseph Hooker for a different
species of Coprosma from New Zealand
(Hooker 1857). Thus, Mueller’s name, when
published by Bentham, was a later homonym
and hence illegitimate under Article 53.1 of the
International Code of Botanical Nomenclature
(ICBN) (Greuter et al. 1994).
Mueller was aware of the problem with
the binomen Coprosma acutifolia F.Muell. ex
Benth. for he published (Mueller 1869) a brief
note on this and provided the new name
Coprosma canthoides F.Muell. for the
Australian species concerned.
However, he apparently overlooked this
note and combination when he named this
species in Morinda as M. acutifolia F.Muell.
(Mueller 1875). There he cited the name
Coprosma acutifolia F.Muell. ex Benth. as a
synonym but did not mention C. canthoides
F.Muell. By including Coprosma acutifolia
F.Muell. ex Benth. (as ‘F.M. in Bentham’), worn.
illeg., in his protologue, Mueller is taken as
providing a new name for the species dating
from 1875 (ICBN Art.58.3) but the type of his
name is the type of Coprosma acutifolia
F.Muell. ex Benth. (ICBN Art.7.3).
As the type of Coprosma acutifolia
F.Muell. ex Benth. is automatically the type of
C. canthoides F.Muell. (ICBN Art.7.3), and is
also the type of Morinda acutifolia F.Muell.,
according to Article 52.1 of the ICBN, Mueller
in 1875 published an illegitimate name which is
to be rejected. The name that Mueller ought to
have provided for the species he circumscribed
as Morinda acutifolia is M. canthoides.
Accordingly, the necessary new
combination is now provided.
Morinda canthoides (F.Muell.) Halford &
R.J.F.Hend. comb. nov. Coprosma
canthoides F.Muell., Fragm. 7:45 (1869);
Coprosma acutifolia F.Muell. ex Benth.,
FI. Austral. 3: 429 (1867), worn, illeg. non
Hook.f., J. Linn. Soc. 1: 128 (1857);
Morinda acutifolia F.Muell., Fragm 9:179
(1875), nom. illeg. Type: [Australia, New
South Wales.] Durandoo, undated, Hb.
F. Mueller (holo: K n.v.\ BRI, photo).
References
Bentham, G (1867) Coprosma. In Flora Australiensis
3:429-430. London: L. Reeve & Co.
Greuter, W., Barrie, F.R., Burdet, H.M., Chaloner, W.G.,
Demoulin, V., Hawksworth, D.L., Jorgensen, P.M.,
Nicolson, D.H., Silva, PC., Trehane, P. &
McNeill, J. 1994. International Code of
Botanical Nomenclature (Tokyo Code).
Regmim Vegetabile 131. Konigstein, Germany:
Koeltz Scientific Books.
Henderson, R.J.F. (ed.) 1997. Queensland Plants:
Names and Distribution. Brisbane; Department
of Environment.
Hooker, J.D. 1857. On the Botany of Raoul Island,
one of the Kermadec group in the South Pacific
Ocean. J. Linn. Soc. (Bot.) 1:125-129.
Mueller, F. 1869. Fragmentia Phytographiae
Australiae 7:45. Melbourne: Victorian
Government.
Mueller, F. 1875. Fragmentia Phytographiae
Australiae 9:179. Melbourne: Victorian
Government.
Accepted for publication 8 October 2000
David A. Halford and Rodney J. F. Henderson,
Queensland Herbarium, Brisbane Botanic Gardens Mt Coot-tha,Toowong, Qld 4066,Australia
Note
Austrobaileya 5(4): 733-734 (2000)
A new combination inProstantheraLabiM. (Lamiaceae)
In 1904, F.M. Bailey named Hemigenia
clotteniana , based on a specimen from
“Herberton” (Bailey 1904). He gave a detailed
description of the plant, including the following
“leaves opposite, simple, or on the branchlets
often in threes, 6 to 9 lines [12-18 mm] long,
scarcely exceeding 2 lines [4 mm] in width” and
later “calyx 2-lipped, lips entire; upper one deep
purple, about 4 lines [8 mm] long”. An
examination of the type showed that it
possesses 4 perfect stamens, each with 2 anther
cells.
In the following year (Bailey 1905), he
described Prostanthera atroviolacea, again
from “Herberton”. His description included the
following “leaves opposite, not exceeding 1
inch [25 mm] long and 2 lines [4 mm] broad”
and later “calyx silky-hairy, the upper lip 5 lines
[10 mm] long,... deep-violet outside”.
Bailey did not compare P. atroviolacea with
H. clotteniana , saying merely that “the nearest
ally of this plant \P atroviolacea] appears to
be P. lithospermoides”.
Between the years 1905 and 1999, only one
collection of either ‘species’ was added to the
Queensland Herbarium. That specimen,
collected by Michael Lockyer in 1974, was
identified as Prostanthera atroviolacea.
In April 1999, K.R. McDonald sent to BRI two
labiaceous specimens, collected from the rocky
hills west of Ravenshoe. The process of
identifying these specimens led to a closer
examination of Hemigenia clotteniana and
Prostanthera atroviolacea , and it was soon
realised that the two are synonymous. There is
some variation in the stamens; some flowers
have two perfect stamens while others have
four. This may have prompted Bailey to place
his two collections into different genera.
The correct genus is undoubtedly
Prostanthera , because the stamens always
have two perfect cells, and the calyx is 2-lobed.
Hence, a new combination is necessary, as
follows:
Prostanthera clotteniana (F.M.Bailey)
ARBeancomb. nov.
Hemigenia clotteniana F.M.Bailey,
Queensl. Agric. J. 15: 493 (1904). Type:
Herberton, undated, J. Stirling (holo:
BRI).
Prostanthera atroviolacea F.M.Bailey,
Queensl. Agric. J. 16: 190 (1905). Type:
Herberton, undated, R.C. Ringrose (holo:
BRI), syn. nov.
Illustration: F.M. Bailey, Comprehensive Cat.
ofQldPl.p. 393(1913).
Additional specimens examined: Queensland. North
Kennedy District: W of Ravenshoe, Dec 1974, Lockyer
s.n. (BRI); TR245 W of Ravenshoe, Apr 1999,
McDonald 22, 32 (BRI); TR 245 near Ravenshoe,
May 1999, Thompson & McDonald (BRI).
Conservation status: H. clotteniana was listed
as presumed extinct (X) on the schedules of
the Queensland Conservation Act, while P.
atroviolacea was listed as rare (R). A re¬
assessment of the conservation status of
Prostanthera clotteniana is now obviously
necessary. It is known only from a few rocky
hilltops in State Forest near Ravenshoe. Three
stands are known with 1,7 and 1 individuals in
each (K. McDonald pers. comm.). Applying the
criteria of the IUCN (Anon. 1994), a category
of‘critically endangered’ is proposed (Criterion
D).
Acknowledgements
I am grateful to Keith McDonald whose dogged
efforts to debunk the ‘X’ status of Hemigenia
clotteniana have finally paid off.
Accepted for publication 10 March 2000
Austrobaileya 5(4): 733-734
734
References
Anonymous (1994). IUCN Red List Categories.
International Union for Conservation of
Nature and Natural Resources: Switzerland.
Bailey, F.M. (1904). Contributions to the Flora of
Queensland. Queensland Agricultural Journal
15: 491-5.
Bailey, F.M. (1905). Contributions to the Flora of
Queensland. Queensland Agricultural Journal
16: 189-93.
A.R. Bean
Queensland Herbarium, Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha Road,Toowong,
Queensland 4066
Austrobaileya 5(4): 735-736 (2000)
Note
A new combination in Corymbia ‘section Politaria’: C. citriodora
subsp. variegata (Myrtaceae).
Introduction
Hill and Johnson (1995) informally established
Corymbia section ‘ Politaria ’ to comprise four
species: Corymbia citriodora (lemon-scented
gum), C. maculata (spotted gum), C. henryi
(large-leaved spotted gum) and C. variegata
(spotted gum). However, a recent study by
McDonald et al. (2000) has shown that the
taxonomic status of C. variegata warrants
reappraisal. They presented evidence that
C. citriodora and C. variegata could not be
distinguished by either allozymes or
morphology. They found that the main attribute
distinguishing them was leaf oil composition
(citronellal is the main leaf oil in C. citriodora
and a-pinene in C. variegata ). The two
‘species’ were thus considered to represent a
single taxon comprising two chemotypes.
Following field and herbarium studies
(particularly by ARB) we concur with
McDonald et al. (2000) and here formally
subsume C. variegata under C. citriodora at
subspecies rank. This will continue to allow a
distinction between the two entities, while more
accurately reflecting the closer relationship of
C. variegata to C. citriodora than to
C. macidata.
Most phytogeographical references on
eucalypts, e.g. Blakely (1934), Boland et al.,
(1984), Chippendale (1988), Brooker et al. (1997)
and Brooker and Kleinig (1999), treated
C. citriodora subsp. variegata as the northern
extension of C. maculata ., although
Chippendale (l.c.) lists Eucalyptus variegata
as a synonym of E. citriodora. The presence
of lemon-scented leaf oils, rather than
morphological attributes, appears to have
preoccupied the taxonomic approach of these
authors. This is despite the findings of Maiden
(1920), McKern (1954) and Larsen (1965), who
also considered the two represented a single
taxon comprising two chemotypes.
Accepted for publication 14 July 2000
McDonald et al. (2000) also concluded
that the two other ‘Politaria’ species,
C. maculata (Hook.) K.D.Hill & L.A.S. Johnson
and C. henryi (S.T.Blake) K.D.Hill &
L.A.S.Johnson, represented vicariads as they
were genetically and morphologically allied,
while genetically distinct from the
C. citriodora-C.variegata alliance. However,
we consider that little would be gained by a
change to infraspecific rank for these species
as their relationship is well established and the
nomenclatural change unnecessarily
disruptive.
Taxonomy
Corymbia citriodora (Hook.) K.D.Hill &
L.A.S.Johnson subsp. variegata
(F.Muell.) A.R.Bean & M.W.McDonald
comb, et stat. nov.
Eucalyptus variegata F.Muell., J. Linn.
Soc., Bot. 3: 88 (1859); Corymbia
variegata (F.Muell.) K.D.Hill &
L.A.S.Johnson, Telopea 6: 389 (1995).
Type: Queensland. Burnett River, 1856,
F. Mueller (holo: MEL; iso: K).
Corymbia citriodora subsp. variegata extends
south of the Springsure-Maryborough region
in Queensland to near Coffs Harbour in New
South Wales and has foliage that lacks a lemon
scent when crushed. C.citriodora subsp.
citriodora extends north from the Springsure-
Maryborough region in central eastern
Queensland to the Atherton Tableland in north
Queensland and has lemon-scented foliage
when crushed. Further details on the two taxa
(as C. citriodora and C. variegata) are given
in Hill and Johnson (1995).
Within C.citriodora subsp. citriodora ,
McDonald et al. (2000) also noted
morphological differences between northern
populations (Mt Janet south to the White
Mountains region) and southern populations
(Mackay region south to the Springsure-
736
Maryborough region). The northern form
differs in having narrower and more densely
hairy juvenile leaves, narrower adult leaves and
Austrobaileya 5(4): 735-736 (2000)
bark with fewer mottles compared to the
southern form. The taxonomic significance of
these differences requires further study.
Key to taxa in Corymbia ‘section Politaria’
1 Adult leaves broad-lanceolate to lanceolate, to 4.5 cm wide . 2
Adult leaves narrow-lanceolate, to 2.8 cm wide. 3
2 Juvenile leaves to 9.0 cm wide, fruit to 2 x 1.5 cm
(from near Grafton, NSW to the Brisbane region, Qld)
Juvenile leaves to 5.5 cm wide, fruit to 1.4 x 1.1 cm
(fromTaree to Eden inNSW and Mottle Range, Vic.).
3 Leaves lemon-scented (N of Maryborough-Springsure to
Atherton Tableland, Qld). C. citriodora subsp. citriodora
Leaves not lemon-scented (S of Maryborugh-Springsure to
NW of Coffs Harbour, NSW). C. citriodora subsp. variegata
. . C. henryi
C. maculata
Acknowledgements
Ian Brooker and Andrew Slee provided useful
comments on a draft of this paper.
References
Blakely, W.F. (1934). A Key to the Eucalypts. Sydney:
The Worker Trustees
Boland, D. J., Brooker, M.I.H., Chippendale, G.M., Hall,
N., Hyland, B.PM., Johnston, R.D., Kleinig, D.A.
and Turner, J.D. (1984). Forest Trees of
Australia , 4 th . edition. Melbourne: Nelson-
CSIRO.
Brooker, M.I.H. and Kleinig, D.A. (1999). Field Guide
to Eucalypts, South-eastern Australia. Volume
1. Second Edition. Melbourne: Bloomings
Books.
Brooker, M.I.H., Connors, J.R. and Slee, A.V (1997).
EUCLID: Eucalypts of South-eastern Australia.
Centre for Plant Biodiversity Research.
Melbourne: CSIRO.
Chippendale, G.M. (1988). Eucalyptus, Angophora
(Myrtaceae), Flora of Australia 19, Canberra:
Australian Government Publishing Service.
Hill, K.D. and Johnson, L.A.S. (1995). Systematic
studies in the eucalypts. 7. A revision of the
bloodwoods, genus Corymbia (Myrtaceae).
Telopea 6: 185-504.
Larsen, E. (1965). A study of the variability of
Eucalyptus maculata Hook, and E. citriodora
Hook. Australian Forestry and Timber Bureau
Leaflet No. 95, Canberra.
Maiden, J.H. (1920). A Critical Revision of the Genus
Eucalyptus. Volume 5: 90. Sydney:
Government Printer.
McDonald, M.W., Butcher, P.A., Bell, J.C. and
Larmour, J.S. (2000). Intra- and interspecific
allozyme variation in eucalypts from the
spotted gum group, Corymbia section Politaria
(Myrtaceae). Australian Systematic Botany 13:
491-507.
McKern, H.H.G, Spie, M.C. and Willis, J.L. (1954).
The essential oils of Eucalyptus maculata
Hooker. Journal and Proceedings of the Royal
Society of New South Wales 88: 15-21.
MW. McDonald
CSIRO, Forestry and Forest Products, Australian Tree Seed Centre, PO Box E4008, Kingston,
Canberra,ACT, 2604,Australia; email: maurice.mcdonald@ffp.csiro.au
A.RBean
Queensland Herbarium, Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha Road,Toowong,
Queensland, 4066,Australia; email: tony.bean@env.qld.gov.au
Index of new names published in Austrobaileya 5(4)
Apatophyllum flavovirens A.R.Bean & Jessup, sp. nov. . 693
Apatophyllum teretifolium A.R.Bean & Jessup, sp. nov .693
Chamaesyce ophiolitica RI.Forst., sp. nov. .711
Clausena smyrelliana RI.Forst., sp. nov .716
Corymbia citriodora subsp. variegata (F.Muell.)
A.R.Bean & M.W.McDonald, comb, et stat. nov . 735
Eucalyptus provecta A.R.Bean, sp. nov .681
Eucalyptus tardecidens (L.A.S.Johnson & K.D.Hill) A.R.Bean comb, et stat. nov .683
Flindersiapimenteliana forma oppositifolia (F.Muell.) K.D.Scott,
W.K.Harris & J.Playford, comb, et stat. nov .668
Homoranthus cor acinus A.R.Bean, sp. nov. .687
Jasminum domatiigerum subsp. australis W.K.Harris & W. J.McDonald, subsp. nov. .699
Lobelia leucotos Albr., sp. nov . 706
Morinda canthoides (F.Muell.) Halford & R. J.F.Hend., comb, nov .731
Prostanthera clotteniana (F.M.Bailey) A.R.Bean, comb, nov .733
Stylidium sect. Alsinoida(MMbr.) A.R.Bean, comb, et stat. nov .634
Stylidium sect. Biloba A.R.Bean, sect, nov .631
Stylidium sect. Tenella (Benth.) A.R.Bean, comb, et stat. nov. .619
Stylidium sect. Uniflora A.R.Bean, sect, nov .613
Stylidium accedens A.R.Bean, sp. nov .605
Stylidium aquaticum A.R.Bean, sp. nov. .622
Stylidium confertum A.R.Bean, sp. nov. .625
Stylidium divergens A.R.Bean, sp. nov .604
Stylidium ensatum A.R.Bean, sp. nov . 603
Stylidium longissimum A.R.Bean, sp. nov . 629
Stylidium oviflorum A.R.Bean, sp. nov. .624
Stylidium stenophyllum A.R.Bean, sp. nov. . 612
Wahlenbergia celata RI.Forst., sp. nov. .661
737
Referees consulted for Volume 5
Acceptance of papers has depended on the outcome of review by referees. Apart from a few
who did not wish to be listed those consulted during the past four years are listed below. Several
were consulted on more than one occasion. Sincere thanks are extended to all these people
whose expertise has helped to maintain journal standards.
Barker, R.M.
Bean,A.R.
Bergstrom, D.
Bostock, P.
Brooker, M.I.H.
Bruhl, J. J.
Jones, D.L.
Jordaan, M.
Keighery, G J.
Kodela, PG
Clayton, W.D.
Conn, B. J.
Cowie, ID.
Craven, L.A.
Cribb, P. J.
Crisp, M.D.
Lavarack, PS.
Lepschi, B. J.
Lyne,A.M.
Makinson, R.O.
May, T.
Morrison, D.A.
Dawson, J.W.
Dixon, D.
Dunlop, C.R
Duretto, M.F.
Panetta, F.D.
Pedley, L.
Playford, J.
Eggli, U.
Quinn, F.C.
Fensham, R. J.
Forster, PI.
Reynolds, S.T.
Schrire, B.
George, A. S.
Goyder, D. J.
Gross, C.L.
Guymer, GP.
Simon, B.K.
Short, PS.
Snow, N.
Toelken, H.R.
Harden, G J.
Harris,W.K.
Henderson, R. J.F.
Hill, K.D.
Holland,A.E.
Hunter, J.T.
Hyland, B.P.M.
Weiller, C.M.
Welzen, van PC.
Weston, PH.
Wiecek, B.
Wilson, Peter G.
Jacobs, S.W.L.
Jessup, L.W.
738
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 or tropical Australia. All papers are refereed
and the editorial committee reserves the right to reject papers.
Manuscripts must be submitted in duplicate to The Editor, Austrobaileya , Queensland
Herbarium, Meiers Road, Indooroopilly, Qld4068, Australia. They must be double-spaced typewrit¬
ten, with 2.5 cm margins in the first instance. After refereeing, the corrected manuscript should be
submitted on an MSDOS disk as an unformatted ASCII file (DOS Text F ile F ormat), and in your word
processing package format, or as a Rich Text Format file. Manuscripts may be e-mailed to the editor
as an attached file (Laurie.Jessup@env.qld.gov.au). All illustrations should be 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 isl3.5cmxl8.5 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 with the text and also adhered
to the overlay or photocopy of the illustration. 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.
StandardAbbreviations
Titles of journals cited in the nomenclatural sections of 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 the ‘References’ section the titles of both journals and books should be
given in full. All entries in the ‘References’ are to be referred to in the narrative text and all
references to papers or books in narrative text are to be included in the ‘References’ section.
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, N.H. Holmgren and L.C. Barrett (eds), Index
HerbariorumPart 1: The Herbaria of the World. New York: New York Botanical Garden (1990).
Other abbreviations which may be used in citing specimens are S.F. (State Forest), S.F.R. (State
ForestReserve),L.A. (Logging Area), T.R. (Timber Reserve) and an AQ number. This number refers
to 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 5(1-4): 1-735 (1997-2000)
Index
New names are in bold, all others are in italics. Page numbers in bold refer to the principal
description, those in italics refer to an illustration. Pages 369-404 do not exist in Volume 5 due to
a pagination error.
A
Acacia Miller 193, 307, 308, 309, 310, 311, 313, 316
sect. Phyllodineae DC., 307
sect. Juliflorae (Benth.) Maiden & Betche 313, 318
sect. Plurinerves (Benth.) Maiden & Betche 310
abbatiana Pedley 307, 313
arbiana Pedley 307 , 308,
argyrotricha Pedley 307, 310 , 311
armillata 311, 312
barakulensis Pedley 307, 308 , 309
burbidgeae Pedley 308, 309
burdekensis Pedley 307, 313 , 314,318
calyculata A.Cunn. ex Benth. 315, 319, 320
concurrens Pedley 317, 320
conferta A. Cunn. ex Benth. 307, 308
convallium Pedley 307, 312 , 313
crassa Pedley 318, 320
cretata Pedley 307, 318
cunninghamii Hook. 158, 317
faucium Pedley 307, 314 , 315, 318
filipes Pedley 307, 315
fodinalis Pedley 307, 316 , 318
gittinsii Pedley 308, 309
glaucescens var. leiocalyx Domin 317
gnidium Benth. 308
granitica Maiden 313
harpophylla F.Muell. ex Benth. 137, 138, 356, 671,
672
hendersonii Pedley 307, 309
ingramii Tindale 167
johannis Pedley 307, 311 , 312
johnsonii Pedley 309, 310
julifera Benth. subsp. julifera 500
lacertensis Pedley 307, 316 , 317
legnota Pedley 311, 312
leiocalyx (Domin) Pedley 307, 313, 314, 315, 316,
317,318,320
subsp. herveyensis 317
subsp. leiocalyx 317
leptostachya Benth. 313, 314, 319
longispicata Benth. 318, 320
subsp. velutina Pedley 307, 320
ommatosperma (Pedley) Pedley 311, 312
pilligaensis Maiden 309
platycarpa F. Muell. 312, 313
proiantha Pedley 307, 318
resinicostata Pedley 309
rigens 311
rubricola Pedley 307, 309 , 310
scopularum Pedley 307, 319 ,
scopulorum Pedley 587
sericata Cunn. ex Benth. 312, 313
shirleyi Maiden 251, 696
solenota Pedley 307, 319 , 320
sp. (Barakula L.A. Neilsen 15) 308
sp. (Blackdown Tableland R.J. Henderson+ HI 199)
309
sp. (Inglewood A.R.Bean 1115) 310
sp. (Mt Abbot A.R.Bean 4873) 313
sp. (Mt Mulligan J.R.Clarkson 8217) 311
sp. (Norwich Park J.Martin AQ349851) 316
sp. (Ropers Peak P.I.Forster PIF7209) 307
sp. (Torrens Creek C.T.White 8725) 314
sp. aff. A. gonoclada 317
spirorbis subsp. solandri (Benth.) Pedley 319
storyi Tindale 309
tingoorensis Pedley 307, 320
tropica Tindale 316, 317
Acanthocereus (A.Berger) Britton & Rose 671
Acanthus ilicifolius L. 652
Acmena smithii (Poir.) Merrill & Perry 536
Acronychia littoralis T.Hartley & J.B.Williams 719
Acrotriche R.Br. 163
Adelia resinosa Blanco 486
Aegiceras corniculatum (L.) Blanco 146
Aeschynomene arborea L. 223
Agaricus cantharellus Schwein. 554
conicus Schaeff. 539
niveus Scop. 549
pratensis Pers. 545
virgineus Wulfen: Fr. 549
var. niveus (Scop.) Fr. 549
Aglaia sp. 55
Albrecht, David. A new species and lectotypification in
Campanulaceae: Lobelioideae 705
Alchornea rugosa (Lour.) Muell.Arg. 488
Alectryon coriaceus 719
ramiflorus S.T.Reynolds 717, 719
reticulatus Radik. 342
Aleurites J.R.Forst. & GForst. 117
Allium L. 345
Allosyncarpia ternata S.T.Blake 187
Alnus 668
Aloaceae 118
Aloe maculata 113, 118
Alysicarpus Necker ex Desv. 235
sect. Desmodiopsis Schindl. 209, 226
campylocaulos (F. Muell. ex Benth.) Schindl. 226, 233
cheelii C.A.Gardner 209, 222, 260
parviflorus Dalzell 235
Amanoa faginea Baill. 411
ovata auct. non (Decne.) Baill. 407
tomentosa (Blume) Baill. 413
Anatherum Beauv. 503
fulvibarbe (Trin.) Keng 503, 525, 528
muricatum (Retz.) Beauv. 522, 528
zizanioides (L.) Hitchc. & Chase 522, 528
Andropogon L. 503
subgen. Chrysopogon (Trin.) Hack. 506
subsp. echinulatus (Steud.) Hack 526
acicularis Willd. 509
aciculatus Retz. 509, 528
anias Llanos 512, 528
argutus Nees ex Steud. 523, 528
aristulatus Hochst. ex Hack. 518, 528
breviaristatus Steud. 518
echinulatus Nees ex Steud. 526,
elongatus (R. Br.) Spreng. 525,
var.filipes Hack. 513
festucoides Presl 512,
fulvibarbis Trin. 525,
fulvus Spreng. 514
fiiscus Presl 527,
gryllus auct. non L. 518,
gryllus L. 524, 525, 579, 580
gryllus L. subsp. echinulatus (Nees ex Steud.) Cope
526
var. philippinensis Merr. 517,
intermedins 527
javanicus Steud. 510,
javanicus L. sect. Vetiveria (Bory) Thouars ex Benth.
506
javanicus L. subgen. Vetiveria (Bory) Benth. ex Hack.
506
lawsonii Hook.f. 515,
leptanthus Steud. 527,
montanum Roxb. 514
monticola Schult. & Schult. f. 514,
var. genuinus Hack. 514,
var. trinii (Steud.) Hook.f. 520,
muricatus Retz. 522,
var. aristatus Buse 512,
nardus Blanco 522,
nemoralis Balansa 517,
nigritanus Benth. 526,
pallidus 579
serrulatum Link 520
sprengelii Kunth 514,
squarrosus auct. non L.f. 512, 522,
squarrosus L.f. 523
var. chrysopogonoides Hack. 523,
var. genuinus auct. non Hack. 522,
var. nigritanus (Benth.) Hack. 526,
var. nigritanus auct. non Hack. 512,
strictus Host. 527
subtilis Steud. 520,
subulatus Presl 510,
trinii Steud. 520,
var. genuinus Hack. 520,
villosulus Nees ex Steud. 527,
wightianus Nees ex Steud. 518,
zizanioides (L.) Urban 522,
zizanioides auct. non Urban 512,
Angophora costata (Gaertn.) Britten 536
Anthurium Schott. 3
Apatophyllum McGillivray 691, 692, 696
constablei McGillivray 692, 693, 696
flavovirens A.R.Bean & Jessup 691, 692, 693, 695,
696
macgillivrayi R.J.Cranfield & N.S.Lander 696
olsenii McGillivray 692, 693, 696, 697
sp. (Bull Creek A. R. Bean 2225) 693
sp. (Expedition Range E.J. Thompson AQ440723) 693
teretifolium A.R.Bean & Jessup 692, 693, 694, 696
Aphelandra R.Br. 652
sinclairiana Nees 652, 653
Aphyllodium (DC.) Gagnep. 209, 210, 211, 212, 219, 225
australiense (Schindl.) Ohashi 216, 219
biarticulatum (L.) Gagnep. 210,211,212,215, 216,
217, 233
glossocarpum Pedley 209,210,211,213, 214, 215,
217, 219
hispidum (Schindl.) Ohashi 217
Austrobaileya 5(1-4): 1-735 (1997-2000)
latifolium Pedley 209, 211, 212, 213, 214, 215, 216,
217
novoguineense (Schindl.) Ohashi 210, 214,217,219
parvifolium Pedley 209, 211, 212, 213, 217
schindleri Pedley 209, 211, 212, 213, 214, 216, 217,
219
sp. (Lockerbie L. J. Brass 18464) 213
stylosanthoides Pedley 209, 212, 215, 216, 217
Apiaceae 145
Araucaria cunninghamii Aiton ex D. Don 110, 342, 408, 422
Archidendropsis thozetiana (F.Muell.) I.C.Nielsen 101, 422
Argyrodendron F.Muell. 110, 719
Aristolochia elegans Mast. 35
Asclepiadaceae 53, 59
Asteromyrtus Schauer 175
Astrebla F.Muell. ex Benth. 234, 242
Astrotricha DC. 63, 65, 66
subg. Hexocenia Domin 63, 64, 65
biddulphiana F.Muell. 64, 65, 66
brachyandra A.R Bean 63, 65, 66, 67, 69
cordata A.R.Bean 63, 64, 66
floccosa 68
glabra Domin 64, 66
intermedia A.R.Bean 63, 64
latifolia Benth. 64, 68, 69
ledifolia var. glabriflora F.M. Bailey 66
longifolia Benth. 64, 66, 68
var. glabrescens 66
pauciflora A.R.Bean 64, 66
pterocarpa Benth. 63, 64
roddii Makinson 63, 65
umbrosa A.R.Bean 63, 64, 66, 67, 68
Austrobuxus Miq. 9
Austrosteenisia R.Geesink 79, 80
blackii (F.Muell.) R.Geesink 81, 83, 85,87, 89
var. astipella D.J.Dixon 79, 85, 86, 87
var. blackii 79, 84, 85
glabristyla Jessup 79, 80, 81, 87, 89, 90
mollitricha D.J.Dixon 79, 81, 87, 88, 89
stipidaris (C.T.White) Jessup 79, 81, 52,83
B
Babingtonia F.Muell. 157, 158, 170
angusta A.R.Bean 157, 159, 162,163, 165, 171
behrii 159
bidwillii A.RBean 157, 158, 160, 161,167, 168, 171
brachypoda A.RBean 157, 158, 159, 168, 169, 170,
171
collina A.RBean 157, 159, 160, 161, 162,163, 166,
167, 168, 171
crassa A.RBean 157, 159, 160, 166,167, 170, 171
crenulata 158
cunninghamii 158
densifolia (Sm.) F.Muell. 159, 500
granitica 159
jncunda 159
leratii (Schltr.) A.R.Bean 160
odontocalyx 159
papillosa A.R.Bean 157, 158, 159, 169,170, 171
pluriflora (F.Muell.) A.R.Bean 157, 158, 159, 163,
165, 166, 171
procera (J.W.Dawson) A.R.Bean 160
prominens 159
silvestris 159
similis A.R.Bean 157, 159, 161, 162, 163, 165, 166,
168, 171
Austrobaileya 5(1-4): 1-735 (1997-2000)
sp. (Atherton A.R. Bean 5707) 163
sp. (Comet P. Rowland AQ634382) 168
sp. (Townsville A.R. Bean 3424) 169
sp. (Yatala P. Grimshaw+ G525) 162
sp. (Yurol A.R. Bean 6803) 161
squarriilosa 159
tozerensis A.R.Bean 158, 169, 170
virgata (J.R.Forst. & GForst.) F.Muell. 157, 158, 159,
160, 170
virgata var. parvula 157
Backhousia kingii Guymer 35, 338, 422
myrtifolia Hook. f. & Harv. 536
Baeckea L. 157, 170, 499
obtusifolia Brongn. & Gris, 160
parvula (Labill.) DC., 160
parvula var. latifolia Brongn. & Gris 160
sp. “Clarence River” 164, 165
virgata (J.R.Forst. & G.Forst.) Andrews 157, 160, 170
var. /?arv?//aF.M.Bailey 161
var. polyandra Maiden & E.Betche 170
Baloghia inophylla (G Forst.) P.S.Green 14
marmorata C.T. White 342
Banksia L.f. 153, 351
aemula 350
integrifolia L.f. 166
spp. 66
Barklya syringifolia F. Muell. 101, 338, 422
Barleria L. 657
Baumea rubiginosa 349
Bean, A.R. & Jessup L.W. Two new species of Apatophyllum
McGillivaray (Celastraceae) from Queensland 691
Bean, A.R. (2000). Homoranthus coracinus 687
Bean, A.R. A conspectus of Astrotricha DC. (Araliaceae) in
Queensland, including two new species 63
Bean, A.R. A new combination in Prostanthera Labill.
(Lamiaceae) 733
Bean, A.R. A revision of Eucalyptus normantonensis Maiden
& Cambage (Myrtaceae) and its allies 679
Bean, A.R. A revision of Rubus subg. Malachobatus (Focke)
Focke and Rubus subg. Diemenicus A.R.Bean (Rosaceae)
in Australia 39
Bean, A.R. A revision of Stylidium sect. Debilia Mildbr., S.
sect. Floodia Mildbr. and S. sect. Lanata A.R.Bean
(Stylidiaceae) 427
Bean, A.R. A revision of Stylidium subg. Andersonia (R.Br. ex
G.Don) Mildbr. (Stylidiaceae) 589
Bean, A.R. A revision of the Babingtonia virgata (J.R.Forst. &
G.Forst.) F.Muell. complex (Myrtaceae) in Australia 157
Bean, A.R. Homoranthus coracinus (Myrtaceae), a new
species from Queensland 687
Bean, A.R. Lilaeopsis brisbanica (Apiaceae), a new species
from Queensland, Australia 145
Bean, A.R. Microcarpaea agonis (Scrophulariaceae), a new
species from south-eastern Queensland 149
Bean, A.R. Notes on Eucalyptus ser.Psathyroxyla Blakely
(Myrtaceae) andother ‘Ash group’ eucalypts 125
Bean, A.R. Ochrosperma obovatum (Myrtaceae), a new
species from south-eastern Queensland 499
Bean, A.R. Two new species of StylidiumWilld. (Stylidiaceae)
from north Queensland 323
Bergia L. 629
Bertrandia astatogala (Heim) Heim 550
Bertya opponens (F.Muell. ex Benth.) Guymer 36
pedicellata F.Muell. 36, 308
Blechum pyramidatum 652
Boronia Sm. 263, 275, 350
sect. Valvatae (Benth.) Engl. 272, 275, 277, 284,
287,291,295
ser. Valvatae Benth. 275
alulata Sol. ex Benth. 288, 291
amabilis 273
artemesiifolia F.Muell. 292
bella Duretto 275, 282, 283, 284, 285, 287, 288, 295
bowmanii F. Muell. 273, 296, 297
chartacea P.H.Weston 278
duiganiae Duretto 273, 286, 290, 292, 294, 295, 587
eriantha 274
excelsa Duretto 282, 274, 275, 283, 284, 285, 287,
288, 295
foetida Duretto 275, 282, 283, 284, 285, 286,
287, 288, 295
forsteri Duretto 274, 276, 278, 279, 280, 281, 282
glabra (Maiden & Betche) Cheel 274, 278, 282, 295
granitica 274
hoipolloi Duretto 273, 278, 288, 289, 290, 291, 292
jensziae Duretto 275, 282, 283, 284, 285, 287, 288,
295
keysii Domin 273, 274, 277, 295
lanceolata F.Muell. 274, 293, 294, 295
lanuginosa White 291,292
ledifolia (Vent.) DC. 273, 275, 277, 295
var. rosmarinifolia (A. Cunn. ex Endl.) Benth. 275
obovata 273, 294
odorata Duretto 273, 274, 277, 278, 281, 284, 285,
287, 288, 290, 292, 293, 294, 295, 296
palasepala Duretto 276, 278, 279, 280,257
quinkanensis Duretto 286, 288, 290, 291, 292
repanda 274
rosmarinifolia A. Cunn. ex Endl. 263, 274, 275, 276,
211, 278, 279, 280, 281, 282, 284, 285, 287
var. albiflora Cheel 277
ruppii Cheel 295
sp. (Hinchinbrook Island S.L. Everist 7786) 282
sp. (Many Peaks Range I.R. Telford CBG7702560)
287
sp. (Massy Creek R.G. Coveny+ 7174) 295
sp. (Mt Mulligan J.R. Clarkson 1) 291
sp. (Mt Walsh PI. Forster+ PIF17253) 286
sp. (Mt Windsor Tableland P.I. Forsterf PIF15225)
285
sp. (Robinson Gorge P.I. ForsterL PIF 11235) 281
splendida Duretto 274, 276, 278, 279, 281
squamipetala Duretto 273, 286, 295, 296, 297
ternata Endl. 295
viridiflora Duretto 291
Bothriochloa bladhii (Retz.) S.T. Blake 527,
Bridelia Willd. 405, 406, 411, 414
erapensis S.Dressier 409, 410
exalt at a F.Muell. 405, 407,406,407,408,413, 414,
418
faginea (Baill.) F.Muell. ex Benth. 406, 411
finalis P.I.Forst. 405, 407, 408, 409,410, 414, 419
glabrifolia Merr. 413, 415
insulana Hance 405, 406, 407, 410, 411, 414, 419
lancifolia Roxb. 413
leichhardtii Baill. ex Muell.Arg. 405, 406, 407, 408,
411, 412, 414, 419
var. glabrata Domin 411
melanthesoides var. australiensis Gehrm. 411, 412
minutiflora Hook.f. 410, 411
ovata Decne. 406
ovata auct. non Decne. 413
var. exaltata (F.Muell.) Muell.Arg. 407
penangiana Hook.f. 406, 410, 411
phyllanthoides W.Fitz. 406, 413
scandens (Roxb.) Willd. 406
sp. 408
tomentosa Blume 405, 406, 407, 413,474, 415, 416,
418
var. eriantha Airy Shaw 406, 413
var. glabrata Domin 406
var. glabrescens Benth. 413
var. glabrifolia (Merr.) Airy Shaw 406, 413
var. lancifolia Muell.Arg. 413
var. ovoidea Benth. 413
var. tomentosa 406
var. trichadenia Muell.Arg. 406, 413
Briedelia 406
Bryaceae351
Bryophyllum Salisb. 114, 115
Brynm billardieri Schwaegr. 351
Bursaria reevesii L.Cayzer & M.D.Crisp 714
C
Cactoblastis cactorum 672
Caesalpinia L. 97, 101
brachycarpa (Benth.) Hattink 101
erythrocarpa Pedley 98 , 99, 100
hymenocarpa 98
nitens (F.Muell. ex Benth.) Pedley 97, 98, 99
scortechinii (F.Muell.) Hattink 98, 99, 101
subtropica Pedley 98, 101
traceyi Pedley 97, 98, 99 , 100
Caleana minor R. Br. 75
Callicoma serratifolia Andrews 163
Callistemon sieberi DC. 167
sp. 308
Calycacanthus K.Schum. 651, 652, 653, 656, 658
magnusianus K.Schum. 652, 653
Calytrix mimiana Craven 615
Camarophyllopsis Herink 535, 537, 539,542,562, 563
subgen. Hodophilus (R. Heim) Bon 562
kearneyi A.M.Young 535, 538, 542, 545, 562, 563
phaeophylla (Romag.) Arnolds 563
schulzeri (Bres.) Herink. 542
Camarophyllus (Fr.) Kummer 539
apricosa sensu Fuhrer & Robinson 546
aurantiopallens Horak 545
lilacinus (Cleland & Cheel) Horak 535, 547
niveus (Scop.) Wunsche 549
pratensis (Pers.: Fr.) Kummer. 545
virgineus (Wulfen: Fr.) Kummer 549
Camphoromyrtus pluriflora F.Muell. 157, 165
Campylopus introflexus (Hedw.) Brid. 350
Canaca Guillaumin 9
Candollea alsinoides 634
anisata (Willd.) Hook.f. ex Benth. 715
capillaris (R.Br.) F.Muell. 620
diffusa (R.Br.) F.Muell. 627
eriorhiza (R.Br.) F.Muell. 323, 451
fissiloba (F.Muell.) F.Muell. 597, 622, 623, 624, 625
floodii (F.Muell.) F.Muell. 433
floribunda (R.Br.) F.Muell. 444
lobuliflora (F.Muell.) F.Muell. 608
muscicola (F.Muell.) F.Muell. 602
pachyrrhiza (F.Muell.) F.Muell. 610, 612
pedunculata (R.Br.) F.Muell. 618
schizantha (F.Muell.) F.Muell. 609
tenerrima 636
trichopoda (F.Muell.) F.Muell. 613, 615
uliginosa (Sw. ex Willd.) F.Muell. 599
Austrobaileya 5(1-4): 1-735 (1997-2000)
Canscora diffusa 629
Cansora drummondii 711, 712
Cantharellaceae 548
Cantharellus 548
lilacinus Cleland & Cheel 547
Canthium Lam. 353, 358, 360
microphyllum F.Muell. 357, 358
sp. (Massy Creek P.I. Forster+ PIF10568) 358
vacciniifolium F.Muell. 353, 354, 355, 357, 358
Capillipedium Stapf 503
leucotrichum (A. Camus) Schmid ex Veldk. 527,
parviflorum (R. Br.) Stapf. 527,
Capparis arborea (F.Muell.) Maiden 107, 110, 111
‘laurifolia ’ 108
nobilis var. pubescens Benth. 107, 108, 110
velutina P.I.Forst. 107, 109, 111
Casearia sp. 55
Cassytha L. 345, 346, 347
glabella R.Br. 345, 346
melantha R.Br. 345, 346
panicidata R.Br. 345
pubescens R.Br. 345, 347
Castanospermum australe A.Cunn. & Fraser ex Hook. 26
Casuarina L. 179, 351
cristata 671, 672
Centrophorum Trin. 506
Centrostemma Decne. 57
multiflorum Blume 57
Ceratopetalum apetalum D. Don 536, 563
Cereus Miller 671
hildmannianus 671, 672
jamacaru 674, 675
penmanus Mill. 671,672, 676
uruguayanus Ritt. ex Kiesl 671, 672, 674,675, 676
Chalcoelytrum Lunell 506
Chamaeraphis muricata (Retz.) Merr. 522,
squarrosa (Retz.) Merr. 522
Chamaesyce S.F.Gray. 711
drummondii (Boiss.)D.C.Hassall 711,712
ophiolitica PLForst. 711, 712, 713, 714
petala (Ewart & L.R.Kerr) P.I.Forst. & RJ.F.Henderson
711, 712
Choricarpia subargentea (C.T.White)L.A.S.Johnson 338, 422
Choriceras Baillon 9
tricorne Benth. 26
Chrysopogon Trin. 503,505, 506, 508, 516, 525, 526, 579
sect. Rhaphis (Lour.) Roberty 506
sect. Vetiveria (Bory) Roberty 506
acicularis Duthie 509
aciculatus (Retz.) Trin. 505, 508, 509, 526,
var. longifolius Buse 510
argutus (Steud.) Trin. ex B.D. Jackson 504, 523
benthamianus Henr. 524, 525, 579
bomeensis Henr. 505, 508, 510, 511
calcaratus (Hack.) Henr. 524
celebicus Veldk. 504, 508, 511
chinense Trin. 506
coll inns Ridl. 518, 519
echinulatus (Nees ex Steud.) Wats. 524,526
elongatus (R. Br.) Benth. 504, 525
elongatus auct. non Benth. 516
var . filipes Benth. 513
fa/lax S.T. Blake 504, 525, 524, 579
festucoides (Presl) Veldk. 505, 507, 508, 512, 513,
523,526
filipes (Benth.) Reeder 504, 506, 509, 513, 514, 515,
517
Austrobaileya 5(1-4): 1-735 (1997-2000)
var. arundinaceus Reeder 513
fulvibarbis (Trin.) Veldk. 504, 525
fidvus (Spreng.) Chiov. 505, 507, 514, 515, 519, 520
subvar. montanus (Trin.) Roberty 514
subvar. serrulatus (Trin.) Roberty 520
fulvns auct. non Chiov. 520
fuscus (Presl) Trin. ex Steud. 527
glabratus Trin. 524
gryllus (L.) Trin. 505, 506, 522, 524,525, 579, 580
grylliis auct. non Trin. 517, 521, 524
subsp. echinulatus (Nees ex Steud) Cope 504, 526
subsp. pallidas (R.Br.) Domin 580
subvar. pilosus Domin 580
var. pallidas (R.Br.) Benth. 524
intercedens Veldk. 504, 505, 509, 515
javaniciim 510
latifolius S.T. Blake 504
lawsonii (Hook.f.) Veldk. 504, 505, 507, 511, 512, 515
leucotrichus A. Camus 527
micrantherus Veldk 504, 509, 516, 517, 525
montanus Trin. 514
var. serrulatus (Trin.) Stapf 520
monticola (Schult. & J.H. Schult.) Haines 514
nemoralis (Balansa) Holttum 504,507, 508, 509, 511,
512, 517, 518, 525
nigritanus (Benth.) Veldk. 513, 526,
nodulibarbis (Steud.) Henr. 504
oliganthus Veldk. 504, 505, 513, 514, 515, 516, 526,
orientalis (Desv.) A. Camus 506, 507, 508, 514, 515,
518, 519, 520, 521,
pallidus (R.Br.) Trin. ex Steud. 524, 580
pauciflorus Vasey 526
perlaxus Bor 507, 519,
philippinensis (Merr.) Henr. 517, 518,
rigidus (B.K. Simon) Veldk. 504, 505, 527,
serridatus Trin. 507, 508, 514, 515,519, 520, 522,
settfolius Stapf 505
sinensis Rendle 518,
squarrosa (L.f.) Merr. 522
strictus (Nees) Jackson 527,
subtilis (Steud.) Miq. 505, 509, 520,521, 522,
subulatus (Presl) Trin. ex Steud. 510,
sylvaticus C.E. Hubb. 504, 505, 513, 514
tadulingamii Sreekumar 505
tenuiculmis Henr. 505, 509, 511, 521, 522, 525,
trinii (Steud.) Watson 520,
trivialis (Lour.) Walker-Amott & Nees 509
verticillatus var. orientalis (Desv.) 518
villosulus (Steud.) Watson 527
wightianus (Nees ex Steud.) Thw. 518
zeylanicus (Steud.) Thw. 504
zizanioides (L.) Roberty 504, 506, 507, 508, 512, 518,
522, 523, 526,
var. fulvibarbis (Trin.) Roberty 525
var. nigritanus (Benth.) Roberty 526
var. nigritanus auct. non Roberty 512
Clausena Burm.f. 715, 716
anisata (Willd.) Hook.f. ex Benth 715
anisum-olens 716
brevistyla Oliver 715, 716, 717
var. brevistyla 715
var. papuana (Lauterb.) J.F.Molino 715
excavata Burm.f 715
harmandiana (Pierre) Pierre ex Guillaumin 715
heptaphylla (Roxb.) Wight & Arn. 715, 719
kanpurensis J.F.Molino 715
lansium (Lour.) Skeels. 715, 716
poilanei J.F.Molino 715
sanki (Perrottet) J.F.Molino 716
smyrelliana PI.Forst. 715, 716,717 ,718, 719
Cleistanthus Hook.f. ex Planch. 405
cunninghamii 717
peninsularis Airy Shaw 410
Clifford, H. Trevor, The seedling of Cassytha glabella R.Br.
345
Coccoceras Miq. 457
Codariocalyx Hassk. 210, 235
motorius (Houtt.) Ohashi 226
Codiaeum variegatum 26
Commelina L. 629
Conospermum sphacelatum Hook. 663
Cooktownia D.L. Jones 74
robertsii D.L. Jones 71, 74, 75, 76, 77
Coprosma J.R.Forst. & G.Forst.731
acutifolia F.Muell. ex Benth. 731
canthoides F.Muell. 731
Cordemoya Baill. 457
Corymbia K.D.Hill & L.A.S.Johnson735
‘section Politaria’ 736
citriodora (Hook.) K.D.Hill & L.A.S.Johnson 662,
735
subsp. citriodorai F. MuelI.)A. R. Bean &
M.W.McDonald 735, 736
confertiflora 527
subsp. variegata 735, 736
bunites (Brooker & A.R.Bean) K.D.Hill &
L.A.S. Johnson 309
hendersonii 696
henryi (S.T.Blake) K.D.Hill & L.A.S.Johnson 735, 736
maculata (Hook.) K.D.Hill & L.A.S.Johnson 735, 736
spp 332
trachyphloia (F.Muell.) K.D.Hill &L.A.S.Johnson 308
variegata (F.Muell.) K.D.Hill & L.A.S.Johnson 735
watsoniana (F.Muell.) K.D.Hill & L.A.S.Johnson 662
xanthope 712
Crantzia lineata (Michx.) Nutt. 145
Crassula L. 115
Crassulaceae 114
Croton mollissimus Geisel. 473, 475
paniculatus Lam. 478
philippense Lam. 480
repandus Willd. 484
ricinoides Pers. 457, 473, 475
Cryptanthus Otto & A. Dietr. 3
Cryptocarya foetida R.T. Baker 342
Cryptocentrum Benth. 4
Cryptolepis 729
lancifolia PI.Forst. 729
papillata PI.Forst. 729
perakensis (Gamble)P.I.Forst. 729
Cuphophyllus 540, 548, 553
Cyclophyllum Hook.f. 353, 354, 355
Cyperus dijformis L. 150
D
Dacrydium Sol. ex G.Forst. 181
Dalibarda repens L. 39
Daniel, Thomas F. Chromosome numbers of some Acanthaceae
from Papua New Guinea 651
Darwinia Rudge 687
Dendrocnide Miq. 121, 123
corallodesme 123
cordata (Warb. ex H.J.Winkl.) Chew 121, 122, 123
cordifolia (L.S.Sm.) B.R.Jackes & M.Hurley 121,122,
123
excelsa 123
moroides 121, 122, 123
photinophvlla (Kunth) Chew 121, 123
Dendrolobium Benth. 209, 210, 219,225, 256, 257
arbuscula (Domin) Ohashi 220,222, 224, 225
cheelii (C.A. Gardner) Pedley 209, 220, 221, 222, 223
hayatae Ohashi 221
multiflorum Pedley 209, 220, 221, 224, 225
polyneurum (S.T. Blake)Pedley 209, 219, 220, 221,222
polyneurum (S.T.Blake) Ohashi 587
quinquepetalum (Blanco) Merr. 224, 225
stipatum S.T. Blake 219, 220, 221, 222, 223
triangulare (Retz.) Schindl. 222, 224
umbellatum (L.) Benth. 219,220,221,222,223,224,225
f. hirsutum (DC.) Ohashi 209, 223, 224
var. hirsutum (DC.) Pedley 223
var. umbellatuni 223
Derris Lour. 79
trifoliata var. macrocarpa Domin 79
Desmodiopsis Desv. 209, 210, 217, 225, 226,234, 235, 246,
259
campylocaulon 234
muelleri 234
Desmodium 226
subsp. acutifoliolum Verde. 250
sect Desmodiopsis (Schindl.) Pedley 209, 226, 227
sect. Dollinera (Endl.) Schindl. 221
sect Heteroloma Benth. 227, 233
sect. Nicolsonia (DC.) Benth. 227, 246
sect Oxytes (Schindl.) Ohashi 227
sect Sagotia (Duchass. & Walpers.)Benth. 227, 246
sect Stenostachys Schindl. 209, 226
ser. Arillata Pedley 209, 226, 227
ser. Sagotia (Duchass. & Walpers) Pedley 226, 227
ser. Stenostachys (Schindl.) Pedley 209, 226, 227,
260
subg. Acanthocladum Pedley 225, 227
subg. Hanslia (Schindl.) Ohashi 226
subg. Oxytes 234
subg. Phyllodium (Desv.) Baker 256
subg. Sagotia (Duchass. & Walpers)Baker 227,246
acanthocladum F. Muell. 209, 226, 227, 228, 231, 233
ancistrocarpum (Ledebour) DC. 256
archboldianum E.G. Baker 247
biarticulatum (L.) F. Muell. 216, 233
biarticulatum var. australiense (Schindl.) Meeuwen
216
var. caudatum 218
brachypodum A. Gray 227, 228, 233, 234
brownii Schindl. 227, 229, 238, 239, 240
campylocaulon F. Muell. ex Benth. 226, 227, 228, 233,
234, 235
canum Schinz & Thellung 255
clavitricha 237
dependens Blume exMiq. 231
deplanchei Harms 234
dietrichiae Domin 252
filiforme Zollinger & Moritzi 209, 226, 227, 229, 236,
237, 238, 239, 240, 243, 244
flagellare Benth. 227, 229, 240, 241, 243
frutescens Schindl. 255
gangeticum (L.) DC. 227, 230, 249
glareosum Pedley 209, 227, 229, 238, 242, 243
gunnii Benth. ex J.D. Hook. 227, 231, 251, 252, 253
hannii Schindl. 227, 229, 238, 241, 242, 243
Austrobaileya 5(1-4): 1-735 (1997-2000)
heterocarpon (L.) DC. 227, 230, 246, 247, 248, 256
heterocarpon (L.) DC. var. heterocarpon 246
var. strigosum Meeuwen 246
heterophyllum (Willd.) DC. 227, 229, 235, 236
incanum DC., Prodr. 226, 227, 230, 255
indigotinum Harms. & K. Schum. 233
intortum (Mill.) Urban 226, 227, 228, 255
limense Hook. 255
macrocarpum Domin 227, 231, 250, 251
megaphyllum 257
microphyllum (Thunb. ex Murray) DC. 209, 226, 227,
228, 235
muelleri Benth. 227, 229, 238, 239, 241, 242, 243
var. minus 246
nemorosum F. Muell. ex Benth. 227, 230, 247, 248
subvar. eboracense Schindl. 247
subvar. whitfordii Schindl. 209, 248
var. nemorosum 248
var. novoguineense Kaneh. & Hatus. 247
var. simplex Schindl. 247
var. whitfordii (Schindl.) Ohashi 248
neurocarpum Benth. 238, 240
var. gracile Benth. 240
var. queenslandicum Domin 238
novae-hollandiae Domin 216
f. latifolium Domin 216
var. caudatum Domin 214
var. novae-hollandiae 218
ormocarpoides DC., 227, 231
ospriostreblum Chiov. 253
parvifolium 235
pilosiusculum DC. 255
polycarpon (Poir.) DC. 246
polyneurum S.T.Blake 209
procumbens (Mill.) Hitchc. 253
pryonii DC. 249
pulchellum (L.) Benth. 257
pullenii Pedley 209, 227, 229, 237, 238, 239, 243, 244
purpureum (Miller) Fawcett & Rendle 253
pycnostachyum Benth. 234
pycnotrichum Pedley 209, 227, 229, 236, 237, 238,
239, 242
quinquepetalum 225
rhytidophyllum F.Muell. ex Benth. 227, 231, 234, 249,
250, 251
sandwicense 255
scorpiurus (Sw.) Desv. 225, 227, 228, 233
spartioides DC., Prodr. 252
strigillosum Schindl. 227, 230, 247
tenax Schindl. 227,231,250
tiwiense Pedley 209, 227, 230, 248
tortuosum (Sw.) DC., 226, 227, 228, 253
trichocaulon DC. 246
trichostachyum Benth. 227, 229, 236, 237, 238, 239,
242
triflorum (L.) DC. 209, 226, 227, 229, 235
umbellatum var. hirsutum DC. 209, 223
uncinatum (Jacq.) DC. 226, 227, 228, 255
unifoliolatum (Merr.) Steen. 233
var. caudatum Domin 218
varians (Labill.) G Don 231, 234, 250, 253
var. angustifolium Benth. 252
var. gunnii (Benth. ex J.D. Hook.) Benth. 251
velutinum (Willd.) DC. 227, 230, 249
var. velutinum 249
whitfordii (Schindl.) Pedley 209, 247
Desmondiastrum A. Pramanik & Thoth. 234
Austrobaileya 5(1-4): 1-735 (1997-2000)
Deuteromallotus Pax & K.Hoffm. 457
Dicerma DC. 209,210, 211, 217,219
sect. Aphyllodium 211
sect. Phyllodium (Desv.) Benth. 256
acanthocladum F.Muell. 231
biarticulatum (L.) DC. 211,216,217
f. australiense Schindl. 219
f. longibracteatum Schindl. 213 ,219
f. plumosum Schindl, 214,219
subsp. australiense Schindl. 214, 216
subsp. biarticulatum 217
var. australiense Schindl. 214, 216
var. biarticulatum 217
var. collettii Schindl. 218
hispidum 217
novoguineense 217 ,219
pulchellum (L.) DC. 257
Dichanthium Willemet 242, 243
Dicliptera Juss. 656
Dicranaceae 350
Diectomis fastigiata 521
Dillwynia J.E.Smith 93
trifoliata Roth 93
Diospyros fasciculosa 717
Diplachne P.Beauv. 137, 299
parviflora (R. Br.) Benth. 300
reptatrix (L.) Druce 300
Dipteracanthus Nees 657
Dischidia R.Br. 53, 725, 726
bengalensis 726
imbricata 726
parvifolia 726
sub ul at a 726
superba Rintz 725
torricellensis (Schltr.) P.I.Forst. 725, 726, 727, 728
Dissiliaria F.Muell. 9, 10
baloghioides F.Muell. ex Baill. 9, 11, 12, 13, 14
indistincta P.I.Forst. 9, 10, 11, 12, 17, 18, 19,
laxinervis Airy Shaw 9, 10, 11, 14, 16, 17, 19, 21
muelleri Baill. 9, 10, 11, 21, 22, 21, 25, 26
surculosa P.I.Forst. 9, 10, 11, 22, 24, 25, 26
tricornis Benth. 26
tuckeri P.I.Forst. 9, 10, 11, 19, 20, 21
Ditrichaceae 350
Ditrichum difficile (Dub.) Fleisch. 350, 351
Dixon, D.J. A taxonomic revision of the genus Austrosteenisia
R.Geesink (Fabaceae: Millettieae) 79
Dodonaea filifolia Hook. 308
spp. 500
Dollinera Endl. 221
Droogmansia De Wild. 259
Drosera L. 601, 619
Duretto, Marco F. Boronia sect. Valvatae (Benth.) Engl.
(Rutaceae) in Queensland, Australia 263
E
Ecbolium Kurz 653
Echidnopsis malum (Lavranos) Bruyns 55
Echinocroton F.Muell. 458
claoxyloides F.Muell. 457, 458, 460
Echinolobium biarticulatum 216
Echinopsis Zucc. 671
multiplex (Pfeiff.) Zucc. 671
Echinus Lour. 458
claoxyloides (F.Muell.) Baill. 460
var. cordata Baill. 457, 460
var. cordatus 463
var. ficifolia Baill. 457, 468
mollissimus (Geisel.) Baill. 473
nesophilus (Muell.Arg.) Baill. 475, 476
philippensis (Lam.) Baill. 480
trisulcus Lour. 458
Elytraria imbricata (Vahl) Pers. 654
Empodisma L.A.S.Johnson & Cutler 349
Epilobium tonkinense H.Lev. 628
Eranthemum L. 657
Eremophila debilis (Andrews) Chinnock 706
Eriocaulon L. 601, 629
Eriorhizum 324
Eucalyptus L’Herit. 125, 175,200
ser. Buxeales Blakely 679
ser. Psathyroxyla Blakely 125, 126, 130
subser. Considenianae Brooker & Slee, M 126
subser. Strictinae L.D.Pryor & L.A.S.Johnson 125,
134
acmenoides Schauer 68
andrewsii Maiden 125, 126, 127, 128, 130, 134
subsp. andrewsii 125
subsp. campanulata (R.T.Baker & H.G.S.) 125, 128
apothalassica L.A.S.Johnson & K.D.Hill 662
approximans Maiden 125, 134
subsp. codonocarpa Blakely & McKie 134
bicolor 680
var. xanthophylla Blakely 680
camaldulensis 625
campanulata R.T.Baker & H.G.Sm. 125, 126,128, 129,
130
chloroclada (Blakely) L.A.S.Johnson & K.D.Hill 150
chlorophylla Brooker & Done 679
clarksoniana 443
cloeziana F.Muell. 65, 68
codonocarpa Blakely & McKie 125, 134
consideniana Maiden 126, 127
crebra F. Muell 251, 316
cullenii 625
dealbata Cunn. ex Schauer 65
decorticans 696
fibrosa F.Muell. 662, 712
grandis W.Hill ex Maiden 44
gummifera (Gaertn.) Hochr. 166
haemastoma 126, 133
var. capitata Maiden 132
var. inophloia C.T.White 130
var. micrantha (DC.) Benth. 132
var. sclerophylla Blakely 132
intermedia R.T.Baker 161
largeana Blakely 679
largiflorens F.Muell. 679
var. xanthophylla (Blakely) Cameron 680
melanophloia F.Muell. 65
mensalis L.A.S. Johnson & K.D. Hill 309
micrantha DC. 132
var. signata (F.Muell.) Blakely 132
microcodon L.A.S.Johnson & K.D.Hill 125, 134
microcorys F.Muell. 48
microtheca F.Muell. 679
miniata A. Cunn. ex Schauer 193, 216
montivaga A. R. Bean 125, 126, 130, 131, 132
multicaulis Blakely, 126, 127
nesophila Blakely 328
normantonensis Maiden & Cambage 679, 680, 681,
682, 685
olida L.A.S.Johnson & K.D.Hill 125, 126, 129, 130
persistens L.A.S.Johnson & K.D.Hill 680, 682, 683,
685
subsp. tardecidens L.A.S.Johnson 683
phoenicea F. Muell. 193, 328
pilularis Sm. 68, 536
polycarpa F.Muell. 437
provecta A.R.Bean 679, 680,681, 682, 685
ptychocarpa F.Muell. 437
racemosa 125, 125, 126, 132, 133
racemosa var. signata (F.Muell.) R.D.Johnst. &
Marryatt 132
remota Blakely 126, 127
resinifera Sm. 161
rossii R.T.Baker & H.GSm., 126, 133, 134
rummeryi Maiden 679
saligna 48
sclerophylla (Blakely)L.A.S.Johnson & Blaxell 132,
133
sieberi L.A.S.Johnson 126, 127, 166
signata F.Muell. 132, 133
sphaerocarpa L.A.S. Johnson & Blaxell 309
spp. 66, 68, 332, 609
stockeri D.J.Carr & S.GM.Carr 328
suffulgens L.A.S.Johnson & K.D.Hill 65
tardecidens (L.A.S.Johnson & K.D.Hill) A.R.Bean.
679, 680, 682, 683, 685
tectifica F.Muell. 679
tenuipes (Maiden & Blakely)Blakely & C.T.White 500,
663
tetrodonta F. Muell. 193, 216, 328, 443
trachyphloia F.Muell. 65, 688
variegata F.Muell. 735
watsoniana F.Muell. subsp. watsoniana 500
lucasii Blakely 679
Euonymus globularis Ding Hou 691
Euphorbia L. 711
Euphorbiaceae 9, 29, 115
Eupomatia lanrina 586
Euroschinus falcata Hook.f. 715, 717
Everistia S.T.Reynolds & RJ.F.Hend. 353,354
vacciniifolia (F.Muell.) S.T.Reynolds & RJ.F.Hend.
353, 354, 355, 356, 358
f. crassa S.T.Reynolds & RJ.F.Hend. 353, 356,357,
358
f. vacciniifolia 356
var. nervosa S.T.Reynolds & RJ.F.Hend. 353, 356,
357, 358, 359
var. vacciniifolia 356, 358, 359
Excoecaria agallocha L. 146
Fasciculochloa B.K. Simon & C.M. Weiller 583
sparshottiorum B.K. Simon & C.M.Weiller 583
Fenzlia Benth. 173,174
bennigseniana (Volk.) Burret 181
microphylla (Benth.) Domin 194
obtnsa Endl. 174, 182, 196
obtusa var. microphylla Benth. 174, 194
phebalioides W. Fitzg., 198
retusa Endl., 174, 198
Ficus albipila (Miq.) King, 24
Finlaysonia Wallich. 729
Flindersia R.Br. 667, 668
oppositifolia (F.Muell.) Hartley & Jessup 667, 668
pimenteliana F.Muell. 667, 668, 669
forma oppositifolia (F.Muell.)K. D. Scott, W.K.Harris
& J.Playford 668
Austrobaileya 5(1-4): 1-735 (1997-2000)
unifoliata Hartley 668
Fontainea Heckel 29, 37
australis Jessup & Guymer 37
borealis P.I.Forst. 29, 30, 31
fugax P.I.Forst. 29, 34, 35, 36, 37
oraria Jessup & Guymer 37
pancheri (Baillon) Heckel 29
picrosperma C.T.White 29, 30, 32, 34, 37
rostrata Jessup & Guymer 34, 35, 37
subpapuana P.I.Forst. 29, 32, 33, 34
venosa Jessup & Guymer 37
Forster, PI. Cryptolepis papillata P.I.Forst. & Sarcolobus
porcatus P.I.Forst. (Asclepiadaceae), newly recorded from
West Papua 729
Forster, Paul I. & Schmeider, Miriam. Cereus uruguayanus
(Cactaceae) and its naturalised occurrence in Queensland,
Australia 671
Forster, Paul I. A taxonomic revision of Bridelia Willd.
(Euphorbiaceae) in Australia 405
Forster, Paul I. A taxonomic revision of Dissiliaria F.Muell.
ex. Baill. (Euphorbiaceae) 9
Forster, Paul I. A taxonomic revision of Mallotus Lour.
(Euphorbiaceae) in Australia 457
Forster, Paul I. Capparis ve/«r/'na(Capparaceae), a new species
from south-eastern Queensland 107
Forster, Paul I. Chamaesyce ophiolitica.( Euphorbiaceae), a
new and endangered species endemic to serpentine
vegetation in central Queensland 711
Forster, Paul I. Clausena smyrelliana (Rutaceae:
Aurantioideae), a new and critically endangered species
from south-east Queensland 715
Forster, Paul I. Marsdenia jensenii (Asclepiadaceae:
Marsdenieae ), a new species from north-eastern
Queensland 59
Forster, Paul I. Notes on the naturalised Flora of Queensland, 3
113
Forster, Paul I. Peperomia hunteriana (Piperaceae), a new
species from the ‘Wet Tropics’ of north-eastern Queensland
573
Forster, Paul I. Rediscovery of Dischidia torricellensis
(Schltr.) P.I.Forst., an unusual epiphytic asclepiad from
New Guinea. 725
Forster, Paul I. Three new species of Fontainea Heckel
(Euphorbiaceae) from Australia and Papua New Guinea 29
Forster, Paul I. Typification and application of the name
Macrozamia macleayi Miq. (Zamiaceae) 577
Forster, Paul I. Wahlengergia celata (Campanulaceae), a new
species from central Queensland 661
Forster, Paul I. Liddle David J. and Liddle, Iris M. Madangia
inflata (Asclepiadaceae: Marsdenieae), a new genus and
species from Papua New Guinea 53
Forstera bellidifolia Hook. 591
Fraxinus L. 668
Frullania rostrata (J.D.Hook. & Tayl.) J.D.Hook. & Tayl. 351
Frullaniaceae 351
Genoplesium apostasioides 75
Gentiana L. 668
Gliophorus chromolimoneus (G Stevenson) Horak 554
graminicolor Horak 556
pallidus Horak 535, 556, 557, 558
viridis (G Stevenson) Horak 560
Glochidion ferdinandi (Mull. Arg.) F. M. Bailey 536
Glossostigma diandrum (L.) Kuntze 150
Gramen aciculatum Rumph 510,
Graptopetalum Rose 114, 115
Austrobaileya 5(1-4): 1-735 (1997-2000)
paraguayense (N.E.Br.) Walther 113, 114
Graptophyllum Nees 651, 653, 654, 655
pictum (L.) Griff. 55, 651, 652, 653, 654, 655
Grevillea longistyla 696
pteridifolia 637
Grewia breviflora Benth. 491
Guioa Cav. 103, 104
crenifoliola Merr. & Perry 103, 104
pteropoda Radik. 103, 104
sarcopterifructa Welzen 104
Gunnessia pepo P.I.Forst. 55
Gymnanthera R.Br. 729
Gymnoschoenus sphaerocephalus (R.Br.) Hook.f. 50
Gymnosporia (Wight & Am.) Hook.f. 691, 692
emarginata 692
inermis Merr. & Perry 692
senegalensis 692
H
Habenaria Willd. 73, 75, 331, 334, 335
divaricata R.S.Rogers & C.T.White 335
elongata R.Br. 335
harroldii D.L.Jones 335
hymenophylla Schltr. 335
macraithii Lavarack 335
ochroleuca R.Br. 331, 332, 334
praecox Lavarack & Dockrill 331, 332, 333, 334, 335
propinquior Rchb.f. 331, 334, 335
rumphii (Brongn.) Lindl. 335
triplonema Schltr. 335
xanthantha F.Muell 334, 335
Halford, D.A. Oldenlandia gibsonii (Rubiaceae:
Hedyotideae), a new species from south east Queensland
337
Halford, David A. & Henderson, R. J.F. A new combination in
Morinda L. (Rubiaceae) for Australia 731
Hanslia Schindl. 231
adhaerens (Poir.) Schindl. 231
Haplostichanthus sp. 26
Harmogia 157
virgata (J.R.Forst. & G.Forst.) Schauer 160
Harris, Wayne K. & McDonald, William J. Jasmimim
domatiigerum subsp. australis (Oleaceae), a new
subspecies from North Queensland 699
Harris, Wayne K. and Holmes Glenn, Jasmimim jenniae
(Oleaceae), new species from south eastern Queensland 341
Harrisia Britton 671
Hedraianthera porphyropetala F.Muell. 691
Hedysarum L. 210, 249
adhaerens Poir. 231
ancistrocarpum Ledebour 256
biarticulatum L. 211, 216, 217 , 219
canum Gmel. 255
diphyllum 247
gangeticum L. 249
heterocarpon L. 246
heterophyllum Willd. 236
incanum Sw. 255
intortum Mill, 255
lutescens Poir. 211
microphyllum Thunb. ex Murray 235
polycarpon Poir. 246
pulchellum L. 211, 247,258
purpureum Miller 253
scorpiurus Swartz 233
tortuosum Sw. 253
triflorum L. 236
tuberculosum Labill. 246
umbellatum 223, 247
uncinatum Jacq. 255
various Labill. 252
Hemigenia clotteniana F.M.Bailey 733
Henderson, Rod. In defence of Chrysopogon fallax S.T. Blake
(Poaceae) 579
Hibbertia exutiacies N.A.Wakef. 500
Hodophilus foetens (Phill.) R. Heim 562
Holcus elongatus R. Br. 525,
gryllus (L.) R. Br. 524, 579, 580
gryllus auct. non R.Br. 524,
pallidus 579
zizanioides (L.) Kuntze ex Stuckert 522,
Holland, A.E. A new species and a new combination in
Rutidosis (Gnaphalieae:Angianthinae: Asteraceae) 565
Holland, A.E. Rothia indica subsp. australis A.E.Holland
(Fabaceae: Crotalarieae), a new subspecies occuring in
Australia. 93
Holographis Nees 652
Holtzea Schindl. 219
umbellata Schindl. 219,222
Homoranthus A.Cunn. ex Schauer 687
coracinus A.R.Bean 687, 688, 689
darwinioides 687, 689
porteri 689
zeteticorum Craven & S.RJones 688
Hoya R.Br. 53,55, 57
heuschkeliana Kloppenburg 55
multiflora Blume 57
telosmoides Omlor 55
Humidicutis 540
Hydrocybe 554
Hygrocybe (Fr.) Rummer 535, 537, 539, 540, 557, 562
subgen. Cuphophyllus Donk 545
subgen. Humidicutis 542
subgen. Humidicutis Singer 560
subgen. Hygrocybe 541, 549
subgen. Pseudohygrocybe 541
subgen. Pseudohygrocybe M. Bon. 550
anomala A.M.Young 541, 543, 544, 545, 551, 552, 553
var. anomala 551
var. ianthinomarginata, A. M. Young 535, 551, 552
apricosa Horak 546
astatogala (Heim) Heinemann 541, 544, 550
aurantiopallens (Horak) A. M. Young 541, 544, 545,
546
aurantipes A. M. Young 538, 541, 544,553
austropratensis A. M. Young 535, 541, 544, 545, 546,
547
batesii A. M. Young 535, 538, 556, 557, 558
cantharellus (Schwein.) Murrill 541, 544,554
cheelii A. M. Young 535, 541, 543, 547
chromolimonea (G. Stevenson) May & Wood 542,
543, 554
coccinea{ Schaeff.: Fr.) Rummer 550
conica (Schaeff.: Fr.) Rummer 549
constrictospora Arnolds 562
erythrocala A. M. Young 542, 543, 544, 555
graminicolor (Horak) May & Wood 535, 538, 542,
543, 556, 557, 558,560
iropus Young 538
kula Grgurinovic 542, 544, 558
lanecovensis A. M. Young 535, 542, 543, 544, 558, 559
lewellinae (Ralchbrenner) A.M.Young 542, 543, 560
lilacina (C. Laest. ex P. Rarst.) M.Moser 547
mavis (G. Stevenson) Horak 561
miniata (Fr.: Fr.) Kummer 538, 558
nivea 549
pallida (A. H. Smith) Singer 556
pallida (Horak) A. M. Young 556
pratensis (Pers.: Fr.) Murrill 547
psittacina sensu Shepherd and Totterdell 560
reesiae 541, 543, 548
rodwayi (Massee) A. M. Young 549
rosella Horak 561
sp. LC1 541, 544, 561, 562
stevensoniae May & Wood 542, 543, 560
virginea (Wulfen: Fr.)Orton & Watling 538, 540, 545,
549
viridis Capelari & Maziero 560
Hygrocybeae{ Schaeff.:Fr.) Kuhner 535, 539
Hygrophoraceae 535, 537, 538, 539, 564
Hygrophoreae P. Henn. 539
Hygrophorus Fr. 539
aurantius Murrill sensu G Stevenson 545
cantharelhis (Schwein.) Fr. 554
chromolimonens G. Stevenson 554
conica Fr. 550
involutus G. Stevenson 549
marginatus Peck 560
niveus (Scop.) Fr. 549
psittacinus sensu Cleland and Cheel 560
virgineus (Wulfen: Fr.) Fr. 549
viridis G Stevenson 560
viridis sensu Young 556
Hylocereus 671
Hypoestes Sol. exR.Br. 651, 655, 658
aristata 656
floribunda R.Br. 652, 653, 655
var. neoguineensis R.M. Barker 655
Hypsela tridens E.Wimm. 707
Hypsophila oppositifolia F.Muell. 668
I
Iseilema spp. 234
J
Jackes, B.R. and Hurley, M. Anew combination in
Dendrocnide (Urticaceae) in north Queensland 121
Jadunia Lindau 651, 656
biroi (Lindau & K.Schum.) Lindau 652,653, 656
Jasminum L. 117, 341, 699
sect. Trifoliolata 700
aemulum R.Br. 342
var. brassii PS.Green 341, 342
calcareum F.Muell. 341, 344, 699
dallachii 344, 700
didymum 344, 699, 700, 701
subsp. didymum 701
subsp. lineare (R.Br.) P.S.Green 341, 699
subsp. racemosum 701
domatiigerum Lingelsh. subsp. australis W.K.Harris
& W.J.McDonald. 699,700
subsp. domatiigerum 700
gilgianum 700
jenniae W.K. Harris & G Holmes 341, 342, 343, 344
mesneyi Hance 113, 117
simplicifolium 344
sp. 342
Jatropha L. 115, 118
curcas L. 115
gossypifolia L. 115
Austrobaileya 5(1-4): 1-735 (1997-2000)
podagrica Hook. 113, 115
Jones, David L. Cooktownia robertsii, a remarkable new
genus and species of Orchidaceae from Australia 7
Justicia L. 653
K
Kalanchoe Adans.114, 115, 118
lateritia 113, 114
Kania Schlechter 174, 182
hirsutula (F.Muell.) A. J. Scott 181
Kudirra-pullu Rheede 510,
Kunstleria Prain 79
blackii (F.Muell.) Polhill 83
stipularis (C.T.White) Polhill 80, 81
Kunzea ambigua (Sm.) Druce 166
ericoides (A.Rich.) Joy Thomps. 166, 170
compacta (Steph.) Grolle 351
L
Lankesteria Lindl 657
Lantana camara L. 423
Laportea cor difolia L.S.Sm., 121, 122
Lavarack, Peter S. and Dockrill, Alick W.A. new species of
Habenaria Willd. (Orchidaceae) from North Queensland
331
Lejeunea flava (Sw.) Nees subsp. orientalis Schust. 351
Lejeuneaceae 351
Lepidagathis Willd. 651, 656, 657
royenii Bremek. 652, 653, 656
formosensis C.B.Clarke ex Hayata 656
Lepidoziaceae 351
Lepironia articulata 349, 350
Leptochloa PBeauv. 5, 137, 141, 299, 300, 301
ciliolata (Jedw.) S.T.Blake 4, 302
decipiens (R.Br.) Stapf ex Maiden 4
digitata 301
divaricatissima 138, 302
dubia (Nees) Kunth 3, 301
fusca (L.) Kunth 299
subsp. fusca 300, 302
subsp. muelleri (Benth.) N. Snow 300, 301
subsp. uninervia 299, 300, 301, 302, 305
gigantea (Launert) T. A. Cope & N. Snow 300
ligulata Lazarides 2, 137, 138, 301, 303
mucronata 302
neesii 301
panicea subsp. brachiata 302
subsp. panicea 301, 303
peacockii (Maiden & Betche) Domin 4, 138, 141
southwoodii N. Snow & B.K.Simon 138, 139, 140,
141, 142, 301, 302
Leptospermum 349
brachyandrum (F.Muell.) Druce 170
lamellatum Joy Thomps. 688
parvulum Labill. 160
petersonii F.M.Bailey 167
sericatum Lindl. 500, 663
virgatum J.R.Forst. & G. Forst. 157, 160
Leucobryum candidum (PBeauv.) Wils. 350
Leucopogon flexifolius R.Br. 688
Lilaeopsis Greene 145, 146
australica 145
brisbanica A.R.Bean 145, 146, 147
chinensis (L.) Kuntze 145
fistulosa A.W.Hill 145, 146
novae-zealandiae (Gand.) A.W.Hill 145, 146, 148
polyantha (Gand.) H.Eichler 145, 146
Austrobaileya 5(1-4): 1-735 (1997-2000)
Lindernia 629
Lithomyrtus F.Muell. 173, 174, 175, 178, 182, 183, 185
cordata (A.J. Scott) N.Snow & Guymer 180, 183, 184,
185, 193
densifolia N.Snow & Guymer 185, 186, 189, 188, 192,
193
dunlopii N.Snow & Guymer 184, 186, 188, 189, 194
grandifolia N.Snow & Guymer 183, 188, 189
hypoleuca N.Snow & Guymer 174, 182, 183, 184, 190,
191, 192, 198
kakaduensis N.Snow & Guymer 183, 193, 183, 193,
192, 190
linariifolia N.Snow & Guymer 174, 183, 189, 190,
192, 193
microphylla (Benth.) N.Snow & Guymer 184, 192,
194, 195, 196
obtusa (Endl.) N.Snow & Guymer 174, 183, 184, 185,
189, 194, 195, 196
repens N.Snow & Guymer 183, 195, 197, 198
retusa (Endl.) N.Snow & Guymer 183, 184, 185, 193,
197, 198, 199
Lobelia darlingensis (E.Wimm.) Albr. 709
leucotos Albr. 705, 706, 707, 709
quadrangularis R.Br. 707, 709
stenophylla Benth. 706, 707
Lobelioideae 705
Logania cordifolia Hook. 663
Lonchocarpus blackii (F.Muell.) Benth. 83
nesiotes 79, 83
stipularis C.T.White 79, 81
Longetia 9
Lophocereus schottii (Englem.) Britt. & Rose 676
Lophostemon confertus (R.Br.) Peter G.Wilson &
J.T. Waterhouse 342
suaveolens (Gaertn.) Peter GWilson & J.T. Waterhouse
161
Lysicarpus angustifolius (Hook.) Druce 663
Mackaya 653
Macromitrium aurescens Hainpe 351
Macrozamia 577
johnsonii D.L. Jones & K.D. Hill 363
johnsonii F.Muell 363
Incida L.A.S.Johnson 577
macleayi Miq. 577
miquelii 577
moorei D.L.Jones and K.D.Hill 363
Madangia P.I.Forst., D.J.Liddle & I.M.Liddle 53, 55
inflata P.I.Forst., D.J.Liddle & I.M.Liddle 53, 54, 55,
56, 57
Mallotus Lour. 457,458
angustifolius Benth. 457, 470
claoxyloides (F.Muell.) Muell.Arg. 457, 460, 461 , 462,
463, 470, 472, 493
f. grossedentata Domin 468
var. angustifolia F.M.Bailey 457
var. cordatus (Baill.) Airy 460, 463
var .ficifolius (Baill.) Benth. 468, 470
var. glabratus Domin 460
var. macrophylla Benth. 463, 468
cochinchinensis Lour. 458, 478
derbyensis W.Fitzg. 491
didymochryseus Airy Shaw 457, 467
didymochryseus auct. non Airy Shaw 465
discolor F.Muell. ex Benth. 457, 458, 460, 463, 464,
465, 493
dispersus P.I.Forst. 457, 458, 460, 465, 466, 467, 495
ficifolius (Baill.) Pax & K.Hoffm. 460, 463, 468, 469,
470, 494
floribundus (Blume) Muell.Arg. 489
megadontus P.I.Forst. 458, 460, 470, 471, 472, 495
mollissimus (Geisel.) Airy Shaw 459, 473, 474, 475,
494
muricatus var. walkerae (Hook.f) Pax & K.Hoffm. 486
nesophilus Muell.Arg. 458, 460, 465, 475, 476, 477,
478, 491,495
oblongifolius (Miq.) Muell.Arg. 491
paniculatus (Lam.) Muell.Arg. 460, 478, 479, 480,
491, 496
philippensis (Lam.) Muell.Arg. 458, 460,
480, 481,496
polyadenos F.Muell. 460, 481, 482, 483, 491, 493
pycnostachys F.Muell. 473, 475
repandus (Willd.) Muell.Arg. 459, 484, 485, 497
resinosus (Blanco) Merr. 458, 460, 486, 487, 488, 497
ricinoides (Pers.) Muell.Arg. 473, 475
sp. 470, 488
surculosus P.I.Forst. 459, 488, 489, 490, 495
tiliifolius (Blume) Muell.Arg. 476, 480, 489, 491
walkerae Hook.f. 486
zippelii (Hassk.) F.Muell. 457
Mammillaria 671
Manihot Mill. 115, 116
esculenta Crantz 113, 115, 116
flabellifolia 116
glaziovii Muell.Arg. 113, 116, 117
grahamii Hook. 113, 115, 116
Maniltoa psilogyne Harms 55
Marsdenia 59, 726
Marsdenia cymulosa 726
hemiptera H.Rch. 59, 61
jensenii P.I.Forst. 59, 60, 61
Maytenus Molina 691, 692
cunninghamii 692
disperma 692
emarginata 692
fasciculiflora Jessup 692
silvestris 692
McDonald, Bill reviewer. Australian Tropical Rainforest Trees
and Shrubs by B.P.M.Hyland, T. Whiffin, B.Gray,
R.W. Elick, &A.J. Ford 585
McDonald, M.W. & Bean, A.R. A new combination in
Corymbia section “ Politaria C. citriodora subsp.
variegata (Myrtaceae) 735
Mecopus Benn. 210
Meibomia acanthoclada (F. Muell.) Kuntze 231
campylocaulon (F. Muell. ex Benth.) Kuntze 234
filiformis (Zoll. & Moritzi) Kuntze 238
flagellaris (Benth.) Kuntze 240
gangetica (L.) Kuntze 249
heterocarpa (L.) Kuntze 246
heterophylla (Willd.) Kuntze 236
intorta (Mill.) Blake 255
microphylla (Thunb.) Kuntze 235
muelleri (Benth.) Kuntze 242
nemorosa (F. Muell. ex Benth.) Kuntze 247
neurocarpa (Benth.) Kuntze 238
pulchella (L) . Kuntze 258
tortuosa (Sw.) Kuntze 253
trichostachya (Benth.) Kuntze 237
triflora (L.) Kuntze 236
uncinata (Jacq.) Kuntze 255
varians (Labill.) Kuntze 252
velutina Willd. 249
Melaleuca L. 157, 163, 175,214,216,238,351, 601, 610
citrolens Barlow 328
leucadendra 450, 633, 635
nervosa 443
quinquinervia 349, 635
spp. 437, 592,617, 637
thymifolia Sm. 500
virgata (J.R.Forst. & GForst.) L.f. 160
viridiflora 431,434,443
viridiflora Sol. ex Gaertn. 328, 332, 603, 609, 619,
620, 622, 625, 627, 629
Mezoneuron 97
brachycarpum 101
Micholitzia 726
Micraira subulifolia 625
Microcarpaea R.Br. 149
agonis A.R.Bean 149, 150, 151
minima 15, 149
muscosa 149
Microloma 726
Micromelum minutum 715, 716, 717
Micromyrtus leptocalyx 696
Millettia 83
blackii F.Muell. 83
Mitrasacme oasena Dunlop 663
Morinda L. 731
acutifolia F.Muell. 731
canthoides (F.Muell.) Halford & R.J.F.Hend. 731
Myristica bicuhyba 345
Myrtaceae 125, 157, 173, 207, 499
Myrtastrum Burret 175
Myrtella F.Muell. 173, 174, 175, 177, 178,183,202, 206, 207
sect. Eufenzlia Burret. 174
beccarii F.Muell. 178,179, 180
bennigseniana (Volk.) Diels 179, 180, 181
cordata A.J. Scott 174, 184, 185
hirsutula F.Muell. 174, 182
microphylla (Benth.) A. J. Scott 194
obtusa (Endl.) A. J. Scott, 196
phebalioides (W. Fitzg.) A.J. Scott, 198
retusa ( Endl.) A. J. Scott 198
rostrata Lauterb., 181
N
Neomyrtus Burret 175
Neotrewia Pax & K.Hoffm. 457
Nothofagus cunninghamii (Hook, f.) Oerst. 538
Nyctocereus 671
O
Ochrosperma Trudgen 499, 500
adpressum 499, 500
citriodorum 499, 500
lineare 499, 500
obovatum A.R.Bean 499, 500, 501
oligomerum 499, 500
sulcatum 499, 500
Octospermum Airy Shaw 457
Odontonema 653
Oldenlandia L.337
gibsonii Halford 337, 338, 339
polyclada (F.Muell.) F.Muell. 337, 338
sp. (Wietalaba N.Gibson 1344) 337
Ole a paniculata R.Br. 342
Oleaceae 117, 341
Oplonia 653
Austrobaileya 5(1-4): 1-735 (1997-2000)
Opuntia Miller 671
stricta 672
tomentosa 671
Orthotrichaceae 351
Ougeinia Benth. 210
P
Paederota minima J.Konig ex Retz. 149
Pandanus spiralis R.Br. 437
spp. 609, 637
Panicnm hians 583
Parasorghum (Snowden) Garber 505
Pedley, Les Desmodium Desv. (Fabaceae) and related genera
in Australia: a taxonomic revision 209
Pedley, Les Notes on Acacia (Leguminosae: Mimosoideae)
chiefly from northern Australia 307
Pedley, Les Notes on Caesalpinia subg. Mezoneuron
(Leguminosae: Caesalpinioideae) in Australia 97
Peperomia Ruiz. & Pav. 573, 574
bellendenkerensis Domin 573, 574
blanda (Jacq.) Knuth 573, 574
enervis C.DC. et F.Muell. 573, 574
hunteriana PI.Forst. 573, 574, 575
leptostachya Hook. & Arn. 573
pellucida 574
tetraphylla 574
Peplidium 149
Pereskia 671
Peristrophe 656
Peristylus 73, 75, 335
banfieldii (F.M. Bailey) Lavarack 335
candidus J.J.Sm. 335
tradescantifolius (Rchb.f.) Kores 335
Persicaria 118
capitata (Buch.-Ham. ex D.Don) H.Gross 113, 118
Petalostigma banksii Britten & S.Moore 328
Phalaris zizanioides L. 522,
Phebalium glandulosum Hook. 308
Phyllodium 209, 210, 211, 220, 225, 256
hackeri Pedley 209,256, 257
insigne 257
lutescens 211
pulchellum (L.) Desv. 211, 256, 257, 259
var. pulchellum 257
var. glabrius Pedley 209, 257, 258, 587
var. pulchellum 258
sp. (Montalbion H.S.McKee 9430) 256
sp. A 258
Pilidiostigma recurvum (C.T.White) A.J.Scott 26
Pittosporum sp. 55
undulatum Vent. 536
Platanthera 73
papuana Schltr. 74, 75
Poaceae 503, 579
Polemoniaceae 174
Pollinia Spreng. 506
fulva Spreng. 514
gryllus (L.) Spreng. 525,
pallida 579
Polyalthia sp. 55
Polygonaceae 118
Pomaderris clivicola E.M.Ross 36
Pongamia pinnata (L.) Pierre 26
Popowia sp. 55
Prainea hackeri 256
insigne (Prain) Schindl. 256
Austrobaileya 5(1-4): 1-735 (1997-2000)
Pratia pedunculata (R.Br.) Benth.705, 706
puberula Benth 705, 706, 707, 709
purpurascens (R.Br.) E.Wimm. 706, 707,709
Premna lignum-vitae (A. Cunn. ex Schauer) W. Piep. 342
Prostanthera Labill. 733
atroviolacea F.M.Bailey 733
clotteniana (F.M.Bailey) A.R.Bean 733
lithospermoides 733
Psammomoya Diels & Loes. 691
Pseudarthria Wight & Am. 210
Pseuderanthemum 653, 654
Psendohygrocybe 540, 553
Pseudoraphis spinescens (R. Br.) Vickery 523
Psidium L. 174
Psydrax Gaertn. 353, 354, 355
lamprophylla (F.Muell.) Bridson 353
Pterolobium 101
nitens F.Muell. ex Benth. 97, 99, 101
Pultenaea 349
Pycnospora 211,235
Pyrostria Comm, ex Juss. 353
R
Racosperma 312
Razisea 653
Reynolds, S.T. and Henderson, R.J.F. Vanguerieae A. Rich, ex
Dum. (Rubiaceae) in Australia, 1. Everistia S.T.Reynolds &
R.J.F.Hend. 353
Rhaphis Lour. 503, 506
acicularis (Retz.) Desv. 509
aciculatus (Retz.) Honda 509
argnta Nees 523
echimdata Nees 526,
elongatus (R. Br.) Chase 525,
elongatus auct. non Chase 516,
var. filipes (Benth.) Keng 503, 513,
javanica Nees 510,
orientalis Desv. 518,
stricta Nees 527,
trivialis Lour. 506, 509,
zizanioides (L.) Roberty 522,
subvar. fulvibarbis (Trin.) Roberty 525,
var. aciculata (Retz.) Roberty 509,
var. nigritana (Benth.) Roberty 526,
Rhinacanthus Nees 653
Rhodamnia Jack 421, 423
angustifolia N.Snow & Guymer 421, 422, 423, 424,
425, 426
costata A.J. Scott 423
dumicola Guymer & Jessup 342, 423
glandulosa 568, 572
maideniana C.T.White 423
whiteana Guymer & Jessup 423
Rhynchosia 241
Rhynchospora pterochaeta F.Muell. 706
Riccia cartilaginosa Steph. 351
Ricciaceae 351
Rinorea horneri (Korth.) O.Kuntze 55
Rockinghamia Airy Shaw 457
Rotala ilecebroides 629
mexicana Cham. & Schltdl. 150, 151
occultiflora Koehne 150, 151
Rothia Pers. 93
hirsuta (Guill. & Perr.) Baker 96
indica (L.) Druce 93, 96
indica subsp. australis A.E. Holland 93, 94, 95
subsp. indica 94
lotoides 93
trifoliata (Roth) Pers. 93
Rottlera oblongifolia Miq. 491
Rottlera Roxb. 458
tiliifolia Blume 489, 491
tinctoria Roxb. 458
Rubiaceae 353
Rubus L. 39, 50
subg. Cylactis (Focke) Focke 39
subg. Dalibarda (L.) Focke 39, 50
subg. Diemenicus A.R.Bean 39, 48
subg. Idaeobatus 48
subg. Malachobatus (Focke)Focke 39,41, 48
subg. Rubus 39
alceifolius Poir. 39, 40, 41, 42
bellobatus 41
capricorni Focke 44, 45
chamaemorus L 50
dalibarda L. 39
dendrocharis (Focke) Focke 42
ellipticus 40
fockeanus Kurz 39
fruticosus 39, 41
gunnianus 39, 40, 49, 50
haskarlii 42
hillii F.Muell. 39, 44, 47
laciniatus 41
lasiococcus Gray 39
moluccanus L. 39, 42, 44, 45
var. alceifolius (Poir.) Kuntze 41
var. austropacificus P.Royen 44
var. moluccanus 39, 40, 44
var. trilobus A.R.Bean 39, 40, 45, 46,47, 48
moorei 40
nebulosus 40
parvifolius L. 41, 48
pedatus Sm. 39
probus 41
queenslandicus 41
rosifolius var. commersonii 41
var. rosifolius 41
rugosus Sm. 39
ursinus 41
x illegitimus Focke 47, 48
x novae-cambriae Gand. 47
x novus Kuntze 39, 41, 47, 48, 50
Ruellia L. 651, 657, 658
macrantha 657
prostrata 657
repens L. 652, 653, 656, 657
tuberosa L. 652, 653, 657
Rutaceae 263
Rutidosis 565, 571
Rutidosis acutiglumis Philipson 567, 571
auricoma F.Muell. 567
crispata 572
glandulosa A.E.Holland 565, 566, 567
helichrysoides DC. 568,567, 570, 571
helichrysoides subsp. acutiglumis (W.R.Philipson)
A.E.Holland 565, 568, 569, 570, 571
subsp. helichrysoides 565, 568, 569, 570
heterogama 572
lanata 572
leiolepis 572
leptorhynchoides 572
leucantha 572
murchisonii 565, 567, 568, 572
sp.(Blackdown Tableland K.A.Williams79082)
565
S
Saffordiella bennigseniana (Volk.) Merrill 178, 181
Sagotia 226, 234
triflora (L.) Duchass. & Walpers 226, 236
Sankowskya stipularis P.I.Forst. 9, 26
Sarcolobus 729
porcatus P.I.Forst. 729
retusns K.Schum.729
secamonoides (Schltr.) P.I.Forst. 729
vittatus P.I.Forst 729
Sarcopteryx 103, 104
Schoenus 619
Scott, Kirsten D., Harris, Wayne K., Playford, Julia. The
identity of Flindersia pimenteliana and F. oppositifolia
(Rutaceae): Evidence from DNA Sequences 667
Scrophulariaceae 149
Sedum 114, 115
praealtum 113, 114
subsp. praealtum 114
Selenicereus 671
Sharma, I.K., Jones, D.L. & Forster, PI. Contribution of
isozymic analysis in differentiating Macrozamia moorei
D.L.Jones and K.D.Hill from M.johnsonii F.Muell.
(Zamiaceae) 363
Simon, Bryan. Steinchisma hians (Elliott) Nash, the correct
name for Fasciculochloa sparshottiorum B.K.Simon &
C.M.Weiller 583
Siphonoglossa Oerst. 653
ramosa Oerst. 654
sessilis (Jacq.) Oerst. 654
Snow, Neil and Guymer Gordon P. Rhodamnia angustifolia
(Myrtaceae), a new and endangered species from south¬
eastern Queensland 421
Snow, Neil and Guymer, Gordon P. Systematic and cladistic
studies of Myrtella F.Muell. and Lithomyrtus F.Muell.
(Myrtaceae) 173
Snow, Neil and Simon, Bryan K. Leptochloa southwoodii
(Poaceae: Chloridoideae), a new species from south-east
Queensland 137
Snow, Neil and Simon, Bryan K. Taxonomic status and
Australian distribution of the weedy neotropical grass
Leptochloa fusca subsp. uninervia, with an updated key to
Australian Leptochloa (Poaceae, Chloridoideae) 299
Snow, Neil. Application of the phylogenetic species concept:
A botanical monographic perspective. 1
Sorghum Pers. 505
elongatum (R. Br.) Beauv. 525,
fuscum (Presl) Miq. 527,
nitidum (Vahl) Pers. 527,
zizanioides (L.) Kuntze 522,
Spathidolepis 725, 726
torricellensis Schltr. 725, 726
Sphagnaceae 350
Sphagnum australe Mitt. 349, 350, 351
Spokes, T. reviewer. Wildflowers of Southern Western
Australia by Margaret G Corrick & Bruce A. Fuhrer 153
Stackhousia tryonii 714
Stapeliopsis neronis Pillans 55
Steinchisma 583
hians (Elliott) Nash 583
Stenandrium Nees 652
Sterculia quadrifida R.Br. 422, 717
Austrobaileya 5(1-4): 1-735 (1997-2000)
Streblacanthus 653
Strychnos psilosperma F.Muell. 342
Stylidiaceae 323, 589
Stylidium Willd. 323, 427, 428, 589
sect. Alsinoida (Mildbr.) A.R.Bean 598, 634
sect. Andersonia R.Br ex G.Don 589, 592, 597, 598
sect. Biloba A.R.Bean 598, 631
sect. Debiles Mildbr. 323
sect. Debilia Mildbr. 427, 428, 440
sect. Floodia Mildbr. 427, 428, 430, 433
sect. Lanata A.R.Bean (Stylidiaceae) 427, 428, 451
sect. Nitrangium (Endl.) Sonder 589
sect. Sparsifolia Benth. 427
sect. Stylidium 427
sect. Tenella (Benth.) A.R.Bean 591, 598, 619
sect. Tolypangium 427
sect. Uniflora A.R.Bean 598, 613
subg. Alsinoida Mildbr. 589, 591, 634
subg. Alsinoides 428
subg. Andersonia (R.Br. ex G.Don) Mildbr. 428,
589, 595, 597
subg. Centridium 428, 595
subg. Tolypangium (Endl.) Mildbr. 427, 428, 595
accedens A.R.Bean 589, 597, 604, 605, 607
adenophorum Lowrie & Kenneally 428, 429, 439,
440, 454
alsinoides R.Br. 589, 634
var. cordifolium Ewart 638
androsaceum 617
aquaticum A.R.Bean 589, 597, 622, 623, 626
austrocapense A.R.Bean 427, 430, 442, 443, 449, 453
barrettorum 428,441, 442
brunonis 598, 599
bryoides F.Muell. 618, 619
candelabrum Lowrie & Kenneally 596, 606, 608, 614
capillare R.Br. 596, 608, 620
clarksonii Lowrie & Kenneally 429, 431,432, 453
claytonioides W.Fitzg. 595, 619
confertum A.R.Bean 589, 597, 625, 630
cordifolium W.Fitzg. 589, 597, 638, 639
costulatum Kenneally & Lowrie 428, 429, 437, 438,
453
curtum Carlquist 618, 619
debile F.Muell. 427,430, 446,447, 448, 450, 451, 455
var. paniculatum Maiden & Betche 446
delicatum A.R.Bean 427, 429, 434, 435, 436, 454
desertorum Carlquist 427, 429, 430, 438, 439, 453
diffusum R.Br. 595, 597, 627 , 629, 630
divergens A.R.Bean 589, 596, 597, 604, 605, 607
dunlopianum Carlquist 589, 595, 632, 633
eglandulosum F.Muell. 323, 427
ensatum A.R.Bean 589, 596,603, 607
ericksoniae J.H.Willis 615, 595, 618, 619
eriorhizum R.Br. 323,324,225, 326, 328, 330, 427,
430, 451
evolutum Carlquist 636
fimbriatum Lowrie & Kenneally 595, 634
fissilobum 622, 623, 624
floodii F.Muell. 427, 429, 433,434, 435, 437, 439,
440, 453
floribundum R.Br. 430, 444, 445, 446, 455
Austrobaileya 5(1-4): 1-735 (1997-2000)
fluminense F.L.Erickson & J.H.Willis 589, 597, 635,
636
foveolatum A.R.Bean 427, 429, 430,431, 432, 436, 453
graminifolium Sw. 427
humphreysii Carlquist 323
inaequipetalum J.M.Black 430, 445, 446, 454
inconspicuum Slooten 597, 629, 631
irriguum W.Fitzg. 631, 632
javanicum Slooten 597, 637, 638
kunthii Wall, ex DC. 591, 595, 596, 598, 599
laricifolium A. Rich. 427
leiophyllum A.R.Bean 323, 325, 326, 327, 328, 330,
430, 451
leptorrhizum auct. non F.Muell. 440
leptorrhizum F.Muell. 428, 430, 441, 442, 443, 444,
453
var. pilosum Benth. 440
lobuliflorum F.Muell. 608, 596, 609, 612
longissimum A.R Bean 589, 597, 629, 639
mitrasacmoides Carlquist 619, 637
mitrasacmoides F.Muell. 636
mucronatum 429, 433, 454
multiscapum O.Schwarz 430,442,443, 444, 454
muscicola F.Muell. 596, 602, 603
nomination Carlquist 596, 619, 620, 621
omatnm S.T.Blake 430, 450, 451, 455
oviflorum A.R.Bean 589, 597, 624
pachyrrhizum F.Muell. 596, 610, 611, 612, 613, 615
paniculatum (Maiden & Betche) A.R.Bean 427, 430,
446, 447, 449, 450, 455
pedunculatum R.Br. 595, 615, 617, 618, 619
var. ericksoniae (J.H.Willis.) Carlquist 617
perizostera Lowrie & Kenneally 591, 595, 618
prophyllum Lowrie & Kenneally 589, 591, 597, 619,
621, 622
pseudotenellum O.Schwarz 623
qnadrifurcatum F.L.Erickson & J.H.Willis 620
ramosissimum A.R Bean 323, 325, 328, 329, 330,429,
451
reductum Carlquist 631
rivulosnm Lowrie & Kenneally 429, 433, 435, 453
roseum Kurz 628, 629
rotundifolium 595,631, 632
rubriscapum W.Fitzg. 429, 432, 433, 455
schizanthum F.Muell. 590, 596, 609, 610
semipartitum F.Muell. 430,440, 441,452, 454
simulans 596, 603, 604
sinicum Hance 599, 600
sp. 434
stenophyllum A.RBean 589, 596, 612, 613, 614, 615
symonii Carlquist 429, 434, 453
tenellum R.Br. 600
tenellnm Sw. ex Willd. 591, 595, 597, 600, 619, 627,
629
var. minimum C.B.Clarke in Hook 628
tenerrimum F.Muell. 597, 598, 636, 637
tenerum Spreng. 589, 595, 596, 597, 600, 601
trichopodum F.Muell. 595, 613, 616, 617
turbinatum Lowrie & Kenneally 429, 435, 437, 438,
453
uliginosum Sw. ex Willd. 589, 599, 600, 606
velleioides A.R.Bean 427, 430, 448, 449, 450, 454
Stylosanthes 216
Syncarpia 284, 285
glomidifera (Sm.) Nied. 161, 536
T
Tabernaemontana aurantiaca Gaud. 55
pandacaqui Lam. 55
Tadehagi Ohashi 210, 225, 258
robustum Pedley 258
triquetrum (L.) Ohashi. 259, 260
triquetrum subsp. triquetrum 259
Tapioca 116
Teliostachya 656
Thunbergia Retz. 657
arnhemica F.Muell. 657
grandiflora Roxb. 652, 653, 657
papuana Bremek. 657
Trewia L. 457
Trifidocanthus unifoliolatus Merr. 233
Triglochin striata Ruiz & Pav. 146
Trigonella indica 93
Triodia mitchellii 696
scariosa N.T.Burb. 500
sp. 609
U
Umbelliferae 145
Uromyrtus Burret. 173, 181
rostrata (Lauterb.) N.Snow & Guymer 181
Urticaceae 121
Utricularia 601, 619
V
van Welzen, Peter C. Guioa sarcopterifructa (Sapindaceae): a
new Australian species 103
VanguerieaeA. Rich. exDum. 353, 354
Veldkamp, J.F. A revision of Chrysopogon Trin. including
Vetiveria Bory (Poaceae) in Thailand and Malesia with
notes on some other species from Africa and Australia
503
Velleia spathidata 450
Verbesina encelioides (Cav.) Benth. & Hook.f. ex A. Gray 35
Vernicia Lour. 117
fordii (Hemsl.) Airy Shaw, 113, 117
Verticordia oculata 153
ovalifolia 153
Vetiveria Bory 503, 504, 505, 506, 516, 525
arguta (Steud.) C.E. Hubb. 523,
elongata (R. Br.) Stapf ex C.E. Hubb. 503, 505, 525,
elongata auct. non C.E. Hubb. 516,
festucoides (Presl) Ohwi 505, 512,
filipes (Benth.) C.E. Hubb. 503, 505, 513,
filipes auct. non C.E. Hubb. 516,
var. arundinacea (Reeder) Jansen 513,
fulvibarbis (Trin.) Stapf 525,
intermedia S.T. Blake 503
lawsonii (Hook.f.) Blatter & McCann 513, 515
lawsonii auct. non Blatter & McCann 512, 517
muricata (Retz.) Griseb. 522
nemoralis (Balansa) A. Camus 517, 523
nemoralis auct. non A. Camus 522
nigritana (Benth.) Stapf 505, 526
nigritana auct. non Stapf. 512
odorata auct. non Virey. 522
odoratissima auct. non Bory. 522
odoratissima Bory 506
Austrobaileya 5(1-4): 1-735 (1997-2000)
pauciflora S.T. Blake 503, 504, 526
rigida B.K. Simon 527
zizanioides (L.) Nash 504, 505, 522
var. chrysopogonoides (Hack.) A. Camus 523
var. genuine/ A. Camus 513, 522
var. nigritana (Benth.) A. Camus 526
var. tonkinensis A. Camus 522
W
Wahlenbergia 661, 662
celata 661, 662, 663
glabra P.J.Sm. 661, 662, 663, 665
islensis P.J.Sm. 661, 662, 663
scopulicola Carolin ex P.J.Sm. 661, 662, 663, 665
Weinmannia 179
Wetria australiensis P.I.Forst. 488
Whyanbeelia Airy Shaw & B.Hyland 9
Windolf, J. A checklist of bryophytes of the wallum habitat of
south-eastern Queensland and north-eastern New South
Wales 349
Windolf, John F.P The first botanical record for Australia 721
Wodyetia bifurcata Irvine 26
X
Xanthorrhoea 349
johnsonii 443
Ximenia americana L. 721, 722, 723
Xyris 629
Young, A.M. The Hygrocybeae (Fungi,Basidiomycota,
Agaricales, Hygrophoraceae) of the Lane Cove Bushland
Park, New South Wales 535
Z
Zamiaceae 363
Zieria 265
aspalathoides A. Cunn. ex Benth. 308
sp. (Binjour Plateau P.I.Forster 14134) 36
Zoopsis argentea (J.D.Hook. & Tayl.) J.D.Hook. & Tayl. 351
Queensland Herbarium
Publications for sale
A Key to Australian Grasses by B.K.Simon (1993), 2nd edition 206pp., card cover.
Ferns of Queensland by S.B.Andrews (1990), 427pp., illustrated, hard cover.
Flora of South-eastern Queensland by T.D. Stanley and E.M.Ross.
Volume 1 (1983), 545pp., illustrated, card cover
Volume 3 (1989), 532 pp., illustrated, hard cover.
Vegetation Survey of Queensland: South West Queensland. Queensland Botany Bulletin No. 4
by D.E.Boyland (1984), 151pp., illustrated, including a map, card cover.
Vegetation Survey of Queensland: South Central Queensland. Queensland Botany Bulletin No.3
by V.J.Neldner (1984),291pp., illustrated, including a map, card cover.
Vegetation Survey of Queensland: Central Western Queensland. Queensland Botany Bulletin
No.9 by V.J.Neldner (1991), 230pp., illustrated, including a map, card cover.
Vascular Plants of Western Queensland. Queensland Botany Bulletin No. 11 by V.J.Neldner (1992),
176pp., illustrated, card cover.
Vegetation of the Sunshine Coast: Description and Management. Queensland Botany Bulletin
No.7 by GN.Batianoff and J.A.Elsol (1989), 107pp., illustrated, including maps, card cover.
Vegetation Map and Description, Warwick, South-eastern Queensland. Queensland Botany
Bulletin No.8 by P A R/Young and T.J.McDonald (1989), 47pp., illustrated, including a map,card
cover.
Vegetation Survey and Mapping in Queensland: Its Relevance and Future, and theContribution of the
Queensland Herbarium. Queensland Botany Bulletin No. 12 by VJ.Neldner (1993), 76pp., card cover.
Suburban Weeds, Third edition by H.Kleinschmidt, A.Holland and P.Simpson (1996), 98pp.,
illustrated, card cover.
Poisonous Plants - a field guide by R.M.Dowling and R.A.McKenzie (1993), 175pp., colour
illustrations and maps, card cover.
The Flora of Girraween and Bald Rock National Parks by Bill McDonald, Colleen Gravatt, Paul
Grimshaw and John Williams (1995), 100pp., colour and line drawing illustrations and maps, card
cover.
Wildflowers of South-eastern Queensland by B.A.Lebler.
Volume 1 (1977) 108pp., illustrated, soft cover.
Volume 2 (1981) 83pp., illustrated, soft cover.
Enquiries regarding the cost and ordering of these publications should be directed to Queensland Herbarium,
Environmental Protection Agency, Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha Road, Toowong
Queensland 4066, Australia.
Austrobaileya 5(1-4): 1-735 (1997-2000)
Contents
Application of the phylogenetic species concept: A botanical monographic
perspective
Snow, Neil. 1
.A taxonomic revision ofDissiliaria F.Muel 1. exBaill. (Euphorbiaceae)
Forster,Pauli. 9
Three new species of Fontainea Heckel (Euphorbiaceae) from Australia and
Papua New Guinea
Forster,Pauli. 29
A revision of Rubus subg. Malachobatus (Focke) Focke and Rubus subg.
Diemenicus A.R.Bean (Rosaceae) in Australia
Bean,A.R. 39
Madangia inflata (Asclepiadaceae: Marsdenieae), a new genus and species
from Papua New Guinea
Forster, Paul I., Liddle David J. and Liddle, Iris M. 53
Marsdenia jensenii (Asclepiadaceae: Marsdenieae), a new species from
north-eastern Queensland
Forster,Pauli. 59
A conspectus of Astrotricha DC. (Araliaceae) in Queensland, including two
new species
Bean,A.R. 63
Cooktownia robertsii , a remarkable new genus and species of Orchidaceae
from Australia
Jones, David L. 71
A taxonomic revision of the genus Austrosteenisia R.Geesink (Fabaceae:
Millettieae)
Dixon, D J. 79
Rothia indica subsp. australis A.E.Holland (Fabaceae: Crotalarieae), a new
subspecies occuring in Australia
Holland,A.E. 93
Notes on Caesalpinia subg. Mezoneuron (Leguminosae: Caesalpinioideae)
in Australia
Pedley, Les. 97
Guioa sarcopterifucta (Sapindaceae): a new Australian species
Van Welzen, Peter C. 103
Capparis velutina (Capparaceae), a new species from south-eastern
Queensland
Forster, Pauli. 107
Notes on the naturalised Flora of Queensland, 3
Forster, Pauli. 113
A new combination in Dendrocnide (Urticaceae) in north Queensland
Jackes, B.R. and Hurley, M. 121
Notes on Eucalyptus ser. Psathyroxyla Blakely (Myrtaceae) and other ‘Ash
group’ eucalypts
Bean,A.R. 125
Austrobaileya 5(1-4): 1-735 (1997-2000)
Leptochloa southwoodii (Poaceae: Chloridoideae), a new species from south¬
east Queensland
Snow, Neil and Simon, Bryan K. 137
Lilaeopsis brisbanica (Apiaceae), a new species from Queensland, Australia
Bean,A.R. 145
Microcarpaea agonis (Scrophulariaceae), a new species from south-eastern
Queensland
Bean,A.R. 149
Book review. 153
Arevision of theBabingtonia virgata (J.R.Forst. & G.Forst.) F.Muell. complex
(Myrtaceae) in Australia
Bean,A.R. 157
Systematic and cladistic studies of MyrtellaY. Muell. and Lithomyrtus F.Muel 1.
(Myrtaceae)
Snow, Neil and Guymer, Gordon P. 173
Desmodium Desv. ( Fabaceae) and related genera in Australia: a taxonomic
revision
Pedley, Les . 209
Boronict sect. Valvatae (Benth.) Engl. (Rutaceae) in Queensland, Australia
Duretto, Marco F. 263
Taxonomic status and Australian distribution of the weedy neotropical grass
Leptochloa fusca subsp. uninervia, with an updated key to Australian
Leptochloa (Poaceae, Chloridoideae)
Snow, Neil and Simon, Bryan K. 299
Notes onAcacza(Leguminosae: Mimosoideae) chiefly from northernAustralia
Pedley, Les . 307
Two new species of Stylidium Willd. (Stylidiaceae) from north Queensland
Bean,A.R. 323
A new species of Habenaria Willd. (Orchidaceae) from North Queensland
Lavarack, Peter S. and Dockrill,Alick W. . 331
Oldenlandia gibsonii (Rubiaceae: Hedyotideae), a new species from south
east Queensland
Halford, D.A. 337
Jasminum jenniae (Oleaceae), a new species from southeastern Queensland
Harris, Wayne K. and Holmes Glenn. 341
The seedling of Cassytha glabella R.Br.
Clifford, H. Trevor. 345
A checklist of bryophytes of the wallum habitat of south-eastern Queensland
and north-eastern New South Wales
Windolf, J. 349
VanguerieaeA. Rich. exDum. (Rubiaceae) inAustralia, 1 .Everistia S.T.Reynolds
&R.J.F.Hend.
Reynolds, S.T. and Henderson, R. J.F. 353
Contribution of isozymic analysis in differentiating Macrozamia moorei
D.L. Jones and K.D.Hill from M.johnsoniiF .Muell (Zamiaceae)
Sharma, I.K., Jones, D.L. & Forster, PI. 363
Pages 369-404 do not exist in volume 5
Austrobaileya 5(1-4): 1-735 (1997-2000)
A taxonomic revision of Bridelia Willd. (Euphorbiaceae) in Australia
Forster,Pauli. 405
Rhodamnia angustifolia (Myrtaceae), a new and endangered species from
south-eastern Queensland
Snow, Neil and Guymer Gordon P. 421
A revision of Stylidium sect. Debilia Mildbr., S. sect. Floodia Mildbr. and
S. sect. Lanata A.R.Bean (Stylidiaceae)
Bean,A.R. 427
A taxonomic revision of Mallotus Lour. (Euphorbiaceae) in Australia
Forster,Pauli. 457
Ochrosperma obovatum (Myrtaceae), a new species from south-eastern
Queensland
Bean,A.R. 499
A revision obChrysopogonTxm. including Vetiveria Bory (Poaceae) in Thailand
and Malesia with notes on some other species from Africa and Australia
Veldkamp, J.F.. 503
The Hygrocybeae (Fungi, Basidiomycota, Agaricales, Hygrophoraceae) of the
Lane Cove Bushland Park, New South Wales
Young,A.M. 535
Anew species and a new combination inRwrz'<i 0 sA(Gnaphalieae: Angianthinae:
Asteraceae)
Holland,A.E. 565
Peperomia huntericma (Piperaceae), a new species from the ‘Wet Tropics’ of
north-eastern Queensland
Forster,Pauli. 573
Typification and application of the name Macrozamia macleayi Miq.
(Zamiaceae)
Forster,Pauli. 577
In defence of Chrysopogon fal 1 ax S.T.Blake (Poaceae)
Henderson, Rod. 579
Steinchisma hians (Elliott) Nash, the correct name for Fasciculochloa
sparshottiorumB.K .Simon &C.M.Weiller
Simon, Bryan. 583
Review
Bill McDonald. 585
A revision of Stylidium subg. Andersonia (R.Br. ex G.Don) Mildbr. (Stylidiaceae)
Bean,A.R. 589
Chromosome numbers of some Acanthaceae from Papua New Guinea
Daniel, Thomas F. 651
Wahlengergia celata (Campanulaceae), a new species from central Queensland
Forster,Pauli. 661
The identity of Flindersia pimenteliana and F .oppositifolia (Rutaceae):
Evidence from DNA Sequences
Scott, Kirsten D., Harris,Wayne K., Playford, Julia. 667
Cereus uruguayanus (Cactaceae) and its naturalised occurrence in Queensland,
Australia
Forster, Paul I. & Watson, Miriam. 671
Austrobaileya 5(1-4): 1-735 (1997-2000)
A revision of Eucalyptus normantonensis Maiden & Cambage (Myrtaceae)
and its allies
Bean,A.R. 679
Homoranthus coracinus (Myrtaceae), a new species from Queensland
Bean,A.R. 687
Two new species of Apatophyllum McGillivaray (Celastraceae) from
Queensland
Bean,A.R. & Jessup L.W. 691
Jasminum domatiigerum subsp. australis (Oleaceae), a new subspecies from
North Queensland
Harris, Wayne K. & McDonald, William J. 699
A new species and lectotypification in Campanulaceae: Lobelioideae
Albrecht, David. 705
Chamaesyce ophiolitica. (Euphorbiaceae), a new and endangered species
endemic to serpentine vegetation in central Queensland
Forster, Paul 1. 711
Clausena smyrelliana (Rutaceae: Aurantioideae), a new and critically
endangered species from south-east Queensland
Forster,Pauli. 715
The first botanical record for Australia
Windolf, John F.P.. 721
Rediscovery of Dischidia torricellensis( Schltr.) PI.Forst., an unusual epiphytic
asclepiad from New Guinea
Forster, Paul 1. 725
Cryptolepis papillata P.I.Forst. & Sarcolobus porcatus P.I.Forst.
(Asclepiadaceae), newly recorded from West Papua
Forster, PI. 729
A new combination in Morinda L. (Rubiaceae) for Australia
Halford, DavidA. & Henderson, Rodney J.F. 731
A new combination mProstantheraL&biM. (Lamiaceae)
Bean,A.R. 733
A new combination in Corymbia section “Politaria”: C. citriodora subsp.
variegata (Myrtaceae)
McDonald, M.W. & Bean,A.R. 735