Volume 9

Number 3 2015

A Journal of Plant Systematics

and

Conservation Biology

Queensland Herbarium

Department of Science, Information Technology and Innovation

Queensland

Government

Editorial Committee

P.I.Forster (editor)

P.D.Bostock (technical advisor)

G.P.Guymer (technical advisor)

Graphic Design

Will Smith

Desktop Publishing

Aniceta Cardoza

Austrobaileya

Vol. 1, No. 1 was published on 1 December 1977

Vol. 9, No. 2 was published on 3 December 2014

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http://www.j stor.org/

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ISSN 0155-4131

© Queensland Herbarium 2015

Web site: http://www.qld.gov.au/herbarium/publications/

Austrobaileya is the journal of the Queensland Herbarium and publishes peer-reviewed research

on plants, algae, fungi and lichens (systematics, morphology, geography, anatomy, karyology,

conservation biology and botanical history), with special emphasis on taxa from Queensland.

Opinions expressed by authors are their own and do not necessarily represent the policies or view

of the Queensland Herbarium, Department of Science, Information Technology and Innovation.

Austrobaileya 9(3): 321-461 (2015)

Contents

A taxonomic revision of Anisomeles R.Br. (Lamiaceae)

A. R. Bean . 321-381

Three new species of Taeniophyllum Blume (Orchidaceae) from northern

Queensland

B. Gray . 382-392

A taxonomic revision of Cynometra L. (Fabaceae) in Australia with a new

species from the Wet Tropics of Queensland and a range extension to the

mainland

WE. Cooper . 393-403

Olearia cuneifolia A.R.Bean & M.T.Mathieson (Asteraceae: Astereae), a

new species from Queensland

A.R.Bean & M.T.Mathieson .404-407

Eremophila woodiae Edginton (Scrophulariaceae), a new species from

Queensland

M.A.Edginton . 408-415

Rhynchospora croydonensis R.Booth (Cyperaceae), a new species from

northern Queensland

R.Booth .416-420

First record of the Gnetales in Australia: Gnetum gnemon L. (Gnetaceae) on

Badu and Mua Islands, Torres Strait, Queensland

D. G.Fell, D.J.Stanton, D.Williams, F.Loban, F.Nona , T.Stow, J.Wigness ,

E. Manas & G.Uiduldam . 421-430

Plectranthus laetus RI.Forst. and P. ventosus RI.Forst. (Lamiaceae), new

species from Cape York Peninsula, Queensland

P. I. Forster . 431-438

Overlooked plant species names associated with the botanical collections of

Eugene Fitzalan

J.L.Dowe .439-444

A conspectus of Polyscias J.R.Forst. & G.Forst. (Araliaceae) in Queensland,

Australia

A.R.Bean . 445-456

Triunia kittredgei Olde (Proteaceae): a name to be rejected

G.P.Guymer & P.I.Forster . 457-460

Validating the name Habenaria vatia D.L.Jones ex M.T.Mathieson

(Orchidaceae) for a threatened orchid species from Queensland

M.T.Mathieson .461

A taxonomic revision of Anisomeles R.Br. (Lamiaceae)

A.R. Bean

Summary

Bean, A.R. (2015). A taxonomic revision of Anisomeles R.Br. (Lamiaceae). Austrobaileya 9(3):

321-381. The genus Anisomeles R.Br., naturally occurring from southern Asia to northern Australia,

is taxonomically revised, and its phylogenetic placement is discussed. 26 species are recognised,

of which 18 species are newly described (A. antrorsa A.R.Bean, A. brevipilosa A.R.Bean, A.

bundeyensis A.R.Bean, A. carpentarica A.R.Bean, A. dallachyi A.R.Bean, A. eriodes A.R.Bean,

A. farinacea A.R.Bean, A. grandibractea A.R.Bean, A. languida A.R.Bean, A. lappa A.R.Bean, A.

leucotricha A.R.Bean, A. macdonaldii A.R.Bean, A. ornans A.R.Bean, A. papuana A.R.Bean, A.

principis A.R.Bean, A. viscidula A.R.Bean, A. vulpina A.R.Bean, A. xerophila A.R.Bean), and one,

A. ajugacea (F.M.Bailey & F.Muell.) A.R.Bean, is a new combination. Diagnostic morphological

characters are discussed, and a comprehensive identification key to the species and four regional keys

are provided. A summary of the chemical properties and cytology of two common Asian species

is presented. Illustrations, images and detailed distribution maps are provided for all species, and

their ecology and phytogeography are discussed. Lectotypes are chosen for Anisomeles candicans

Benth., A. cuneata J.Jacq. ex Fenzl, A. heyneana Benth., A. inodora R.Br., A. moschata R.Br. and A.

salviifolia R.Br.

Key Words: Lamiaceae, Anisomeles, Anisomeles ajugacea, Anisomeles antrorsa, Anisomeles

brevipilosa, Anisomeles bundeyensis, Anisomeles carpentarica, Anisomeles dallachyi, Anisomeles

eriodes, Anisomeles farinacea, Anisomeles grandibractea, Anisomeles languida, Anisomeles

lappa, Anisomeles leucotricha, Anisomeles macdonaldii, Anisomeles ornans, Anisomeles papuana,

Anisomeles principis, Anisomeles viscidula, Anisomeles vulpina, Anisomeles xerophila, Epimeredi,

Asia flora, Australia flora, Malesia flora, nomenclature, new species, indumentum, morphology,

identification keys, distribution maps, conservation status

A.R. Bean, Queensland Herbarium, Department of Science, Information Technology and Innovation,

Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. E-mail: Tony.

Bean@dsiti.qld.gov.au

Introduction

Anisomeles R.Br. is a genus belonging to

Lamiaceae subfamily Lamioideae (Harley et

al. 2004). Its species are short-lived perennial

shrubs distributed in northern Australia,

Malesia and southern Asia. The genus was

described by Robert Brown (Brown 1810),

with three species, A. salviifolia R.Br., A.

moschata R.Br. and A. inodora R.Br., based

on specimens that he himself collected

from northern Australia, and a specimen

collected earlier by Banks and Solander. Soon

afterwards, it was realised that two species

named by Linnaeus from India (Nepeta indica

L. and Nepeta malabarica L.) were referable

to Anisomeles (Sims 1819).

Bentham (1848) provided a comprehensive

taxonomic treatment of Anisomeles , where he

enumerated eight species - the three species

of Brown from Australia, and five from India

and south-east Asia. Later, Bentham (1870)

recognised only one species (A. salviifolia) for

Australia, but at the same time documented

four ‘forms’, with Brown’s three species

associated with two of these forms. He

commented, “with the very large number [of

specimens] from various localities now before

me, I am unable to assign any positive limits

to any of the following [forms]”. Bentham’s

taxonomy was subsequently followed,

and hence A. salviifolia was applied to all

Australian populations of Anisomeles for

many years. Domin (1928) maintained A.

salviifolia as the sole Australian species, but

erected several varieties based partly on the

Accepted for publication 13 August 2015

322

differing indumentum patterns. None of these

varieties was accepted by Australian botanists

of the time.

Keng (1969) used the name Anisomeles

salviifolia for some Malesian material, but

later he (Keng 1978) considered that even A.

salviifolia could not be maintained, reducing

it to a synonym of A. malabarica (L.) R.Br.

On this basis, the name A. malabarica was

subsequently adopted by all Australian

herbaria for nearly all Australian Anisomeles

specimens, and this has been mainly the

status quo until this revision. Bhatti &

Ingrouille (1996) used the name A. salviifolia

for specimens originating in Australia and

New Guinea, although this was partly based

on the erroneous belief that all four anthers

in A. salviifolia are dithecous. In fact, all

Anisomeles species have the same pattern of

two dithecous anthers and two monothecous

anthers in each flower.

Harley et al. (2004) accepted three

species for the genus, but did not name

them. Govaerts et al. (2013) have accepted

five Anisomeles species, with four occurring

in southern Asia and Malesia (A. candicans

Benth., A. heyneana Benth., A. indica (L.)

Kuntze, A. malabarica ), and two occurring

in Australia (A. malabarica, A. salviifolia).

In the current paper, the four Asian/Malesian

species accepted by Govaerts et al. (2013) are

maintained, and two are added; A. principis

A.R.Bean (found in Timor) and A. papuana

A.R.Bean (found inNewGuinea, the Moluccas

and Torres Strait, Queensland). For Australia,

23 species are recognised, being A. salviifolia

and 22 other species, including 18 species

newly named. The present study clearly

demonstrates that A. malabarica does not

occur in Australia. The taxonomic Tumping’

currently applied to Australian Anisomeles is

a legacy of the statements of Bentham (1870),

reinforced by Keng (1978), to the effect

that Anisomeles is a diffuse genus, where

intergradation is rife and there are no discrete

taxa. The current author has found that this is

not the case. While there is certainly evidence

of intergradation between some species,

many are eminently discrete and separable

from each other by discontinuities in readily

Austrobaileya 9(3): 321-381 (2015)

observable morphological characters. The

differences are often subtle, but they are at

least as robust as differences used to separate

species in other genera of Lamiaceae, and in

other related families.

Rothmaler (1944) maintained that

the generic name Epimeredi Adans. is

synonymous with Anisomeles , and that

the former should replace the latter. As a

consequence, Epimeredi was taken up for

a time by some Australian herbaria. An

unsuccessful proposal to conserve Anisomeles

against Epimeredi was made by Subramanyam

& Henry (1969). In the meantime, Backer

and Backhuisen van den Brink (1965) had

decided that Adanson’s original material

of Epimeredi probably did not include any

specimen referable to Anisomeles , and so

they maintained Anisomeles as the accepted

generic name. Parkinson (1987) considered

that none of Adanson’s unconserved generic

names were validly published. However, this

conclusion was refuted by Wilbur (1989), and

the current consensus amongst the botanical

community is that Adanson’s genera are

validly published.

I have examined high quality images of

the four Lamiaceae specimens in the Adanson

Herbarium (P-Ad.) that have Epimeredi

written on the label; these four specimens

comprise three different species. Only one of

these specimens (P00680377) can be linked

to the protologue, as one of its labels includes

the notation “h. Reg Par 1754”. This is a

reference to the Hortus Regius Parisiensis ,

cited in the protologue. The date 1754

precedes the publication date for Epimeredi ,

and the specimen is therefore original

material. This specimen is readily identifiable

as Anisomeles malabarica. The inevitable

conclusion is that Epimeredi Adans. (1763)

and Anisomeles R.Br. (1810) are congeneric.

A proposal to conserve Anisomeles against

Epimeredi is currently under consideration

by the Nomenclature Committee for Vascular

Plants (Bean 2015).

Materials and methods

This revision is based mainly on a

morphological examination of 1300 herbarium

Bean, Revision of Anisomeles

323

specimens, including 370 specimens held at

BRI. Specimen loans were obtained from

A, BKF, BM, CANB, DNA, E, G, GH, K,

L, MEL, NT, NY, P, PERTH, PR and SING.

Specimen images from E, K, MH, P and W

have either been received or viewed on the

internet. In most cases, the delicate corolla

has been examined after reconstitution

in boiling water, although spirit material

and photographic images were available

for several species, especially those from

Queensland. Close-up photographic images

of flowers have assisted in assessing corolla

characters. The author has made limited field

studies in Queensland and Northern Territory.

Data on 72 morphological characters for

the 26 species were entered into a matrix

using the Delta editor (Dallwitz et al. 1999

onwards). INTKEY software (Dallwitz et al.

1995 onwards) facilitated the development

of an interactive key, and the retrieval of

diagnostic characters for each taxon. Natural

language descriptions were also derived

from the data stored in Delta format. Where

sequential measurements are given as e.g.

5-7.5 or 8.3-10 this means that the 5 or 10 is

equivalent to 5.0 or 10.0 respectively.

The distribution maps were compiled

using DIVA-GIS Version 7.5.0, using localities

or geocodes given on the labels of specimens

and specimen images from the herbaria listed

above.

Commonly used abbreviations in the

specimen citations are HS (Homestead), NP

(National Park), Mt (Mount or Mountain;

some place or locality names are correctly

Mountain in terms of official names

[Geoscience Australia 2015]) and SF (State

Forest). Species treatments are arranged

alphabetically.

Conservation assessments are made

using the IUCN (2012) criteria but are only

recommendations as they have not been

enacted into Legislation.

Phylogenetic relationships

Cantino (1992) hypothesised a close

relationship between Anisomeles and

Pogostemon Desf. based on the shared

presence of minute leaf epidermal glands

with a unicellular cap, ‘bearded’ staminal

filaments and a lustrous pericarp. This close

relationship was further supported by the

pollen study of Abu-Asab & Cantino (1994),

with both genera shown to have very similar

pollen grains, with regular polygonal lumina

and large perforations.

Phylogenetic reconstruction using

molecular data has determined that

Anisomeles belongs in the subfamily

Lamioideae, tribe Pogostemoneae Briq.

(Scheen et al. 2010; Bendiksby et al. 2011),

and is sister to Pogostemon. Other genera in

the ‘Pogostemoneae a’ clade are Colebrookea

Sm., Craniotome Reichb. and Microtoena

Prain; these three genera are endemic to

south-east Asia. Pogostemon has a similar

distribution to Anisomeles, but with its species

diversity centred in India.

Uses and chemical properties

Indian people have used both Anisomeles

malabarica (Malabar catmint) and A. indica

(Indian catmint) as medicinal herbs for

centuries. A. malabarica has been traditionally

used to treat amentia, anoxeria, fevers,

halitosis, intestinal worms, swellings and

rheumatism (Chopra et al. 1956; Warrier et al.

1994). In recent years, chemical investigations

have sought to discover the reasons for

the perceived efficacy of Anisomeles spp.

Jeyachandran et al. (2007) claimed that an

extract from A. malabarica has anti-cancer

properties based on a study of liver disease

in mice. Kavitha et al. (2012) reported that

ethanol ic extracts and diethyl ether extracts

of A. malabarica had a statistically significant

inhibitory effect against a range of bacteria,

including Staphylococcus aureus and

Escherischia coli. Similarly, Mohanraj et al.

(2012) found an inhibitory effect against four

pathogenic bacteria, using a methanol extract

from A. malabarica.

Baranwal etal. (2012) stated that A. indica

is a source of medically active compounds

having various positive pharmacological

effects. They made reference to more than

a dozen scientific papers that document the

bioactivity of A. indica , as an analgesic, a

324

natural herbicide, an antioxidant, an anti¬

microbial agent, an anti-inflammatory, an

inhibitor of the HIV virus, and an inhibitor of

tumour cell proliferation.

Chemical analysis of Anisomeles indica

has revealed the presence of numerous

terpenoid compounds, including Anisomelic

Acid (Arisawa et al. 1986), and 15 essential

oil constituents (Yadava & Barsainya

1998). Anisomeles malabarica has been

shown to possess Anisomelic Acid, as

well as Anisomelolide, Malabaric Acid,

2-Acetoxymalabaric Acid, Anisomelyl

Acetate and Anisomelol (Preethy et al. 2013),

and essential oils.

Surprisingly, the available literature on

Australian Aboriginal ethnobotany makes no

mention of Anisomeles spp. Nor have there

been any chemical analyses performed on

Australian Anisomeles spp.

Ecology and phytogeography

All Anisomeles species are short-lived

perennial shrubs. In areas where the climate

is mesic, there may be continual growth for

some years. In places that are very dry for

part of the year, the leaves abscise and the

stems die back to ground level, then new

stems sprout from a woody rootstock upon the

arrival of the wet season. The same response

occurs after a fire. Anisomeles spp. do not

seem to be pioneer plants that colonise newly

burnt ground, but they are certainly fire-

adapted.

The present author was unable to

germinate any Anisomeles seeds. I tried

fresh field-collected seeds, as well as seeds

extracted from recently collected herbarium

specimens. A few seeds from each seedlot

were dissected to confirm the presence of

a plump embryo. Seedlots were variously

untreated, hot-water treated, or scarified. All

seeds were placed in petri dishes on blotting

paper and kept moist for five weeks, with

temperatures ranging from 17° to 27° C. No

germination was recorded for any treatment

or seedlot. Aluri (1992) similarly reported

failure in germinating seeds of A. indica ,

using different unspecified treatments in a

greenhouse, and he concluded that there are

Austrobaileya 9(3): 321-381 (2015)

unknown barriers for breaking dormancy

and subsequent germination. In contrast,

Fryer (2006) recorded that Anisomeles seeds

germinated at Kings Park, Perth, in 9-42

days with no special treatment.

Anisomeles species inhabit a wide range of

soil types, from sands to clays, but in all cases

freely draining. In some instances, the soil

is skeletal in rocky gorges and escarpments.

The parent material can be sandstone, granite,

limestone or basalt. The distribution of some

species extends into littoral areas including

continental islands, but they usually occur

away from the direct influence of salt-laden

winds.

Aluri (1992) reported that Anisomeles

malabarica and A. indica are pollinated

mainly by sunbirds ( Nectarinia sp.) and

carpenter bees (Xylocopa spp.). The bees

land on the lower corolla lip and insert their

proboscis into the base of the flower, and in

so doing the stigma brushes against the back

of its head and its thorax. The sunbird lands

on the inflorescence rachis and probes several

flowers in the vicinity, contacting the stigma

with its pollen-laden bill and forehead.

No hybrids between Anisomeles spp. have

been reported in Australian literature, nor

indicated on specimen labels, nor observed

by the current author. However, Aluri

(1992) has reported the existence of a single

individual with morphological characteristics

intermediate between A. malabarica and

A. indica , naturally occurring at Turimella,

India. Aluri & Subba Reddi (1989) bagged

unmanipulated flowers of A. malabarica and

A. indica , and found that the plants were self-

and cross-compatible and self- and cross-

pollinating. Fruit and seed production was

higher in cross-pollinated plants, but plants

can produce limited seed in the absence of

pollinators.

Anisomeles is distributed in northern

Australia, Malesia and southern Asia. It

extends west as far as Pakistan, north to the

Himalayan Range, and east to the Ryukyu

Islands of Japan. It is widespread in India

and south-east Asia, and occurs, somewhat

sporadically, in the islands of Malesia. It

Bean, Revision of Anisomeles

325

is found in Australia as far south as the

Queensland-New South Wales border. Most

members of the genus are confined to the

tropics; only A. moschata extends south

of the Tropic of Capricorn, while only A.

indica extends north of the Tropic of Cancer.

The species diversity is greatest in tropical

Australia, especially the ‘Top End’ of Northern

Territory and in northern Queensland. The

genus has not been recorded on any of the

islands of the southern Pacific Ocean, except

as a naturalised alien.

Ctyology

Krishnappa & Basavaraj (1982) reported

2 n = 34 for Anisomeles indica , and several

other studies have confirmed this. The same

authors reported 2 n = 34 for A. malabarica;

this is supported by most other studies, but

Thoppil & Jose (1998) recorded 2n = 32. No

chromosome counts have been published for

Australian Anisomeles spp.

Discussion of morphological characters

1. Habit

Anisomeles species may be prostrate (Fig.

IB) or may reach three metres in height,

according to labels of specimens from both

Asia and Australia, but more usually they

are in the height range 0.6-1.5 m (Fig. 1C).

Upright plants may become top-heavy and

lean over with large stems resting on the

ground, but continuing to grow; such plants

have here been termed procumbent.

2. Stem indumentum

Anisomeles species display a range of hair

types, and the structure, direction, density

and size of hairs are all diagnostic. The three

common hair types are 1. the hispid hair (Fig.

2A). This hair type is erect (± perpendicular

to the stem), relatively straight, eglandular,

multicellular and longer than 1 mm. The

presence of this hair type is partly related

to ontogeny. Most species have dense hispid

hairs throughout the stem on young plants, but

as the plant ages these are usually lost, at least

from the upper portions of the plant. From the

small proportion of herbarium specimens that

include the entire plant, it is evident that in the

majority of species, hispid hairs persist at the

basal part of the stem on mature plants. They

are apparently absent in A. farinacea and A.

ornans. In this study, comparisons between

taxa involving this hair type were made only

from the upper stems, 1-3 nodes below the

most proximal verticil. 2. the short, curved,

eglandular hair (Fig. 2B). This type is found

on the majority of taxa, and varies in density

from very sparse to dense, though on any

given taxon, the density is relatively uniform.

On the stems of a few species, these hairs

are antrorse; but in most species the hairs

are retrorse. In some taxa with very densely

tomentose stems, the hairs are somewhat

variable in direction. 3. the stalked glandular

hair (Fig. 2C). These are 0.1-0.3 mm long

and erect. They are frequently present on

the calyx and rachis, but on the stems they

are usually (depending on the taxon) either

abundant or absent.

The hair types present, in combination with

their density, allows the partial identification

of many species from sterile material.

3. Leaf morphology and indumentum

Leaf shape is more or less consistent for

any given taxon, and the leaf base may be

attenuate, narrowly cuneate, broadly cuneate

or obtuse (Figs. 3, 4, 5). The leaves of all

species are lobed, with the margins being

crenate, dentate or serrate. The shape, number

and the depth of the lobes are diagnostic for

some species. At the peak of each lobe is a

prominent gland. The length of the petiole

relative to the lamina is also a useful measure

that discriminates some taxa.

For the purposes of this paper, the ‘cauline

leaves’ are defined as those that are three or

more nodes below the most proximal verticil.

The ‘upper leaves’, i.e. those adjacent to or 1

or 2 nodes below the most proximal verticil,

have been measured only when cauline

leaves are not available. The ‘leaves’ or

‘bracts’ subtending a verticil (except the most

proximal) are referred to as ‘floral bracts’.

The indumentum types present on the

leaves are often the same as on the stems, but

the direction is never retrorse - they are either

antrorse, erect, or flexuose with no fixed

direction. The difference in density between

326

Austrobaileya 9(3): 321-381 (2015)

Fig. 1 . Anisomeles spp. photographed in the wild. A. flower of A. languida. B. habit of A. languida. C. flowering

plant of A. carpentarica. D. flower of A. macdonaldii. A & B from McDonald KRM14297 (BRI); C from Cowie 10489

(DNA); D from McDonald KRM14083 & Guymer (BRI).

Bean, Revision of Anisomeles

327

Fig. 2. Indumentum types of Anisomeles spp. A. stem

of A. languida with hispid hairs and short curved

eglandular hairs B. stem of A. moschata with short

curved eglandular hairs only. C. stems of A. viscidula

with stalked glandular hairs only. A from McDonald

KRM14297 (BRI), B from Batianoff 940683 & Price

(BRI), C from Egan 5038 (DNA).

the upper and lower surfaces is sometimes

diagnostic, as is the length of the hairs on both

surfaces.

4. Inflorescence structure

In all species, the inflorescence comprises

few to many verticils arranged in a spike¬

like structure at the end of each branch. Each

verticil comprises two cymose inflorescences

on opposite sides of the stem. There is much

variation in the branching pattern of the

cymes, the number of flowers per cyme, and

the distance between the individual flowers

or fruits. The leaves or bracts subtending the

verticil (except the lowest one) are referred to

as ‘floral bracts’. All other leaf-like structures

within the cymes are termed ‘bracteoles’.

There are three cyme types (Fig. 6): 1. entirely

monochasial, where every flower is borne on

328

Austrobaileya 9(3): 321-381 (2015)

Fig. 3. Cauline leaves of Anisomeles spp. (all x0.8): A. A. ajugacea (Wannan 3568 & Verdec, BRI). B. A. brevipilosa

(i O’Neill 72 , DNA). C. A. bundeyensis (Fensham 559 , DNA). D. A. candicans (Mokim 586 , G). E. A. carpentarica

(Russell-Smith 2866 & Lucas , DNA). F. A. dallachyi ( Dallachy s.n., MEL 1551746). G. A. grandibractea (Dunlop

4438 , DNA). H. A. eriodes (Forster PIF32572 & McDonald , BRI). I. A. farinacea (Edinger 496 , PERTH). Del. W.

Smith.

Bean, Revision of Anisomeles

329

Fig. 4. Cauline leaves of Anisomeles spp. (all x0.8): A. A. grandibractea {Lazarides 9106 , DNA). B. A. heyneana

{Sinclair 4562 , E). C. A. inodora {Forster PIF32933 , BRI). D. A. indica {Lei 110 , NY). E. A. languida {McDonald

KRM14297, BRI). F. A. lappa {Forster PIF12777 & Bean , BRI). G. A. leucotricha {Michell & Risler 1520 , DNA). H. A.

macdonaldii {McDonaldKRM3899 & Little , BRI). I. A. malabarica {Wight 2173 , NY). J. A. moschata {Bean 26462 ,

BRI). Del. W. Smith.

330

Austrobaileya 9(3): 321-381 (2015)

Fig. 5. Cauline leaves of Anisomeles spp. (all x0.8): A. A. moschata (McDonald KRM938, BRI). B. A. moschata

(Fensham 5361 & Butler , BRI). C. A. ornans {Hubbard & Winders 7710 , BRI). D. A. papuana (Pullen 7199 , BRI). E.

A. principis (Kenneally 8922 , PERTH). F. A. salviifolia (Randell 821 , DNA). G. A. viscidula (Eichler 22501 , DNA).

H. A. vulpina (Jensen 3350 & Kemp , BRI). I. A. antrorsa (Bean 13601 , BRI). J. A. xeropliila (Albrecht 7633 & Latz ,

NT). Del. W. Smith.

a single zig-zag rachis (i.e. two monochasia

per verticil); 2. once-dichasial, where the

first branch of the cyme is dichasial, and

thereafter monochasial (i.e. four monochasia

per verticil); and 3. twice-dichasial, where

the first two basal branches of the cyme are

dichasial, and thereafter monochasial (i.e.

eight monochasia per verticil). The cyme type

has a direct influence on the overall shape

of the verticil. The ‘entirely monochasial’

Bean, Revision of Anisomeles

331

Fig. 6. Diagrams of infloresence types in Anisomeles. A. entirely monochasial. B. once dichasial at base. C. twice

dichasial at base.

cyme results in a lax slender inflorescence

where each flower or fruit is clearly visible.

The ‘twice-dichasiaf cyme results in a

compact sub-globose inflorescence where the

branching pattern is difficult to see and many

of the flowers or fruits are obscured. The

‘once-dichasiaf cyme is of an intermediate

form. Inflorescence characters have been

assessed using fruiting material, where the

internodes are fully expanded, and it is easier

to detect the patterns involved. The number

of flowers per monochasium is typically

quite variable within species, but is of some

diagnostic value. The distance between the

individual flowers or fruits (it is the fruiting

distance that is measured in this paper) is

diagnostic for some species. Adjacent fruits

can be as much as 12 mm apart, giving rise to

a very lax cyme, or as little as 0.7 mm apart,

resulting in a congested cyme.

5. Corolla colour, shape and indumentum

(Figs. 1A, ID, 7A-C).

The corolla is predominantly pink or mauve

to purple for most species with areas of white

usually present. In Anisomeles heyneana , the

corolla is often pure white. The descriptions of

flower colour given on herbarium labels are so

varied or generalised that it makes any analysis

of the character quite pointless. The corolla

shape is remarkably consistent throughout

the genus; it consists of a small entire upper

lip, adjacent to or in contact with the stamens;

and a large lower lip, generally somewhat

recurved. The two terminal lobes are obtuse

and mostly fused, with an emarginate apex

indicating their fusion; the lateral lobes are

similarly obtuse and scarcely extend past the

base of the lateral lobes. Between the lateral

lobes is an area here dubbed the ‘platform’,

a roughly rectangular area bounded by the

lateral lobes, terminal lobes, and the throat.

The platform is sometimes flat, but often

somewhat raised or puckered, and in some

species, it bears relatively long stiff hairs. The

number of these hairs is of diagnostic value.

Within the corolla tube is the annulus, which

is surmounted by many tiny stiff erect hairs.

6. Fruiting calyx morphology

The calyx in Anisomeles grows in size from

the budding stage to anthesis and through

to fruiting, and measurements of the calyx

near anthesis will vary greatly depending on

the exact stage of floral development. All of

the features of the flowering calyx are also

present on the fruiting calyx, and are more

easily seen and assessed on the latter, and

measurements are more reliable as the calyx

has by then stopped growing.

The fruiting calyx offers a number of very

diagnostic morphological characters, which

have been assessed and measured from dried

pressed material at the fruiting stage, where

the nutlets (Fig. 7H) are mature or nearly so.

332

Austrobaileya 9(3): 321-381 (2015)

Fig. 7. Anisomeles floral morphology. A. face-on view of corolla, showing lateral and terminal lobes of the lower lip,

the corolla platform (dappled light and dark area) and the upper lip enclosing the stamens x4. B. lateral view of the

corolla and stamens x4. C. opened out flower showing the corolla lobes, the annulus (near base of stamens) and the

hairy filaments x8. D. stigma, showing unequal lobes x40. E. upper part of stamens and style showing hairy filaments,

monothecous anthers and dithecous anthers xl2. F. cylindrical calyx x8. G. obconical calyx x4. H. nutlet xl6. A-D

from A. moschata (Forster PIF28444, BRI); E from A. lappa (Forster PIF12777, BRI); F from A. moschata (Elsol 9,

BRI); G from A. indica (Newman et al. LA0486 , E); H from A. moschata (Stanley 592 , BRI). Del. W. Smith.

Bean, Revision of Anisomeles

333

The overall shape of the fruiting calyx is

very often cylindrical (Fig. 7F), or at times

narrowly campanulate, but in the Asian

species, A. indica, it is obconical (Fig. 7G).

The shape may be quantified by using the

ratio of the length versus the width (measured

as the maximum distance across calyx lobe

apices on dried material). The external surface

of the calyx is more or less smooth, apart from

the 5 or 10 rather faint longitudinal ribs, and

the external indumentum tends to reflect

that of the stems and leaves, and is often

diagnostic. The inner surface of the calyx is

often glabrous, but there is frequently a zone

of long dense hairs distal from the nutlets.

The calyx lobes are very often acute i.e.

tapering evenly to the apex, but in a few

species, most notably A. malabarica , they are

acuminate to attenuate, and the lobes have

a long and slender apex. The inner surface

of the calyx lobes is notable for the network

of anastomosing raised veins, including

an intramarginal vein that runs close to the

margin but not confluent with it. Between the

margin and this intramarginal vein are found

the calyx ‘fringe’ hairs; densely clustered

erect eglandular multicellular hairs. The

fringe hairs are highly diagnostic, because

they are, with very few exceptions, consistent

in length and pattern for any given species

(Figs. 8, 9,10,11). In some species the length

of the fringe hairs is ± constant from calyx

lobe sinus to calyx lobe apex, while in other

species the hair length decreases from sinus

to apex.

Taxonomy

Anisomeles R.Br., Prodr. 503 (1810), nom.

cons, prop.. Bean (2015). Lectotype: A.

moschata R.Br., chosen by Subramanyam &

Henry (1969).

Epimeredi Adans., Fam. PI. (Adanson)

192 (1763), nom. cons. rej. Lectotype: E.

malabaricus (R.Br.) Rothm., chosen by

Subramanyam & Henry (1969).

Perennial herbs or shrubs, with erect,

procumbent or prostrate stems arising from

a woody rootstock, usually aromatic. Sessile

glands present on nearly all plant parts.

Stems square in cross-section; indumentum

of unbranched multicellular, eglandular

hairs (short or long, curved or straight)

and/or short erect unicellular gland-tipped

hairs. Leaves petiolate, opposite, decussate,

margins crenate to dentate; the cauline leaves

transforming abruptly or gradually to the

floral bracts of the verticil. Inflorescences

terminal, spicate, indeterminate, of several

verticils at the apex of each branch; lowest

verticil borne in axil of cauline leaves, all other

verticils subtended by floral bracts. Verticils

few to many flowered, congested or lax;

branching of cymes initially often dichasial,

then monochasial, with a single flower at

each node, bracteoles persistent. Flowers

bisexual. Calyx gamosepalous, symmetric,

lengthening after anthesis, then somewhat

swollen at the proximal end; tube obconical

to narrowly campanulate or cylindrical, outer

surface with 5 or 10 longitudinal ribs, inner

surface frequently with dense multicellular

hairs on medial section; lobes 5, acute,

acuminate or attenuate, equal; outer surface

more or less smooth, inner surface with

several raised anastomosing veins, including

an intramarginal vein, and with a fringe

of straight multicellular hairs between the

intramarginal vein and the inner margin.

Corolla zygomorphic, 2-lipped, upper lip

entire, hooded, white; lower lip 3-lobed,

median lobe large, reflexed, broadening

distally and usually emarginate or bipartite.

Stamens 4, all fertile; staminal filaments

raised, adjacent to upper corolla lip, coherent

near apex, with many spreading multicellular

hairs distally or medially. Anterior anthers

transversely 2-celled, cells parallel; posterior

anthers transversely 1-celled. Style gynobasic,

glabrous; stigma branches 2, unequal. Nutlets

brown to black, smooth, glossy, with a small

basal scar.

26 species in southern Asia, Malesia and

Australia, 23 species occurring in Australia

(20 endemic).

Etymology

From the Greek anisos meaning unequal, and

melos meaning ‘a limb, or a part’, presumably

in reference to the corolla, in which the upper

lip is far smaller than the lower lip.

334

Austrobaileya 9(3): 321-381 (2015)

Fig. 8. Inner surface of fruiting calyx lobes, showing fringe hairs. A. A. ajugacea (Wannan 3568 & Verdec, BRI). B.

A. antrorsa (Bean 13601, BRI). C. A. brevipilosa (O’Neill 72, DNA). D. A. bundeyensis (Fensham 559, DNA). E. A.

candicans (Mokim 586, G). F. A. carpentarica (Russell-Smith 2866 & Lucas, DNA). G. A. dallachyi (Dallachy s.n.,

MEL 684770). H. A. grandibractea (Dunlop 4438, DNA).

Bean, Revision of Anisomeles

335

Fig. 9. Inner surface of fruiting calyx lobes, showing fringe hairs. A. A. eriodes {Forster PIF32572 & McDonald , BRI).

B. A. farinacea (Edinger 496 , PERTH). C. A. grandibractea {Lazarides 9106 , DNA). D. A. heyneana {Sinclair 4562,

E). E. A. indica {Singh 287, NY). F. A. inodor a {Forster PIF32933, BRI). G. A. languida {McDonald KRM14297, BRI).

H. A. lappa {Forster PIF30742 & McDonald, BRI).

336

Austrobaileya 9(3): 321-381 (2015)

Fig. 10 . Inner surface of fruiting calyx lobes, showing fringe hairs. A. A. leucotricha (Michell & Risler 1520 , DNA).

B. A. macdonaldii ( McDonaldKRM3899 & Little , BRI). C. A. malabarica ( Wight 2173 , NY). D. A. moschata ( Bean

26462 , BRI). E. A. ornans ( Bean 18853 , BRI). F. A. papuana ( Pullen 7199 , BRI). G. A. principis ( Kenneally 8922,

PERTH). H. A. salviifolia (Randell 821, DNA).

Bean, Revision of Anisomeles

337

Fig. 11. Inner surface of fruiting calyx lobes, showing fringe hairs. A. A. viscidula (Eich/er 22501 , DNA). B. A.

vulpina (Jensen 3350 & Kemp , BRI). C. A. xerophila (Albrecht 7633 & Latz , NT).

Key to species of Anisomeles

1 Upper stems white, indumentum very dense, hairs completely obscuring

stem at *40 magnification.2

1. Upper stems not white, indumentum very sparse to dense, stem surface

visible at *40 magnification.7

2 Leaf and stem indumentum appressed, retrorse, hairs 0.05-0.2 mm long.3

2. Leaf and stem indumentum woolly, hairs without fixed direction, 0.4-1.2 mm long . . . .4

3 External calyx hairs moderately dense to dense, spreading, 0.3-0.8 mm

long; style 12-13 mm long; leaves often narrow (3-6.6 times longer than

wide).3. A. brevipilosa

3. External calyx hairs very dense, appressed, 0.15-0.25 mm long;

style 9.5-10.5 mm long; leaves often broader (2.8-3.5 times longer than

wide).9. A. farinacea

4 Fruiting calyx lobes attenuate, 3.1-5.3 mm long; external calyx hairs

1.2-1.5 mm long.18. A. malabarica

4. Fruiting calyx lobes acute, 1.8-3.3 mm long; external calyx hairs 0.5-1 mm long.5

338 Austrobaileya 9(3): 321-381 (2015)

5 Cauline leaves with 7-14 pairs of marginal lobes; low shrub 30 to 60 cm;

verticils widely separated.20. A. ornans

5. Cauline leaves with 12-20 pairs of marginal lobes; shrub (40—) 60-200 cm;

verticils not widely separated, sometimes overlapping.6

6 Bracteoles 0.3-0.8 mm wide; corolla platform glabrous; calyx fringe

hairs 0.2-0.35 mm long at apical end.8. A. eriodes

6. Bracteoles 0.9-1.2 mm wide; corolla platform with 20-100

hairs; calyx fringe hairs 0.5-1 mm long at apical end.23. A. salviifolia

7 Erect glandular hairs (0.1-0.3 mm long) frequent to abundant on upper

stems and inflorescence rachis.8

7. Erect glandular hairs absent from upper stems and rachises, although

sessile glands usually present.11

8 Patent hispid hairs absent from upper stems; cymes twice dichasial at

base; cauline leaves 120-150 mm long; bracteoles 6.4-11 mm long. . .4. A. bundeyensis

8. Patent hispid hairs present (often frequent) on upper stems; cymes

monochasial or once dichasial at base; cauline leaves 51-110 mm long;

bracteoles 2.5-7.5 mm long.9

9 Cauline leaves 1.2-1.9 times longer than wide, base obtuse; corolla upper

lip 5.8-67 mm long.17. A. macdonaldii

9. Cauline leaves 2-3.8 times longer than wide, base cuneate; corolla

upper lip 3.5-57 mm long.10

10 Glandular hairs not extending to lower stems; hairs sparse on upper

leaf surface; corolla platform glabrous; nutlets 17-1.9 mm long.7. A. dallachyi

10. Glandular hairs extending to lower stems; hairs moderately dense to dense

on upper leaf surface; corolla platform with 1-20 hairs; nutlets

1.9-2.2 mm long.24. A. viscidula

11 All calyx fringe hairs < 0.4 mm long.12

11. Calyx fringe hairs > 0.4 mm long, at least at the sinus end.19

12 Floral bracts consistently longer than verticils.13

12. Floral bracts not consistently longer than verticils.14

13 Hispid hairs frequent on upper stems; 4-10 pairs of marginal lobes on

cauline leaves; plants prostrate or procumbent; Qld.15. A. lappa

13. Hispid hairs absent from upper stems; 12-27 pairs of marginal

lobes on cauline leaves; plants erect; N.T.10. A. grandibractea

14 Fruiting calyces obconical, 5-7.5 mm wide at lobe apices; corolla

platform with > 100 hairs.12. A. indica

14. Fruiting calyces cylindrical to narrowly campanulate, 2-5 mm wide

at lobe apices; corolla platform glabrous or with many fewer than 100 hairs.15

15 Petioles 0-3 mm long; plants prostrate.16

15. Petioles 3.5-31 mm long; plants erect.17

Bean, Revision of Anisomeles 339

16 Marginal lobes of cauline leaves 2.5-5.1 mm deep; longest leaf hairs 1-2.5

mm long; corolla annulus hairs 0.4-0.5 mm long; corolla platform

glabrous.1. A. ajugacea

16. Marginal lobes of cauline leaves 0.8-1.7 mm deep; longest leaf hairs

0.6-0.9 mm long; corolla annulus hairs 0.15-0.2 mm long; corolla

platform with 1-20 hairs.2. A. antrorsa

17 Hairs on upper leaf surface 0.9-1.3 mm long; leaves 27-50 mm long;

peduncles on lowermost verticil 0-1 mm long.25. A. vulpina

17. Hairs on upper leaf surface 0.2-0.8 mm long; leaves 44-152 mm

long; peduncles on lowermost verticil (0—)1—15 mm long.18

18 Corolla platform with 20-100 hairs; style 10-12 mm long;

fruiting calyces 7-8.9 mm long.6. A. carpentarica

18. Corolla platform with 1-20 hairs; style 12-14 mm long; fruiting calyces

8.2-11.0 mm long.22. A. principis

19 Longest hairs on the upper leaf surface 0.1-0.25 mm long.20

19. Longest hairs on the upper leaf surface 0.25-2 mm long.22

20 Leaves and stems green, indumentum sparse to moderately dense;

corolla platform glabrous.13. A. inodora

20. Leaves and stems grey or grey-green, indumentum moderately dense

to dense; corolla platform with at least some hairs.21

21 Marginal lobes of cauline leaves 4-16 pairs, lobes serrate; calyx fringe

hairs 0.15-0.3 mm long at apical end.3. A. brevipilosa

21. Marginal lobes of cauline leaves 15-37 pairs, lobes crenate; calyx fringe

hairs 0.7-1.4 mm long at apical end.16. A. leucotricha

22 Longest hairs on leaves 0.9-2 mm long.23

22. Longest hairs on leaves 0.25-0.7 mm long.24

23 Stems with patent hispid hairs; plants procumbent; external hairs on

fruiting calyx 1.1-2.4 mm long.14. A. languida

23. Stems without patent hispid hairs; plants erect; external hairs

on fruiting calyx 0.6-1 mm long.25. A. vulpina

24 Lower leaf surface glabrous or with sparse hairs (> 0.2 mm apart);

outside of fruiting calyx with a mix of glandular and

eglandular hairs.11. A. heyneana

24. Lower leaf surface with moderately dense to dense hairs (< 0.2 mm

apart); outside of fruiting calyx with eglandular hairs only.25

25 Nutlets 1.5-1.7 mm long; cauline leaves with 19-33 pairs of marginal

lobes.21. A. papuana

25. Nutlets 1.8-2.2 mm long; cauline leaves with 3-18 pairs of marginal lobes.26

26 Fruiting calyces 8.9-10.2 mm long; fruiting calyx lobes 3-3.5 mm long . . 5. A. candicans

26. Fruiting calyces 5.8-8.6 mm long; fruiting calyx lobes 1.5-2.5 mm long.27

27 Corolla platform glabrous; petioles 21-38% of lamina length.19. A. moschata

27. Corolla platform with 20-100 hairs; petioles 13-22% of lamina length . . 26. A. xerophila

340

Austrobaileya 9(3): 321-381 (2015)

Key to the Asian and Malesian species of Anisomeles

1 Upper stems white, indumentum very dense, hairs completely obscuring

stem at *40 magnification.18. A. malabarica

1. Upper stems not white, indumentum very sparse to dense, stem surface

visible at x40 magnification.2

2 All calyx fringe-hairs < 0.4 mm long.3

2. Calyx fringe-hairs > 0.4 mm long, at least at the sinus end.4

3 Fruiting calyces obconical, 5-7.5 mm wide at lobe apices; corolla platform

with more than 100 hairs.12. A. indica

3. Fruiting calyces cylindrical to narrowly campanulate, 2.0-5 mm wide at

lobe apices; corolla platform glabrous or with fewer than 20 hairs . . . . 22. A. principis

4 Lower leaf surface glabrous or with sparse hairs (> 0.2 mm apart);

outside of fruiting calyx with a mix of glandular and eglandular hairs . . 11. A. heyneana

4. Lower leaf surface with moderately dense to dense hairs (< 0.2 mm apart);

outside of fruiting calyx with eglandular hairs only.5

5 Nutlets 1.5-1.7 mm long; fruiting calyx 6.2-8.3 mm long; cauline

leaves with 19-33 pairs of marginal lobes.21. A. papuana

5. Nutlets 1.8-2.2 mm long; fruiting calyx 8.9-10.2 mm long; cauline

leaves with 3-18 pairs of marginal lobes.5. A. candicans

Key to the Western Australian species of Anisomeles

1 Upper stems white, indumentum very dense, hairs completely obscuring

stem at x40 magnification.2

1. Upper stems not white, indumentum very sparse to dense, stem surface

visible at x40 magnification.3

2 External calyx hairs moderately dense to dense, spreading, 0.3-0.8 mm

long; style 12-13 mm long; leaves often narrow (3-6.6 times longer than

wide).3. A. brevipilosa

2. External calyx hairs very dense, appressed, 0.15-0.25 mm long;

style 9.5-10.5 mm long; leaves often broader (2.8-3.5 times longer than

wide).9. A. farinacea

3 Erect glandular hairs (0.1-0.3 mm long) frequent to abundant on upper

stems and inflorescence rachis.24. A. viscidula

3. Erect glandular hairs absent from upper stems and rachises, although

sessile glands usually present.4

4 Corolla platform with 1-20 hairs; hairs on the leaves 0.25-0.8 mm long . . 22. A. principis

4. Corolla platform glabrous; hairs on the leaves 0.15-0.25 mm long.13. A. inodora

Bean, Revision of Anisomeles

341

Key to the Northern Territory species of Anisomeles

1 Upper stems white, indumentum very dense, hairs completely obscuring

stem at x40 magnification.2

1. Upper stems not white, indumentum very sparse to dense, stem surface

visible at x40 magnification.4

2 Leaf and stem indumentum appressed, retrorse, hairs 0.05-0.2 mm long.3

2. Leaf and stem indumentum woolly, hairs without fixed direction,

0.4-1.2 mm long.23. A. salviifolia

3 External calyx hairs moderately dense to dense, spreading, 0.3-0.8 mm

long; style 12-13 mm long; leaves often narrow (3-6.6 times longer than

wide).3. A. brevipilosa

3. External calyx hairs very dense, appressed, 0.15-0.25 mm long;

style 9.5-10.5 mm long; leaves often broader (2.8-3.5 times longer than

wide).9. A. farinacea

4 Erect glandular hairs (0.1-0.3 mm long) frequent to abundant on upper

stems and inflorescence rachis.5

4. Erect glandular hairs absent from upper stems and rachises, although

sessile glands usually present.6

5 Patent hispid hairs absent from upper stems; cymes twice dichasial at

base; cauline leaves 120-150 mm long; bracteoles 6.4-11 mm long. . .4. A. bundeyensis

5. Patent hispid hairs present (often frequent) on upper stems; cymes

monochasial or once dichasial at base; cauline leaves 51-110 mm long;

bracteoles 2.5-7.5 mm long.24. A. viscidula

6 Fruiting calyx fringe-hairs < 0.4 mm long.7

6. Fruiting calyx fringe-hairs > 0.4 mm long, at least at the sinus end.8

7 Floral bracts consistently longer than verticils; fruiting calyx

8.7-11 mm long.10. A. grandibractea

7. Floral bracts not consistently longer than verticils; fruiting calyx

7-8.9 mm long.6. A. carpentarica

8 Longest hairs on the upper leaf surface 0.25-0.4 mm long.26. A. xerophila

8. Longest hairs on the upper leaf surface 0.1-0.25 mm long.9

9 Leaves and stems green, indumentum sparse to moderately dense; corolla

platform glabrous.13. A. inodora

9. Leaves and stems grey or grey-green, indumentum moderately

dense to dense; corolla platform with at least some hairs.10

10 Marginal lobes of cauline leaves 4-16 pairs, lobes serrate; fruiting calyx

fringe hairs 0.15-0.3 mm long at apical end.3. A. brevipilosa

10. Marginal lobes of cauline leaves 15-37 pairs, lobes crenate; fruiting

calyx fringe hairs 0.7-1.4 mm long at apical end.16. A. leucotricha

342

Austrobaileya 9 ( 3 ): 321-381 ( 2015 )

Key to the Queensland species of Anisomeles

1 Upper stems white, indumentum very dense, hairs completely obscuring

stem at x40 magnification.2

1. Upper stems not white, indumentum very sparse to dense, stem surface

visible at x40 magnification.3

2 Cauline leaves with 7-14 pairs of marginal lobes; low shrub 30 to 60 cm

tall; verticils widely separated.20. A. ornans

2. Cauline leaves with 12-20 pairs of marginal lobes; shrub (40—) 60-200

cm tall; verticils not widely separated, sometimes overlapping.8. A. eriodes

3 Erect glandular hairs (0.1-0.3 mm long) frequent to abundant on upper

stems and inflorescence rachis.4

3. Erect glandular hairs absent from upper stems and rachises, although

sessile glands usually present.5

4 Cauline leaves 1.2-1.9 times longer than wide, base obtuse; corolla upper

lip 5.8-67 mm long; glandular hairs extending to lower stems . . . . 17. A. macdonaldii

4. Cauline leaves 2.4-3.8 times longer than wide, base cuneate; corolla upper

li 3.8-57 mm long; glandular hairs not extending to lower stems .... 7. A. dallachyi

5 Fruiting calyx fringe-hairs < 0.4 mm long.6

5. Fruiting calyx fringe-hairs > 0.4 mm long, at least at the sinus end.10

6 Floral bracts consistently longer than verticils.15. A. lappa

6. Floral bracts not consistently longer than verticils.7

7 Petioles 0-3 mm long; plants prostrate.8

7. Petioles 3.5-31 mm long; plants erect.9

8 Marginal lobes of cauline leaves 2.5-5.1 mm deep; longest leaf hairs

1-2.5 mm long; corolla annulus hairs 0.4-0.5 mm long; corolla

platform glabrous.1. A. ajugacea

8. Marginal lobes of cauline leaves 0.8-17 mm deep; longest leaf hairs

0.6-0.9 mm long; corolla annulus hairs 0.15-0.2 mm long; corolla

platform with 1-20 hairs.2. A. antrorsa

9 Hairs on upper leaf surface 0.9-1.3 mm long; leaves 27-50 mm long;

peduncles on lowermost verticil 0-1 mm long.25. A. vulpina

9. Hairs on upper leaf surface 0.2-0.8 mm long; leaves 44-152 mm long;

peduncles on lowermost verticil (0—)1—15 mm long.6. A. carpentarica

10 Fongest hairs on the upper leaf surface 0.1-0.25 mm long.11

10. Fongest hairs on the upper leaf surface 0.25-2.0 mm long.12

11 Feaves and stems green, indumentum sparse to moderately dense;

corolla platform glabrous.13. A. inodora

11. Feaves and stems grey or grey-green, indumentum moderately dense

to dense; corolla platform with at least some hairs.3. A. brevipilosa

12 Fongest hairs on leaves 0.9-2 mm long.13

12. Fongest hairs on leaves 0.25-0.7 mm long.14

Bean, Revision of Anisomeles

343

13 Stems with patent hispid hairs; plants procumbent; external hairs on

fruiting calyx 1.1-2.4 mm long. 14. A. languida

13. Stems without patent hispid hairs; plants erect; external hairs on fruiting

calyx 0.6-1 mm long. 25. A. vulpina

14 Nutlets 1.5-1.7 mm long; cauline leaves with 19-33 pairs of marginal lobes 21. A. papuana

14. Nutlets 1.8-2.2 mm long; cauline leaves with 3-18 pairs of marginal lobes. 15

15 Corolla platform glabrous; petioles 21-38% of lamina length. 19. A. moschata

15. Corolla platform with 20-100 hairs; petioles 13-22% of lamina length . . 26. A. xerophila

I. Anisomeles ajugacea (F.M.Bailey &

F.Muell.) A.R.Bean comb, nov.; Teucrium

ajugaceum F.M.Bailey & F.Muell., Synop.

Queensl. FI. Second Suppl. 48 (1888). Type:

Australia: Queensland. Cook District:

Musgrave Electric Telegraph Station,

undated, T. Bard ay-Millar s.n. (holo: BRI

[AQ340432], 2 sheets).

Anisomeles sp. (Big Coleman River

J. R.Clarkson+ 7119); Henderson (2002).

Procumbent shrub, 0.05-0.2 m high. Upper

stems and rachises with patent hispid hairs;

short curved hairs absent or retrorse, sparse;

stalked glandular hairs absent; sessile glands

8-32 mm 2 . Cauline leaves 16-29 mm long,

14-24 mm wide, 1.1-1.4 times longer than

wide, base narrowly cuneate (< 60°) or

attenuate; marginal lobes dentate, irregular

or regular, 2-4 on each side, acute, 2.5-5.1

mm deep; petioles 0-3 mm long, 0-13%

of lamina length. Lamina upper surface

indumentum appressed, eglandular, 1-2.2

mm long, moderately dense (hairs 0.1-0.2

mm apart) or dense (hairs < 0.1 mm apart),

sessile glands 8-112 mm 2 ; lower surface

indumentum appressed, eglandular, 1-2.5

mm long, moderately dense (hairs 0.1-0.2 mm

apart) or dense (hairs < 0.1 mm apart), sessile

glands 8-96 mm 2 ; transition from leaves to

floral bracts gradual. Floral bracts ovate or

broadly ovate, 5-13 mm long, 3-7 mm wide,

not consistently exceeding verticils. Verticils

(inflorescence clusters) all separated on

rachis, cymes entirely monochasial, with 3-5

flowers per monochasium, peduncles 0-3 mm

long on lowermost cluster; bracteoles obovate

or spathulate, 5-10 mm long, 1-3 mm wide.

Corolla tube same length as calyx; annulus

3.3-5 mm from base of corolla, annulus

hairs 0.4-0.5 mm long; upper lip elliptical,

4-6.6 mm long, with glandular hairs on outer

surface or with eglandular hairs on outer

surface; lower lip pink, 5.6-67 mm long to

end of lateral lobes, 9.8-11.3 mm long overall,

glabrous on platform. Longest stamens 12-14

mm long from base of corolla tube; filament

hairs 0.35-0.5 mm long, mainly along middle

part or mainly at distal end. Style 12.7-15.5

mm long; longer stigma lobe 0.85-1.2 mm

long, shorter stigma lobe 0.4-0.65 mm long.

Fruiting calyces 2-5 mm apart on rachilla;

fruiting calyx narrowly campanulate, 7-8 mm

long, 3.4-5 mm wide at lobe apices, 1.5-2.1

times longer than wide, exterior surface with

all hairs same size and type, hairs eglandular,

1.5-2.1 mm long, sessile glands 48-128 mm 2 ;

lobes acute, 2.6-3 mm long. Fruiting calyx

fringe hairs about the same length throughout,

0.2-0.35 mm long at apical end, 0.1-0.25 mm

long at sinus end, sinus hairs absent, inner

surface of tube glabrous. Nutlets 2.3-2.6 mm

long. Figs. 3A, 8A.

Additional selected specimens examined : Australia:

Queensland. Cook District: Near Little Laura River,

SSW of Laura, Jul 1990, Bean 1906 (BRI); 28.3 km from

the Kennedy River on the Jedda Creek track to King

River Station, Jun 1981, Clarkson 3717 (BRI, K, MO,

PERTH); 2 km S of the Big Coleman River on the Coen

to Musgrave Road, May 1987, Clarkson 7119 & Simon

(BRI, NSW); 3.5 km N of South Five Mile Creek on the

Peninsula Developmental Road, Apr 1991, Clarkson

8929 & Neldner (BRI, K); 27 km N of Musgrave, Cape

York Peninsula, Jun 2004, Foley s.n. (BRI [AQ610577]);

21.4 km E of Musgrave, May 2004, Gray 8910 (BRI,

CANB, CNS, NSW); Musgrave Electric Telegraph

Station, May 1891, Jacobson s.n. (BRI [AQ161262]);

25 km by road towards Jowalbinna from Peninsula

Development Road, Jul 2007, McDonald KRM6826 &

Johnson (BRI); 6.7 km along Pormpuraaw Road from

Gulf Development Road junction near Musgrave, May

2010, McDonald KRM9188 (BRI, CNS); N of Musgrave

on Cape York Development Road, just S of Red Blanket

Creek, May 2004, Wannan BSW3496 & Graham (BRI);

344

S of Musgrave, May 2004, Wannan 3564 & Verdec

(BRI); N of Musgrave, May 2004, Wannan 3568 &

Verdec (BRI); W of Musgrave, Jun 2004, Wannan

3598 (BRI); 0.5 km S of Bamboo Creek, 6 km SW of

Spion Kop, Jun 2005, Wannan 4018 & Beasley (BRI);

Telegraph Track, ‘Bamboo’, Cape York Peninsula, Jun

2008, Wannan 5223 (BRI).

Distribution and habitat : Anisomeles

ajugacea is endemic to Queensland. Most

records are within a 40 km radius of Musgrave,

Cape York Peninsula, and there are a couple

of records further south near Laura (Map 1).

It inhabits low rises or flats in woodland of

Eucalyptus tetrodonta F.Muell. and Corymbia

spp. on white sandy soils.

Phenology : Flowers are recorded from April

to July; fruits between May and August.

Notes : Anisomeles ajugacea is a highly

distinctive species because of its short

truncate leaves with just 2-4 pairs of lobes,

the petioles 0-3 mm long, the exterior calyx

hairs to 2.0 mm long, and the abundant

antrorse to erect hispid hairs on the stems and

leaves. It can produce prostrate stems up to 3

metres long ( Wannan 3564 & Verdec).

This species was placed in the genus

Teucrium by F.M. Bailey and F. Mueller, but

it clearly belongs in Anisomeles , due to its

gynobasic style, 2-lipped corolla, 2-celled

anterior anthers, and the nutlets with a very

small, basal areole.

Conservation status : Least Concern.

2. Anisomeles antrorsa A.R.Bean sp. nov.

habitu prostrato serpente, pilis antrorsis

insidentibus cauli, foliis parvis, cymis

perfecte monochasialibus et nuculis 2.5-27

mm longis distinguitur. Typus: Australia:

Queensland. Cook District: 13.7 km along

New Pennefather River Road, N of Weipa, 11

July 1998, A.R. Bean 13601 (holo: BRI).

Prostrate shrub, 0.1-0.4 m high. Upper stems

and rachises without patent hispid hairs, or

with patent hispid hairs; short curved hairs

antrorse, moderately dense or dense; stalked

glandular hairs absent. Cauline leaves 17-33

mm long, 9-21 mm wide, 1.4-2.4 times longer

than wide, base obtuse or broadly cuneate (>

60°) or narrowly cuneate (< 60°) or attenuate;

marginal lobes crenate or dentate, regular,

Austrobaileya 9(3): 321-381 (2015)

3-8 on each side, acute or obtuse, 0.8-17

mm deep; petioles 1.5-4 mm long, 5-15%

of lamina length. Lamina upper surface

indumentum erect or curved, eglandular, 0.6-

0.9 mm long, sparse (hairs > 0.2 mm apart)

or moderately dense (hairs 0.1-0.2 mm apart),

sessile glands 64-80 mm 2 ; lower surface

indumentum erect or curved, eglandular,

0.6-0.8 mm long, sparse (hairs > 0.2 mm

apart) or moderately dense (hairs 0.1-0.2 mm

apart), sessile glands 48-112 mm 2 ; transition

from leaves to floral bracts gradual. Floral

bracts ovate or broadly ovate, 6-25 mm long,

5-18 mm wide, not consistently exceeding

verticils or consistently exceeding verticils.

Verticils (inflorescence clusters) all separated

on rachis, cymes entirely monochasial, with

3-7 flowers per monochasium, peduncles 0-4

mm long on lowermost cluster; bracteoles

spathulate or linear, 4-5.8 mm long, 0.4-1.1

mm wide. Corolla tube same length as calyx;

annulus 1.9-2.5 mm from base of corolla,

annulus hairs 0.15-0.2 mm long; upper lip

ovate or elliptical, 5-5.6 mm long, with

eglandular hairs on outer surface; lower

lip 4.5-5.5 mm long to end of lateral lobes,

9-10 mm long overall, with 1-20 eglandular

hairs on platform. Longest stamens 11.5-12

mm long from base of corolla tube; filament

hairs 07-1.1 mm long, mainly at distal end.

Style 11.5-12.5 mm long; longer stigma

lobe 0.65-0.75 mm long, shorter stigma lobe

0.4-0.5 mm long. Fruiting calyces 2-3.5 mm

apart on rachilla; fruiting calyx cylindrical,

7-7.3 mm long, 2-3.3 mm wide at lobe apices,

2-3 times longer than wide, exterior surface

with hairs of different sizes or types or with

all hairs same size and type, hairs glandular

or hairs eglandular, 07-1.3 mm long, sessile

glands 80-192 mm 2 ; lobes acute, 2.3-3 mm

long. Fruiting calyx fringe hairs about the

same length throughout, 0.15-0.4 mm long

at apical end, 0.15-0.4 mm long at sinus end,

inner surface of tube with sparse long hairs

in medial section or glabrous. Nutlets 2.5-27

mm long. Figs. 51, 8B.

Additional specimens examined : Australia:

Queensland. Cook District: 4.3 km NW of Beagle

North Camp, c. 41 km NNE of Aurukun, May 1982,

Clarkson 4331 (BRI); Red Beach, Weipa area, in 1980,

Herrman s.n. (CANB); Andoom, E of Botchitt Swamp,

17.5 km NW of Lorim Point, Jan 1981, Morton AM1021

Bean, Revision of Anisomeles

345

(BRI); Community orchard and garden, Mapoon (Old

Mapoon), Mar 2005, Waterhouse BMW7207 (BRI,

CANB); Mapoon, near dump, Jun 2008, Wannan 5283 &

Graham (BRI, NSW).

Distribution and habitat : Anisomeles

antrorsa is endemic to Queensland. It is

confined to the western coast of Cape York

Peninsula between Old Mapoon and North

Camp, over a distance of c. 120 km (Map

2). It grows in sandy soil in open forest

dominated by Eucalyptus tetrodonta and

Corymbia nesophila (Blakely) K.D.Hill &

L.A.S.Johnson.

Phenology : Flowers are recorded from

January to July; fruits from May to July.

Notes : Anisomeles antrorsa is distinguished

by the antrorse hairs on the stems, the small

leaves with short petioles, the prostrate

trailing habit, the long nutlets (2.5-27 mm),

and the short calyx fringe hairs. It is most

closely related to A. ajugacea , but differs by

the shallower leaf marginal lobes, the shorter

hairs on the leaves and the exterior of the

calyx, the corolla platform with 1-20 hairs

(versus glabrous) and the hairs of the corolla

tube annulus only 0.15-0.2 mm long (versus

0.4-0.5 mm long).

Conservation status : Least Concern.

Etymology : From the Latin antrorsus ,

meaning ‘forward pointing’, and given in

reference to the antrorsely directed hairs on

the stems in this species.

3. Anisomeles brevipilosa A.R.Bean sp.

nov. pilis cauli et paginae inferiori foliorum

insidentibus longitudine minus quam 0.2

mm, pilis exterioribus calycis fructificantis

0.3-0.7 mm longis, pedunculis longis

cymarum imarum et foliis crebro lanceolatis

distinguitur. Typus: Australia: Northern

Territory. Limmen National Park, c. 1 km N

of southern lost city turnoff, southern park

boundary, 22 April 2008, D.L. Lewis 733

(holo: DNA; iso: MO).

Erect or spreading shrub, 0.6-2 m high.

Upper stems and rachises without patent

hispid hairs; short curved hairs retrorse,

dense or very dense, obscuring stem surface

at x40 magnification; stalked glandular hairs

absent; sessile glands 96-192 mm 2 . Cauline

leaves 57-93 mm long, 13-23 mm wide,

3-6.6 times longer than wide, base narrowly

cuneate (< 60°) or attenuate; marginal lobes

serrate, irregular or regular, 4-16 on each

side, obtuse, 0.2-2 mm deep; petioles 8-20

mm long, 9-35% of lamina length. Lamina

upper surface indumentum erect or curved,

eglandular or appressed, eglandular 0.1-0.2

mm long, sparse (hairs > 0.2 mm apart),

moderately dense (hairs 0.1-0.2 mm apart) or

dense (hairs < 0.1 mm apart), sessile glands

8-112 mm 2 ; lower surface indumentum erect

or curved, eglandular or appressed, eglandular

0.1-0.2 mm long, moderately dense (hairs

0.1-0.2 mm apart) or dense (hairs <0.1 mm

apart) or very dense, obscuring surface at x40

magnification, sessile glands 48-256 mm 2 ;

transition from leaves to floral bracts abrupt.

Floral bracts linear or lanceolate, 4-28 mm

long, 1-6 mm wide, not consistently exceeding

verticils. Verticils (inflorescence clusters)

overlapping near apex or all separated on

rachis, cymes entirely monochasial, or once

dichasial at base then monochasial, or twice

dichasial (±globose), with 3-12 flowers per

monochasium, peduncles 1-20 mm long on

lowermost cluster; bracteoles spathulate or

linear, 2.5-75 mm long, 0.3-1 mm wide.

Corolla tube longer than calyx, or same length

as calyx; annulus 3.5-3.8 mm from base of

corolla, annulus hairs 0.2-0.25 mm long;

upper lip elliptical, 4.5-5.2 mm long, with

glandular hairs on outer surface; lower lip

5.5-6.8 mm long to end of lateral lobes, 11.7-

13.1 mm long overall, with 20-100 eglandular

hairs on platform. Longest stamens 12-13

mm long from base of corolla tube; filament

hairs 07-1.2 mm long, mainly at distal end.

Style 12-13 mm long; longer stigma lobe

0.6-07 mm long, shorter stigma lobe 0.4-0.5

mm long. Fruiting calyces 0.9-3 mm apart on

rachilla; fruiting calyx cylindrical, 6.6-10 mm

long, 2.9-4.2 mm wide at lobe apices, 2.3-2.8

times longer than wide, exterior surface with

all hairs same size and type, hairs eglandular,

0.3-0.8 mm long, moderately dense to dense,

sessile glands 96-176 mm 2 ; lobes acute,

17-3.9 mm long. Fruiting calyx fringe hairs

longer at sinus end than at apical end, 0.15—

0.3 mm long at apical end, 07-1.2 mm long at

346

sinus end, sinus hairs absent, inner surface of

tube with dense ring of long hairs in medial

section or with sparse long hairs in medial

section. Nutlets 2-2.3 mm long. Figs. 3B, 8C.

Additional specimens examined : Australia: Western

Australia. Nimbing Range, NNW of Kununurra, May

1996, Keighery 15255 (PERTH); banks of Packsaddle

Creek, near Kununurra, Aug 1974, Kenneally 1938

(CANB, PERTH); The Gorge, Station Creek, 35 km NNE

of Carlton Hill HS, Mar 1978, Lazarides 8475 (CANB,

PERTH); c. 97 km N of Kununurra on road to coast N

of Ningbing, Jul 1995, Mitchell 4005 (CANB, PERTH);

Summit of Poompangala Hill, c. 8 km W of Kalumburu,

Apr 1991, Willing 265 (PERTH). Northern Territory,

near Caranbirini Waterhole, SW of Borroloola, Jun 1999,

Bean 15048 (BRI); 33 km N of Victoria River Downs

HS, Jun 2005, Bean 24129 (BRI, DNA, MEL); 25 miles

[41.7 km] E of Timber Creek, May 1968, Byrnes NB743

(BRI); 35 miles [56 km] SW of Borroloola, Mar 1959,

Chippendale 5553 (BRI, CANB, L, MEL, PERTH); c.

1.9 km NNE of Pungalina HS on Calvert River Road,

Pungalina Wildlife Sanctuary, Jun 2011, Jensen 2375

(BRI, DNA); 17 km SW of Kalkarmdji, May 2010, Latz

25553 (DNA, NT); Nicholson River area, Jun 1974,

Maconochie 1992 (AD, BRI, CANB, K, MO, NT);

Site 141, Kidman Springs Research Station, Aug 1990,

Manning V627 (DNA); 50 km SW of Borroloola, Jun

1977, Must 1551 (BRI, CANB, DNA, NT); 7.5 km S of

Timber Creek racecourse, Jul 1977, Must 1564 (DNA,

CANB, NT); Gregory NP, beside tributary of Bullock

Paddock Creek, Apr 1996, O’Neill 72 (DNA, MO); 4

miles [6 km] SSE of Coolibah Station, Jun 1952, Perry

2865 (BRI, CANB, NT); 14 km from Settlement Creek

on road to Calvert Hills, 1 km E of jump-up. May 1978,

Simon 3124 & Farrell (BRI); Daly River Aboriginal

Reserve, c. 95 km SE from Port Keats Mission, May

1994, Walsh 3727 (DNA, MEL); 5 km W of Robinson

River, Gulf of Carpentaria, May 1985, Wightman 1886

& Leach (BRI, CANB, DNA, K, PERTH); Fish River

station, Apr 2012, Wirf 710 (CANB, DNA). Queensland.

Burke District: Settlement Creek, 25 miles [40 km]

from coast. Gulf of Carpentaria, Jun 1948, Perry 1201

(BRI, CANB, PERTH); Gregory River crossing near

Riversleigh Station, Jun 1989, Purdie 3585 (CANB);

Bowthorn Station, 33.5 km NNW of HS, Jun 2006,

Thompson WES796 & Morgan (BRI, NSW); 100 km

WSW of Burketown, May 2008, Thompson WES1484 &

Wilson (BRI, DNA).

Distribution and habitat: Anisomeles

brevipilosa is widespread in the Northern

Territory, and extends to near Kununurra in

Western Australia, and the extreme north¬

west of Queensland (Map 3). It grows in dry

watercourses or on sandy colluvium, over

sandstone or limestone substrates.

Phenology : Flowers are recorded from March

to June; fruits from April to June.

Austrobaileya 9(3): 321-381 (2015)

Notes : Anisomeles brevipilosa can often be

distinguished solely by its narrow leaves,

up to 6.6 times longer than broad. However,

populations from the Victoria River Downs

area have broader leaves than usual, i.e. 3-4

times longer than broad. Very often, the stem

indumentum is extremely dense, obscuring

the stem surface at *40 magnification, but in

some localities the stem indumentum is dense

or moderately dense.

Anisomeles brevipilosa is closely allied to

A. far inace a; they both have extremely short

hairs (< 0.2 mm) on their leaves and upper

stems. A. brevipilosa is distinguished from A

farinacea by the presence of hispid hairs at

the base of the plant, hairs 0.3-0.8 mm long

on the outside of the calyx (versus 0.15-0.25

mm long), and not obscuring the calyx surface

(surface obscured in A. farinacea ), the lower

corolla lip 11.7-13.1 mm long (versus 6.4-10

mm long), and style 12-13 mm long (versus

9.5-10.5 mm long).

Conservation status: Least Concern.

Etymology: From the Latin brevis meaning

short and pilosus hairy. This is in reference to

the very short hairs on the stems and leaves of

this species.

4. Anisomeles bundeyensis A.R.Bean

sp. nov. pilis glandularibus abundantibus

caulibus insidentibus, foliis basalibus 125—

145 mm longis, bracteolis longissimis, cymis

bis dichasialibus in verticillis et calycibus

fructificantibus 11.4-12.6 mm longitudine

distinguitur. Typus: Australia: Northern

Territory. Mt Bundey East, 14 May 1987, R.

Fensham 559 (holo: DNA).

Erect or spreading shrub, 1-1.5 m high. Upper

stems and rachises without patent hispid hairs;

short curved hairs absent; stalked glandular

hairs abundant; glandular hairs extending

to lower stems; sessile glands 8-48 mm 2

Cauline leaves 120-150 mm long, 55-67

mm wide, 2-2.4 times longer than wide, base

obtuse or broadly cuneate (> 60°); marginal

lobes dentate or serrate, regular, 26-35 on

each side, acute, 1.5-2.2 mm deep; petioles

23-30 mm long, 18-21% of lamina length.

Lamina upper surface indumentum of erect

glandular hairs and curved eglandular hairs

Bean, Revision of Anisomeles

347

0.25-0.3 mm long, moderately dense (hairs

0.1-0.2 mm apart), sessile glands 8-48 mm 2 ;

lower surface indumentum of erect glandular

hairs 0.25-0.3 mm long, moderately dense

(hairs 0.1-0.2 mm apart), sessile glands 32-

80 mm 2 ; transition from leaves to floral bracts

abrupt. Floral bracts elliptical or ovate, 5-15

mm long, 2-4.5 mm wide, not consistently

exceeding verticils. Verticils (inflorescence

clusters) overlapping near apex, cymes twice

dichasial (± globose), with 4-7 flowers per

monochasium, peduncles 0-1 mm long on

lowermost cluster; bracteoles spathulate or

linear, 6.4-11 mm long, 0.8-1.6 mm wide.

Corolla tube shorter than calyx; annulus 3.4-4

mm from base of corolla, annulus hairs 0.25-

0.35 mm long; upper lip ovate or elliptical,

4.5-4.8 mm long, with glandular hairs on

outer surface; lower lip 7-7.2 mm long to end

of lateral lobes, 12-13.4 mm long overall, with

more than 100 eglandular hairs on platform.

Longest stamens 12-13 mm long from base

of corolla tube; filament hairs 1.2-1.3 mm

long, mainly at distal end. Style 12.5-13.5

mm long; longer stigma lobe 0.65-0.75 mm

long, shorter stigma lobe 0.55-0.7 mm long.

Fruiting calyces 0.7-1.1 mm apart on rachilla;

fruiting calyx cylindrical, 11.4-12.6 mm

long, 3.2-4 mm wide at lobe apices, 3.1-3.6

times longer than wide, exterior surface with

all hairs same size and type, hairs glandular,

0.25-0.35 mm long, sessile glands 8-32 mm 2 ;

lobes acute, 3.3-4.4 mm long. Fruiting calyx

fringe hairs about the same length throughout,

0.15-0.25 mm long at apical end, 0.15-0.25

mm long at sinus end, sinus hairs absent,

inner surface of tube with sparse long hairs

in medial section or glabrous. Nutlets 2.3-2.4

mm long. Figs. 3C, 8D.

Additional specimens examined: Australia: Northern

Territory. Mt Bundey, Arnhem Highway, Mar 1987,

Dunlop 6884 & Wightman (DNA).

Distribution and habitat: Anisomeles

bundeyensis is endemic to the Northern

Territory. It is known only from Mt Bundey

and nearby Mt Bundey East, SE of Darwin

(Map 2). It grows on granite outcrops.

Phenology : Flowers are recorded for March

and May; fruits recorded in May.

Notes: Anisomeles bundeyensis is

distinguished by the abundant glandular hairs

on the stems, the very long bracteoles, the

twice-dichasial cymes on the verticils, and

the long fruiting calyces. It is closely related

to A. viscidula, but differs by the lack of

retrorse eglandular hairs and hispid hairs on

the stems, the considerably larger corolla, the

twice-dischasially branched cymes and the

fruiting calyces only 1-1.5 mm apart.

Conservation status: This species has a very

restricted Area of Occupancy (<20 km 2 ), and

is known from just two locations. A status of

Vulnerable, criterion D2, is recommended.

Etymology: Named for the location of Mount

Bundey.

5. Anisomeles candicans Benth. in Wall.,

PI. Asiat. Rar. 1: 59 (1830); Epimeredi

candicans (Benth.) Rothm., Repert. Spec.

Nov. Regni Veg. 53: 12 (1944). Type citation:

Yenangheun, ripae Irawaddi. Type: Burma.

Yenanghuen [Yenangyaung], ripae Irawaddi,

in 1826, [Wallich Cat. No. 2038] (lecto [here

designated]: K 000846316, image!).

Anisomelesmalabaricavm. nigrescens Benth.

in Wall., PI. Asiat. Rar. 1: 59 (1830). Type

citation: ad ripas Irawaddi. Types: Burma.

Prome, in 1826, [Wallich Cat. No. 2037] (syn:

K); Irawaddi River, [1826], [Wallich Cat. No.

2037/4] (syn: K; syn: BM 000984304).

Shrub, height unknown. Upper stems and

rachises without patent hispid hairs; short

curved hairs retrorse, moderately dense;

stalked glandular hairs absent; sessile glands

8-32 mm 2 . Cauline leaves 65-91 mm long, 15-

32 mm wide, 2.6-4.3 times longer than wide,

base narrowly cuneate (< 60°) or attenuate;

marginal lobes crenate or dentate, irregular or

regular, 11-18 on each side, acute or obtuse,

0.6-2.1 mm deep; petioles 9-17 mm long, 13-

20% of lamina length. Lamina upper surface

indumentum erect or curved, eglandular,

0.3-0.4 mm long, moderately dense (hairs

0.1-0.2 mm apart), sessile glands 48-80 mm 2 ;

lower surface indumentum erect or curved,

eglandular, 0.4-0.6 mm long, dense (hairs

< 0.1 mm apart), sessile glands 8-32 mm 2 ;

transition from leaves to floral bracts abrupt.

Floral bracts lanceolate or elliptical, 6-45

348

mm long, 2-16 mm wide, not consistently

exceeding verticils. Verticils (inflorescence

clusters) all separated on rachis, cymes once

dichasial at base then monochasial, with 6-9

flowers per monochasium, peduncles 4-24

mm long on lowermost cluster; bracteoles

spathulate or linear, 3.8-8.5 mm long, 0.4-1.3

mm wide. Corolla unknown. Fruiting calyces

1.5-1.8 mm apart on rachilla; fruiting calyx

narrowly campanulate or cylindrical, 8.9-10.2

mm long, 3.6-5 mm wide at lobe apices, 1.9-

2.5 times longer than wide, exterior surface

with all hairs same size and type, hairs

eglandular, 0.4-0.6 mm long, sessile glands

48-96 mm 2 ; lobes acute, 3-3.5 mm long.

Fruiting calyx fringe hairs longer at sinus end

than at apical end, 0.2-0.5 mm long at apical

end, 1-1.4 mm long at sinus end, sinus hairs

present, inner surface of tube with dense ring

of long hairs in medial section. Nutlets 2-2.1

mm long. Figs. 3D, 8E.

Additional specimens examined: Burma. Shingaung

road, Minbu, Nov 1902, Mokim 575 (G); Minbu district,

Nov 1902, Mokim 586 (G).

Distribution and habitat : The only

specimens or specimen images I have seen

are from central Burma, all close to the

Irrawaddy River (Map 4). Murata (1971)

recorded this species from Thailand, citing

several specimens held at TI or KYO, but I

have not had the opportunity to view these.

The habitat is unknown.

Phenology : Unknown.

Notes: Anisomeles candicans is probably most

closely related to A. malabarica , but differs

from the latter by the stem indumentum being

only moderately dense; the shorter hairs on

the leaves and outside of the calyx; the calyx

fringe hairs 1-1.4 mm long at the sinus end

(0.5-0.7 mm for A. malabarica ); the lower

verticils with long peduncles; and the leaves

more deeply toothed. Wallich’s specimen

from Prome has shorter petioles than the other

specimens, but is otherwise typical.

Conservation status: Data Deficient.

6. Anisomeles carpentarica A.R.Bean

sp. nov. affinis A. moschatae sed foliis

angustioribus, pilis marginalibus loborum

calycis brevioribus, numero maximo pilorum

Austrobaileya 9(3): 321-381 (2015)

labio inferiori corollae insidentibus differens.

Typus: Australia: Northern Territory. Gray’s

Bay, in Caledon Bay, 21 June 1972, D.E.

Symon 7804 (holo: BRI; iso: AD, DNA, NT).

Erect or spreading shrub, 0.45-2.5 m high.

Upper stems and rachises without patent hispid

hairs; short curved hairs retrorse, sparse to

dense; stalked glandular hairs absent; sessile

glands 8-128 mm 2 . Cauline leaves 44-152

mm long, 20-53 mm wide, 1.6-4.1 times

longer than wide, base obtuse or broadly

cuneate (> 60°) or narrowly cuneate (< 60°) or

attenuate; marginal lobes crenate or serrate,

irregular or regular, 12-30 on each side, acute

or obtuse, 0.5-1.5 mm deep; petioles 10-31

mm long, 15-31% of lamina length. Lamina

upper surface indumentum erect or curved,

eglandular, 0.2-0.5 mm long, sparse (hairs

> 0.2 mm apart) or moderately dense (hairs

0.1-0.2 mm apart), sessile glands 8-112 mm 2 ;

lower surface indumentum erect or curved,

eglandular, 0.2-0.5 mm long, moderately

dense (hairs 0.1-0.2 mm apart) or dense

(hairs < 0.1 mm apart), sessile glands 8-128

mm 2 ; transition from leaves to floral bracts

abrupt. Floral bracts elliptical or ovate, 6-13

mm long, 2.5-7 mm wide, not consistently

exceeding verticils. Verticils (inflorescence

clusters) overlapping near apex or all separated

on rachis, cymes entirely monochasial or once

dichasial at base then monochasial or twice

dichasial (± globose), with 4-12 flowers per

monochasium, peduncles 1-12 mm long on

lowermost cluster; bracteoles spathulate or

linear, 3.5-6.5 mm long, 0.3-1.2 mm wide.

Corolla tube longer than calyx, or same length

as calyx; annulus 3.3-3.6 mm from base of

corolla, annulus hairs 0.15-0.2 mm long;

upper lip ovate or elliptical, 2.6-4.4 mm long,

with glandular hairs on outer surface or with

eglandular hairs on outer surface; lower lip

4-5.5 mm long to end of lateral lobes, 8.5-

11.7 mm long overall, with 20-100 eglandular

hairs on platform. Longest stamens 10-11.5

mm long from base of corolla tube; filament

hairs 0.7-1.2 mm long, mainly at distal end.

Style 10-12 mm long; longer stigma lobe 0.5-

0.65 mm long, shorter stigma lobe 0.3-0.4

mm long. Fruiting calyces 1-2.3 mm apart on

rachilla; fruiting calyx cylindrical, 7-8.9 mm

long, 3.3-3.8 mm wide at lobe apices, 2.1-2.4

Bean, Revision of Anisomeles

349

times longer than wide, exterior surface with

all hairs same size and type, hairs eglandular,

0.2-0.6 mm long, sessile glands 32-160 mm 2 ;

lobes acute, 1.9-2.8 mm long. Fruiting calyx

fringe hairs about the same length throughout,

0.15-0.35 mm long at apical end, 0.15-0.35

mm long at sinus end, sinus hairs absent or

present, inner surface of tube with dense ring

of long hairs in medial section or with sparse

long hairs in medial section or glabrous.

Nutlets 1.9-2.2 mm long. Figs. 1C, 3E, 8F.

Additional specimens examined : Australia: Northern

Territory. Groote Eylandt, 6.5 km SSE of Alyangula,

Apr 1992, Cowie 2557 (CANB, DNA); Arnhem Bay,

mouth of Cato River, May 1992, Cowie 2879 (DNA);

South Bay, Bickerton Island, Apr 1993, Cowie 3883

& Leach (DNA); Stevens Island, southern end, Apr

1996, Cowie 6769 (DNA, MEL); Groote Eylandt, near

Malkiyangwa Beach, Mar 2005, Cowie 10489 (DNA, L);

Walker River, May 1993, Dunlop 9531 & Leach (DNA);

Cape Arnhem, Sep 1993, Dunlop 9734 & Wightman

(DNA); N of Nhulunbuy, Oct 1993, Egan 2719 (DNA);

track to Cape Arnhem, Oct 1993, Egan 2772 (DNA);

Wessel Island, Sep 1972, Latz 3283 (CANB, DNA);

Wigram Island, English Company Islands, Jul 1992,

Leach 3061 (DNA, NT); South side of Rimbija Island,

Wessel Islands, Nov 2007, Roberts 1026 (BRI, CANB);

4 Mile Jungle, Umbakumba, Groote Eylandt, Jul 1987,

Russell-Smith 2866 & Lucas (DNA); 10 km N of Harris

Creek, Blue Mud Bay, Sep 1987, Russell-Smith 3116

& Lucas (BRI, DNA); Warangaya, Elcho Island, Sep

1987, Russell-Smith 3278 & Lucas (DNA, PERTH);

Hemple Bay, Groote Eylandt, Apr 1948, Specht 276

(BRI, CANB, L, MEL, PERTH); Port Bradshaw, Jul

1948, Specht 781 (BRI); 63 miles [101 km] west of Giddy

River crossing, Jun 1972, Symon 7748 (DNA, NT);

mouth of Bing Bong Creek, Bing Bong Station, May

1984, Thomson 643 (DNA, NT). Queensland. Burke

District. North Bountiful Island, South Wellesley

Group, Nov 2002, Thomas BOW & Pedley (BRI); 178

km NW of Burketown on Wollogorang Station (Gulf site

392), May 2008, Thompson MORN058 & Wilson (BRI);

Karumba, Jul 1960, Trapnell 186 (BRI); Bountiful

Islands, Wellesley Island Group, Mar 2008, Waterhouse

BMW7648 (BRI, CANB).

Distribution and habitat : Anisomeles

carpentarica is distributed from Karumba in

Queensland to Elcho Island in the Northern

Territory (Map 5). It mainly occurs in

littoral areas on old beach dunes, but it does

occasionally grow further inland on sandy

substrate.

Phenology : Flowers are recorded from

February to November; fruits are recorded

from April to November.

Notes : Anisomeles carpentarica is close to A.

moschata, but differs by the narrower leaves

(L/B ratio 27-4.1 times, compared to 2.1-2.8

times for A. moschata ), the longer bracteoles,

the calyx fringe hairs of uniform length

throughout, 0.15-0.35 mm long (longer

towards the sinus in A. moschata, ; 0.5-1 mm

long), and the numerous (20-100) trichomes

on the platform of the corolla (glabrous or <

20 hairs for A. moschata).

Conservation status : Least Concern.

Etymology : The epithet refers to the

distribution of this species which pre¬

dominantly fringes the Gulf of Carpentaria.

7. Anisomeles dallachyi A.R.Bean sp. nov.

pilis glandularibus abundantibus rachibus

calycibus insidentibus, foliis indumento

sparso in pagina superiore praeditis, pilis

marginalibus brevissimis loborum calycis

et nuculis 1.7-1.9 mm longis distinguitur.

Typus: Queensland. [North Kennedy

District]: Rockingham Bay, 27 April 1866, J.

Dallachy s.n. (holo: MEL 684769; iso: MEL

684766, MEL 684768, MEL 684770, MEL

684773, MEL 684774, MEL 684775, MEL

684776, MEL 684777, MEL 684778, MEL

684788).

Anisomeles salviifolia var. subtomentosa

Domin, Biblioth. Bot. 89: 567 (1928), pro

parte. Type: Queensland. [North Kennedy

District]: Rockingham Bay, undated, J.

Dallachy s.n. (syn: K).

Erect or spreading shrub, height unknown.

Upper stems and rachises without patent

hispid hairs, or with patent hispid hairs; short

curved hairs retrorse, sparse or moderately

dense; stalked glandular hairs abundant;

sessile glands 16-80 mm 2 Cauline leaves

46-96 mm long, 16-44 mm wide, 2-3.3 times

longer than wide, base narrowly cuneate (<

60°) or attenuate; marginal lobes crenate or

dentate, irregular or regular, 14-23 on each

side, acute or obtuse, 0.5-2.4 mm deep;

petioles 12-33 mm long, 26-45% of lamina

length. Lamina upper surface indumentum

of erect or appressed, curved eglandular

hairs, 0.2-0.55 mm long, sparse (hairs > 0.2

mm apart), sessile glands 8-48 mm 2 ; lower

surface indumentum of erect or curved,

350

eglandular hairs, 0.15-0.3 mm long, sparse

(hairs > 0.2 mm apart) or moderately dense

(hairs 0.1-0.2 mm apart), sessile glands 48-

96 mm 2 ; transition from leaves to floral bracts

abrupt or gradual. Floral bracts lanceolate or

elliptical, 4-27 mm long, 1.5-8 mm wide,

not consistently exceeding verticils. Verticils

(inflorescence clusters) all separated on

rachis, cymes entirely monochasial or once

dichasial at base then monochasial, with 4-7

flowers per monochasium, peduncles 0-11

mm long on lowermost cluster; bracteoles

linear, 3.2-4.5 mm long, 0.3-0.6 mm wide.

Corolla tube same length as calyx; annulus

3.8-4.2 mm from base of corolla, annulus

hairs 0.25-0.3 mm long; upper lip elliptical,

3.5-47 mm long, with eglandular hairs on

outer surface or glabrous; lower lip 5.5-6.5

mm long to end of lateral lobes, 12-14 mm

long overall, glabrous on platform. Longest

stamens 13-13.5 mm long from base of

corolla tube; filament hairs 0.3-1.5 mm long,

mainly at distal end. Style 13.5-14 mm long;

longer stigma lobe 0.8-0.9 mm long, shorter

stigma lobe 0.4-0.6 mm long. Fruiting

calyces 1.8-4 mm apart on rachilla; fruiting

calyx cylindrical, 7.1-9 mm long, 27-3.8 mm

wide at lobe apices, 1.9-2.8 times longer than

wide, exterior surface with hairs all glandular,

or with some curved eglandular hairs 0.2-0.4

mm long also present, sessile glands 32-96

mm 2 ; lobes acute, 2.2-3 mm long. Fruiting

calyx fringe hairs about the same length

throughout, 0.1-0.15 mm long at apical end,

0.1-0.2 mm long at sinus end, sinus hairs

absent, inner surface of tube with sparse long

hairs in medial section or glabrous. Nutlets

17-1.9 mm long. Figs. 3F, 8G.

Additional specimens examined : Australia:

Queensland. North Kennedy District. Beside Old

Dalrymple Track near Cardwell, May 1975, Andrews 158

& Simon (BRI); Rockingham Bay, in 1870, Dallachy s.n.

(MEL 1551746, MEL 1551726).

Distribution and habitat : Anisomeles

dallachyi is endemic to Queensland. It is

known only from the Cardwell area on the

north-eastern coast (Map 1). It inhabits

eucalypt forest on sandy soils.

Phenology : Flowers and fruits have been

recorded in April and May.

Austrobaileya 9(3): 321-381 (2015)

Typification: There are 13 sheets of this

species at MEL that were collected by J.

Dallachy from Rockingham Bay. A close

examination of the 13 sheets of A. dallachyi

reveals that they are separable into two

“groups”, based on details of their morphology

and overall similarity. One group of 11 sheets

has plant material so similar that I regard it as

originating from a single gathering; one of the

sheets has an original Dallachy label giving

the date of collection as 27 th April 1866; this

sheet is selected as the holotype, and the

other 10 sheets are regarded as isotypes. The

labels of all 11 sheets have a “B” written on

the corner indicating that they were seen by

Bentham for Flora Australiensis.

The second group of two sheets has

material closely matching, but different in

leaf shape and indumentum to the first group;

one of these sheets has the date ‘1870’ on

the label, and neither sheet has a “B” on the

corner of its label.

The syntypes of Anisomeles salviifolia

var. subtomentosa at K comprise a mixture

of A. moschata and A. dallachyi. It is the

K specimens that Domin examined before

describing that variety (Orchard 1999); he did

not see material at MEL or R

Notes : The location of Dallachy’s type

collection can be inferred. On the 26 th April

1866, he collected Ficus copiosa Steud. at

Meunga Creek (a few km north of Cardwell)

(AVH 2015). On the 27 th April 1866, he

collected Cyperus decompositus (R.Br.)

F.Muell. (AVH 2015). This species is known

to occur on sand-ridges at the southern end

of Edmund Kennedy NP, about 2 km north of

Meunga Creek (specimen at BRI, Bean 3893).

This is a likely place for Dallachy’s collection

of the Anisomeles.

Anisomeles dallachyi is probably most

closely related to A. viscidula , but A. dallachyi

differs by the lack of glandular hairs on the

lower stems, the sparse hairs of the upper

leaf surface, the glabrous corolla platform,

the mostly shorter calyx fringe hairs, and the

shorter nutlets. The only other Anisomeles

species in the Cardwell area is A. moschata, A.

dallachyi is distinguished from A. moschata

Bean, Revision of Anisomeles

351

by the stalked glandular hairs on the upper

stems, rachises and calyces; the sparsely hairy

leaves; and the much shorter calyx fringe

hairs.

Conservation status : A status of Vulnerable,

criterion D2, is recommended (IUCN 2012).

Etymology : Named for John Dallachy

(18087-1871), collector of the type specimen.

Dallachy was a very well-known botanical

collector in Victoria, New South Wales and

Queensland. In the latter state, most of his

collections are from near Rockingham Bay

(Cardwell), where he lived from 1864 until his

death in 1871.

8. Anisomeles eriodes A.R.Bean sp.

nov. indumentis densissimis in caulibus

superioribus, folii indumentis lanatis

intricatis, tubo corollae quam calyce breviore,

labio inferiore corollae glabro et calycibus

fructificantibus 8.3-97 mm longitudine

distinguitur. Typus: Australia: Queensland.

Cook District: Olive River Environmental

Reserve, 0.5 km W by road of Bromley

Homestead, 14 June 2007, P.I. Forster

PIF32572 & K.R. McDonald (holo: BRI; iso:

MEL, NSW).

Anisomeles sp. (Agnew J.R.Clarkson 4993);

Bostock & Holland (2010, 2014).

Procumbent shrub, or erect or spreading

shrub, 0.4-1 m high. Upper stems and rachises

without patent hispid hairs; short curved hairs

retrorse or no fixed direction, very dense,

obscuring stem surface at x40 magnification;

stalked glandular hairs absent. Cauline leaves

50-100 mm long, 21-40 mm wide, 1.9-3.2

times longer than wide, base narrowly cuneate

(< 60°) or attenuate; marginal lobes dentate

or serrate, regular, 15-20 on each side, acute

or obtuse, 0.3-1.7 mm deep; petioles 8-14

mm long, 13-19% of lamina length. Lamina

upper surface indumentum lanate, tangled,

0.4-0.7 mm long, dense (hairs < 0.1 mm

apart) or very dense (obscuring surface at

x40 magnification), sessile glands 32-96

mm 2 ; lower surface indumentum lanate,

tangled, 0.4-0.7 mm long, dense (hairs <0.1

mm apart) or very dense, obscuring surface

at x40 magnification; transition from leaves

to floral bracts abrupt, or gradual. Floral

bracts elliptical, 8-34 mm long, 4-13 mm

wide, not consistently exceeding verticils.

Verticils (inflorescence clusters) overlapping

near apex or all separated on rachis, cymes

entirely monochasial or once dichasial at

base then monochasial, with 6-13 flowers per

monochasium, peduncles 0-3 mm long on

lowermost cluster; bracteoles spathulate or

linear, 3.5-6.1 mm long, 0.3-0.8 mm wide.

Corolla tube shorter than calyx; annulus

2.4-3 mm from base of corolla, annulus hairs

0.15-0.25 mm long; upper lip elliptical, 4.5-

5.5 mm long, with glandular hairs on outer

surface; lower lip pink or white, 5.6-73 mm

long to end of lateral lobes, 10.6-12.5 mm

long overall, glabrous on platform. Longest

stamens 11-12.5 mm long from base of corolla

tube; filament hairs 0.5-0.9 mm long, mainly

at distal end. Style 11-13 mm long; longer

stigma lobe 07-0.9 mm long, shorter stigma

lobe 0.4-0.5 mm long. Fruiting calyces 1-1.6

mm apart on rachilla; fruiting calyx narrowly

campanulate or cylindrical, 8.3-97 mm long,

4-4.9 mm wide at lobe apices, 17-2.4 times

longer than wide, exterior surface with all

hairs same size and type, hairs eglandular,

0.6-1 mm long; lobes acute, 2.5-3.3 mm long.

Fruiting calyx fringe hairs longer at sinus end

than at apical end, 0.2-0.35 mm long at apical

end, 0.8-1.1 mm long at sinus end, sinus hairs

absent, inner surface of tube with sparse long

hairs in medial section or glabrous. Nutlets

2-2.4 mm long. Figs. 3H, 9A.

Additional selected specimens examined: Australia:

Queensland. Cook District: 23.2 km from Peninsula

Development Road, towards Iron Range, Jul 1998, Bean

13618 (BRI); Archer River, Wenlock - Coen Road, Aug

1948, Brass 19773 (BRI, CANB, K); c. 60 km directly

NE of Weipa, Jun 2008, Byrd CA31 (BRI); 1 km N of

Batavia Landing on the Weipa - Mapoon Road, Aug

1983, Clarkson 4934 (BRI, CNS, K, MEL); Batavia

Downs, Shilling paddock, c. 0.5 km W of HS, Apr 1990,

Clarkson 8574 & Neldner (AD, BRI, DNA, L); 10.8

km east of Agnew airstrip on the track to Bertihaugh

Station, Aug 1983, Clarkson 4993 (BRI, PERTH);

Merapah Holding, corridor between Rokeby and Archer

Bend NP, 25 km NW of Ranger Station, Aug 1990,

Fell DF2154 (BRI); 2 km E of Mt Gibson, 16 km SSE

of Lakeland Downs, West Normanby River catchment.

May 1993, Fell DGF3295 & Daunt (BRI); slope of St

George granites, 9.3 km E of Maitland Downs HS, Jul

2003, Fox IDF2090 (BRI); Archer Bend NP, May 1988,

Godwin C3854 (BRI); 31.3 km along Pormpuraaw

Road from Peninsula Development Road, Aug 2008,

McDonald KRM7879 (BRI); Oyala Thumotang NP, 23

352

km by road W of Peninsula Development Road, Geike

Range, May 2013, McDonald KRM14197 & Winter

(BRI, DNA, HO); Running Creek Nature Refuge, Jun

2013, McDonald KRM14489 (BRI); Lama Lama NP,

between Goose and Bull Lagoons, Jun 2013, McDonald

KRM14578 (BRI, MEL); 50 km N of Weipa airport. Little

Scrub Creek, Feb 2009, Mitchell 6-190 & Massey (BRI,

MEL, PERTH); E of Falls Creek crossing on Kennedy

Road, Jun 1982, Morton AM1732 (BRI); Nichol Creek,

Kaanju nation, central Cape York, Aug 2007, Smith 5236

& Nelson (BRI, NSW); 30 km ESE of Weipa Mission,

Sampling Point 19, Jul 1974, Specht W377 & Salt (BRI);

E of Weipa on the Peninsula Development road, Feb

2000, Wannan 1608 & Jago (BRI); East of Musgrave,

Jun 2004, Wannan 3593 (BRI, CANB); Telegraph track,

‘Bamboo’, Cape York Peninsula, Jun 2008, Wannan 5219

& Graham (BRI).

Distribution and habitat: Anisomeles

eriodes is endemic to Queensland. It is

widely distributed on Cape York Peninsula

from NE of Weipa to Maitland Downs

(Map 5). It commonly grows in woodland

with Eucalyptus tetrodonta , E. cullenii or

Corymbia nesophila on hillsides or low

pebbly rises. The soil is sandy, often reddish

in colour.

Phenology: Flowers are recorded from

February to August; fruits from June to

August.

Notes: Two poor quality specimens from

hilly terrain south-west of Cooktown {Fell

DGF3295 & Daunt ; Fox IDF2090 ; cited

above) are disjunct from the main area of

A. eriodes , and are somewhat atypical in

appearance, but are included here with it. A.

eriodes is most likely to be confused with A.

ornans. See notes under that species.

Conservation status: Least Concern.

Etymology: The epithet is from the Greek

word eriodes , meaning Tike wool, woolly’

(Brown 1956). It refers to the woolly tomentum

of the leaves and stems in this species.

9. Anisomeles farinacea A.R.Bean sp. nov.

tomento densissimo caulibus et paginae

inferiori foliorum insidente, pilis 0.05-0.15

mm in paginae inferiori foliorum et pilis

20-100 labio inferiori corollae insidentibus

distinguitur. Typus: Australia: Western

Australia. Flint Creek Gorge, 9 km NW of

homestead, 28 June 1987, D.J. Edinger 496

(holo: BRI; iso: DNA, PERTH).

Austrobaileya 9(3): 321-381 (2015)

Erect or spreading shrub, 1-2 m high. Upper

stems and rachises without patent hispid

hairs; short curved hairs retrorse, very dense,

obscuring stem surface at *40 magnification;

stalked glandular hairs absent; sessile glands

96-240 mm 2 Cauline leaves 52-111 mm

long, 15-42 mm wide, 2.8-3.5 times longer

than wide, base narrowly cuneate (< 60°) or

attenuate; marginal lobes dentate or serrate,

irregular or regular, 8-24 on each side, acute

or obtuse, 0.6-2 mm deep; petioles 13-23 mm

long, 12-21% of lamina length. Famina upper

surface indumentum appressed, eglandular,

0.05-0.2 mm long, sparse (hairs > 0.2 mm

apart) or moderately dense (hairs 0.1-0.2 mm

apart) or dense (hairs < 0.1 mm apart), sessile

glands 16-96 mm 2 ; lower surface indumentum

erect or curved, eglandular or appressed,

eglandular 0.05-0.15 mm long, dense (hairs

< 0.1 mm apart), sessile glands 48-192 mm 2 ;

transition from leaves to floral bracts abrupt.

Floral bracts linear or lanceolate, 7-24

mm long, 1-5 mm wide, not consistently

exceeding verticils or consistently exceeding

verticils. Verticils (inflorescence clusters) all

separated on rachis, cymes once dichasial at

base then monochasial or twice dichasial (±

globose), with 2-9 flowers per monochasium,

peduncles 2-7 mm long on lowermost cluster;

bracteoles linear, 4-7.5 mm long, 0.35-0.6 mm

wide. Corolla tube longer than calyx; annulus

2.5-3.2 mm from base of corolla, annulus

hairs 0.2-0.25 mm long; upper lip ovate or

elliptical, 2-3.6 mm long, with glandular and

eglandular hairs on outer surface; lower lip

3-6.5 mm long to end of lateral lobes, 6.4-10

mm long overall, with 20-100 eglandular

hairs on platform. Fongest stamens 9.5-10

mm long from base of corolla tube; filament

hairs 0.5-0.8 mm long, mainly at distal end.

Style 9.5-10.5 mm long; longer stigma lobe

0.8-0.9 mm long, shorter stigma lobe 0.5-0.6

mm long. Fruiting calyces 1-2 mm apart on

rachilla; fruiting calyx cylindrical, 8.2-9 mm

long, 27-4.4 mm wide at lobe apices, 1.9-3

times longer than wide, exterior surface with

all hairs same size and type, hairs eglandular,

0.15-0.25 mm long, very dense, sessile glands

8-128 mm 2 ; lobes acute, 2-3.1 mm long.

Fruiting calyx fringe hairs longer at sinus end

than at apical end, 0.2-0.3 mm long at apical

Bean, Revision of Anisomeles

353

end, 0.8-1.1 mm long at sinus end, sinus hairs

absent, inner surface of tube with sparse long

hairs in medial section or glabrous. Nutlets

2-2.2 mm long. Figs. 31, 9B.

Additional specimens examined: Australia: Western

Australia. King Leopold Range, Gibb River road, 42.7

km SW of turnoff to Mt House Station, May 1985, Aplin

1020 et al. (NSW, PERTH); near headwaters of King

Creek, SSW of Mt Humbert, Yampi Peninsula, Yampi

Sound Defence Training Area, Mar 2001, Barrett RLB

2135 (PERTH); 13 miles [21 km] S of Halls Creek, Jul

1968, Beard 5533 (PERTH); between Silent Grove and

the Bell Gorge parking area, Aug 2005, Byrne 1610

(PERTH); S side of Cockburn Range, c. 6.5 km W of

King River, Jul 1974, Carr 3359 & Beauglehole 47138

(MEL, NSW, PERTH); 1 km NNW of Barker River

Gorge, Napier Range, Apr 1988, Cranfield 6466 (CANB,

K, PERTH); Fern Creek, King Leopold Range, Apr 1988,

Dunlop 7773 & Simon (BRI, DNA, PERTH); Devil’s

Pass, Napier Range, May 1905, Fitzgerald606 (PERTH);

near Mt Eliza, May 1905, Fitzgerald 734 (PERTH);

Silver Gull Creek at spring, c. 14 km SE of Cockatoo

Island, Apr 1983, Fryxell & Craven 3877 (CANB, MEL,

PERTH); between McDonald Range and Glenelg River,

Jul 1950, Gardner 9583 (NSW, PERTH); Fossil Downs,

Apr 1951, Gardner 10074 (PERTH); Lower western

slopes of Mt Bell, King Leopold Ranges, May 1988,

Goble-Garratt 617 (PERTH); Near Ord River, in 1884,

Johnston s.n. (MEL); 14 km SE of Mt Kitchener, Jun

1987, Kenneally 10549 & Hyland (PERTH); Wulwuldji

Spring, Bungle Bungle NP, Nov 1989, Menkhorst 907

(DNA, PERTH); Windjana Gorge NP, in immediate

vicinity of Carpenters Gap rockshelter, Jul 1997,

Wallis LW97A/13 (PERTH); Napier Range, south side,

Windjana Gorge NP, Jun 1988, Wilson 12799 (BRI,

PERTH); March Fly Creek, 85 km NE of Lennard River

crossing, Jun 1988, Wilson 12895 (PERTH). Northern

Territory. Gregory NP, tributary of Upper East Baines

River, 50 km from Bullita Outstation, Apr 1996, Walsh

4502 & Jones (DNA).

Distribution and habitat: Anisomeles

farinacea is widespread in the Kimberley

region of Western Australia, from Cockatoo

Island in the west to south-east of Kununnurra,

and is known from a single collection in the

Northern Territory (Map 2). It frequently

inhabits watercourses or creek-banks, with

genera such as Pandanus, Terminalia and

Livistona, but it also grows on limestone

hills or on sandstone (or even granite) scree

slopes with closed-forest species or woodland

species. Soil varies from sandy loam to black

clay.

Phenology: Flowers are recorded from March

to August; fruits are recorded from April to

October.

Notes: The colour of the corolla has been

variously described on herbarium labels as

mauve, purple, purple and white, blue, or

violet.

Anisomeles farinacea is closely related to

A. brevipilosa. Distinguishing features are

given under that species.

Conservation status: Least Concern.

Etymology: The epithet is from the Latin

farina , meaning flour, given in reference to

the very pale colour of the stems and lower

leaf surfaces.

10. Anisomeles grandibractea A.R.Bean

sp. nov. bracteis floralibus maximis, pilis

brevibus in pagina superiore, monochasiis

paucifloris et pilis marginalibus brevibus

insidentibus lobis calycis distinguitur. Typus:

Australia: Northern Territory. Deaf Adder

Gorge, 23 February 1977, C.R. Dunlop 4438

(holo: DNA; iso: MEL).

Erect or spreading shrub, 0.5-1.5 m high.

Upper stems and rachises without patent

hispid hairs, or with patent hispid hairs; short

curved hairs retrorse or antrorse, sparse or

moderately dense; stalked glandular hairs

absent; sessile glands 8-96 mm 2 Cauline

leaves 44-116 mm long, 27-62 mm wide,

1.5-3.5 times longer than wide, base obtuse,

broadly cuneate or narrowly cuneate (< 60°)

or attenuate; marginal lobes crenate, dentate

or serrate, regular, 12-27 on each side, acute

or obtuse, 0.3-2.2 mm deep; petioles 12-31

mm long, 16-40% of lamina length. Lamina

upper surface indumentum erect or curved,

eglandular, 0.2-0.7 mm long, sparse (hairs

> 0.2 mm apart) or moderately dense (hairs

0.1-0.2 mm apart), sessile glands 8-48 mm 2 ;

lower surface indumentum erect or curved,

eglandular, 0.3-0.6 mm long, moderately

dense (hairs 0.1-0.2 mm apart) or dense (hairs

< 0.1 mm apart), sessile glands 8-112 mm 2 ;

transition from leaves to floral bracts gradual.

Floral bracts elliptical, ovate or broadly-ovate,

12-78 mm long, 9-30 mm wide, consistently

exceeding verticils. Verticils (inflorescence

clusters) all separated on rachis, cymes once

dichasial at base then monochasial, or twice

dichasial, with 2-8 flowers per monochasium,

peduncles 0-7 mm long on lowermost cluster;

354

bracteoles spathulate or linear, 4.3-9.5 mm

long, 0.5-1.3 mm wide. Corolla tube longer

than calyx or the same length as calyx;

annulus 3.1-4 mm from base of corolla,

annulus hairs 0.2-0.3 mm long; upper lip

elliptical, 37-5.9 mm long, with eglandular

hairs on outer surface; lower lip 4.8-76 mm

long to end of lateral lobes, 8.5-14.5 mm

long overall, platform glabrous or with 1-20

eglandular hairs. Longest stamens 12.5-14.5

mm long from base of corolla tube; filament

hairs 0.5-1.4 mm long, mainly at distal end.

Style 13-15 mm long; longer stigma lobe 0.4-

0.7 mm long, shorter stigma lobe 0.3-0.55

mm long. Fruiting calyces 1-1.5 mm apart

on rachilla; fruiting calyx cylindrical, 8.7-11

mm long, 2.8-4.3 mm wide at lobe apices,

2.3-37 times longer than wide, exterior

surface with all hairs same size and type,

hairs eglandular, longest ones 0.25-0.6 mm

long, sessile glands 8-112 mm 2 ; lobes acute,

1.8-3.3 mm long. Fruiting calyx fringe hairs

about the same length throughout, 0.15-0.35

mm long at apical end, 0.15-0.4 mm long at

sinus end, sinus hairs absent, inner surface of

tube with dense ring of long hairs in medial

section or with sparse long hairs in medial

section or glabrous. Nutlets 1.8-2.1 mm long.

Figs. 3G, 4A, 8H, 9C

Additional specimens examined : Australia: Northern

Territory. Koolpin Gorge, Mar 1990, Brennan Brel90

(DNA); 20 km SE of Twin Falls, May 1980, Craven

5880 (CANB); Mt Brockman, Feb 1973, Dunlop 3323

(BRI, CANB, DNA, NT); SE of Mt Howship, Arnhem

land, Feb 1984, Dunlop 6628 & Russell-Smith (DNA);

Nourlangie Rock, along path to lookout, Feb 1987,

Gartrell & Cunliffe UNSW19974 (BRI, CANB, DNA);

14.5 km WNW of Twin Falls, May 1980, Lazarides

9106 (DNA, CANB, DNA); walking trail, Obiri Rocks,

Apr 1987, Purdie 3175 (CANB); top of Twin Falls,

Kakadu NP, Jun 1983, Russell-Smith 711 (DNA); 5 km

E of Winwuyurr Creek crossing. Kakadu NP, Feb 1984,

Russell-Smith 1102 (DNA); 6 km S of Oenpelli, Jul

1983, Russell-Smith 1167 (DNA); 10 km SW of Oenpelli

Aboriginal Settlement, May 1988, Weher 9890 (AD,

BRI, DNA).

Distribution and habitat : Anisomeles

grandibractea is endemic to the Northern

Territory. It is confined to the eastern parts

of Kakadu NP, and the Oenpelli area of

Arnhem Land (Map 7). It inhabits sandstone

plateaux and gorges, in eucalypt woodland or

Acacia scrub, and is often recorded from the

Austrobaileya 9(3): 321-381 (2015)

margins of monsoon vine forest in sandstone

gorges, with at least some sites dominated by

Allosyncarpia ternata S.T.Blake. Soils are

skeletal or sandy.

Phenology : Flowers are recorded between

February and July; fruits from March to July.

Notes : This species is characterised by

seemingly axillary inflorescences, due to the

floral bracts greatly exceeding the verticils.

One collection (Lazarides 9106 ) has antrorse

stem hairs, while other specimens have

retrorse hairs on the stem.

Anisomeles grandibractea has two forms.

The typical form has relatively shorter and

broad floral bracts, often brown tomentum

on new growth, long hairs (0.45-0.7 mm

long) on the upper leaf surface, long petioles

(27-40% of lamina length), and obtuse or

broadly cuneate leaf bases. The other form

(represented by e.g. Lazarides 9106, Dunlop

6628 & Russell-Smith ) has very long floral

bracts, no brownish tomentum, short hairs

(0.2-0.35 mm long) on the upper leaf surface,

shorter petioles (16-28% of lamina length),

and narrowly cuneate leaf bases. Further

study may reveal these forms to be separate

species.

Conservation status : Least Concern.

Etymology : The epithet alludes to the floral

bracts of this species, which consistently

exceed the verticils.

11. Anisomeles heyneana Benth. in

N.Wallich, PI. Asiat. Rar. 1: 59 (1830);

Anisomeles secunda O.Kuntze, Revis. Gen. PI.

2: 512 (1891), nom. illeg:, Epimeredi secundus

Rothm., Repert. Spec. Nov. Regni Veg. 53:

12 (1944), nom. illeg. Type: India, undated,

B. Heyne s.n. [Wallich Cat. No. 2028] (lecto

[here designated]: K 001057386, image!).

Teucrium secundum Heyne, In Numer. List

[Wallich] n. 2028 (1829), nom. nud.

Erect or spreading shrub, 1.2-3 m high.

Upper stems and rachises without patent

hispid hairs; short curved hairs retrorse,

sparse; stalked glandular hairs absent; sessile

glands 8-16 mm 2 Cauline leaves 72-116 mm

long, 30-38 mm wide, 2.4-3.1 times longer

Bean, Revision of Anisomeles

355

than wide, base narrowly cuneate (< 60°) or

attenuate; marginal lobes dentate or serrate,

irregular or regular, 12-22 on each side,

acute, 0.5-2.3 mm deep; petioles 17-43 mm

long, 24-37% of lamina length. Lamina upper

surface indumentum appressed, eglandular,

0.6-0.7 mm long, sparse (hairs > 0.2 mm

apart), sessile glands 8-48 mm 2 ; lower surface

indumentum erect or curved, eglandular,

0.6-0.8 mm long, confined to veins or sparse

(hairs > 0.2 mm apart), sessile glands 48-128

mm 2 ; transition from leaves to floral bracts

abrupt. Floral bracts ovate, 3.5-14 mm long,

1.5- 7.5 mm wide, not consistently exceeding

verticils. Verticils (inflorescence clusters) all

separated on rachis, cymes once dichasial at

base then monochasial, with 4-8 flowers per

monochasium, peduncles 8-39 mm long on

lowermost cluster; bracteoles spathulate or

linear, 2.5-4.8 mm long, 0.3-1.2 mm wide.

Corolla tube same length as calyx; annulus

2.5- 3.5 mm from base of corolla, annulus

hairs 0.1-0.15 mm long; upper lip elliptical,

4.1-4.9 mm long, with eglandular hairs on

outer surface; lower lip 5-5.5 mm long to

end of lateral lobes, 9-10 mm long overall,

glabrous on platform or with 1-20 eglandular

hairs on platform. Longest stamens 11-12 mm

long from base of corolla tube; filament hairs

0.7-1.1 mm long, mainly along middle part.

Style 12-13 mm long; longer stigma lobe 0.6-

0.8 mm long, shorter stigma lobe 0.15-0.5

mm long. Fruiting calyces 1.3-3 mm apart on

rachilla; fruiting calyx cylindrical, 8-9.1 mm

long, 37-4.5 mm wide at lobe apices, 1.9-

2.2 times longer than wide, exterior surface

with hairs of different sizes or types, hairs

glandular and hairs eglandular, 0.4-0.9 mm

long, sessile glands 48-112 mm 2 ; lobes acute,

2.5- 3 mm long. Fruiting calyx fringe hairs

about the same length throughout or longer at

sinus end than at apical end, 0.2-0.3 mm long

at apical end, 0.2-0.8 mm long at sinus end,

sinus hairs absent or present, inner surface of

tube with sparse long hairs in medial section.

Nutlets 1.8-2.1 mm long. Figs. 4B, 9D.

Western Hill catmint.

Additional specimens examined : India. Karjat, North

Konkan, Jan 1949, Fernandes 26 (A); near Dudh

Sagor, Nov 1949, Fernandes 5430 (K); Londa, Bombay

Presidency, Jan 1950, Fernandes 1012 (K); Nil-Gherries,

1857-58, Perrottet (G); Hosangadi, Karnataka State, Jan

1939, Raja 6340 (MH); below Phonda Ghat, Jan 1853,

Ritchie 540 (E); near Kanheri Caves [between Thane &

Borivali], Bombay presidency, Oct 1945, Sinclair 4562

(E); Concan [Konkan], undated. Stocks s.n. (BRI, G, L,

MH, NY, P).

Distribution and habitat : Anisomeles

heyneana is endemic to India. It occurs close

to the west coast (Western Ghats), from

Hosangadi to Mumbai (Map 6). None of

the specimen labels include information on

habitat; however, Efloraofindia (2007-2015)

gives its habitat as “near forest clearing on

hills and slopes”.

Phenology : Flowers are recorded for January,

October, and November; fruits in January.

Notes : Bentham, in the protologue,

distinguished Anisomeles heyneana

by the secund (one-sided) pedunculate

inflorescences, the long ascending branches

and the small leaves. It is true that in A.

heyneana , one of the cymes at each verticil

sometimes does not develop forming a

secund inflorescence, but this is not a reliable

difference.

Anisomeles heyneana is most likely to be

confused with the sympatric A. indica , from

which it differs by the narrower and often

smaller leaves, the verticils all separated

along the rachis, the long peduncle of the

lower verticils, the white corolla, and the

glabrous or sparsely hairy corolla platform.

Conservation status: Data Deficient.

12. Anisomeles indica (L.) Kuntze, Revis.

Gen. PI. 2: 512 (1891); Nepeta indica L.,

Sp. PI. 2: 571 (1753); Epimeredi indicus (L.)

Rothm., Repert. Spec. Nov. Regni Veg. 53: 12

(1944). Type: Habitat in India (lecto: LINN

726.28, fide Cramer 1981).

Ballota disticha L., Mant. PI. 1: 83 (1767);

Ajuga disticha (L.) Roxb., Hort. Bengal. 44

(1814); Anisomeles disticha (L.) B.Heyne ex

Roth, Nov. PI. Sp:. 254 (1821); Nepeta disticha

(L.) Blume, Bijdr. 823 (1826). Type: Habitat

in India (lecto: LINN 1211, fide Cramer 1981).

356

Monarda zeylanica Burm.fi, FI. Ind.

(N.L.Burman) 12 (1768). Type citation:

Habitat in Zeylona. Types: Ceylon, 1672-

1677, P. Hermann (syn: BM 000621817); t.

71, J. Burman, Thesaurus Zeylanica (syn: the

illustration).

Marrubium indicum Burm.f., FI. Ind.

(N.L.Burman) 127 (1768). Type citation:

Habitat in Zeylona & Java. Types: Ceylon,

1672-1677, P. Hermann (syn: BM 000621817);

t. 71, J. Burman, Thesaurus Zeylanica (syn:

the illustration).

Nepeta amboinica L.f., Suppl. PI. 273 (1782).

Type citation: Habitat in Amboina. Types:

Nepeta indica rotundiore folio, PI. Hist. Univ.,

pt. 3, p. 415 (1699) (syn: the illustration, n.v).

Lamium garganicum Lour., FI. Cochinch.

365 (1790), nom. illeg., non L. (1763). Type

citation: incultum in Cochinchina & China.

Ballota mauritiana Pers., Syn. PI. [Persoon]

2: 126 (1806). Type: Isle de France, P.

Commerson s.n. (syn: P, image!).

Anisomeles ovata R.Br. in W.T.Aiton, Hortus

Kew. ed. 2, 3: 364 (1811), nom. illeg. [Ballota

disticha L. cited in synonymy]. Type:

probably a cultivated plant, n.v.

Anisomeles ovata var. glabrata Benth. in

Wall., PI. Asiat. Rar. 1: 59 (1830). Types:

Nepal. Hetaundah, 13 December 1826, N.

Wallich s.n. [Wallich Cat. No. 2041] (syn: K

000674248); India. Goalpara, 14 November

1808, F. Buchanan-Hamilton s.n. [Wallich

Cat. No. 2041] (syn: K 000674247).

Anisomeles ovata var. mollissima Benth. in

Wall., PI. Asiat. Rar. 1: 59 (1830); A. indica

var. mollissima (Benth.) Backer, FI. Java 2:

624 (1965). Type citation: “Prome, Taong

Dong et Sillet” Type: Bangladesh. Sylhet, N.

Wallich s.n. [Wallich Cat. No. 2039] (syn: K

[3 specimens]).

Phlomis alba Blanco, FI. Filip. 474 (1837),

nom. illeg., non Forrsk. (1775).

Anisomeles cuneata J.Jacq. ex Fenzl, Eel.

PI. Rar. 2: no. 27, t. 127 (1844). Type:

the illustration, t. 127 (lectotype [here

designated]).

Austrobaileya 9(3): 321-381 (2015)

Anisomeles malabarica var. albiflora Hassk.,

PI. Jav. Rar.: 485 (1848); Anisomeles albiflora

(Hassk.) Miq., FI. Ned. Ind. 2: 976 (1859); A.

indica var. albiflora (Hassle.) Backer, FI. Java

2: 624 (1965). Type: not cited.

Anisomeles ovata var. serratifolia Miq.,

FI. Ind. Bat. 2: 975 (1859). Type: Java, C.L.

Blume (syn: ?L, n.v)\ Borneo, P.W. Korthals

(syn: ?L, n.v).

Lophanthus argyi H.Lev., Repert. Spec.

Nov. Regni Veg. 12: 181 (1913). Type: China.

Kiang-sou [Jiangsu Sheng]: Zou-se, Vous-sie,

Sou-Tcheou, undated, P.C. dArgy s.n. (holo:

E, fruiting material only).

Anisomeles tonkinensis Gand., Bull. Soc. Bot.

France 65: 65 (1918). Type: Vietnam. Tonkin,

ad Hanoi, August 1908, A.C. DAlleizette 184

(syn: P 04443001).

Erect or spreading shrub, 0.8-2 m high. Upper

stems and rachises without patent hispid

hairs; short curved hairs retrorse, sparse to

dense; stalked glandular hairs absent; sessile

glands 8-64 mm 2 Cauline leaves 55-123 mm

long, 27-76 mm wide, 1.3-2 times longer than

wide, base obtuse or broadly cuneate (> 60°);

marginal lobes crenate or dentate or serrate,

irregular or regular, 11-22 on each side, acute

or obtuse, 0.8-4 mm deep; petioles 12-40

mm long, 22-35% of lamina length. Lamina

upper surface indumentum erect or curved,

eglandular or appressed, eglandular 0.6-1.2

mm long, sparse (hairs >0.2 mm apart) or

moderately dense (hairs 0.1-0.2 mm apart) or

dense (hairs < 0.1 mm apart), sessile glands

8-32 mm 2 ; lower surface indumentum erect or

curved, eglandular or appressed, eglandular

0.5-1.4 mm long, confined to veins or sparse

(hairs > 0.2 mm apart) or moderately dense

(hairs 0.1-0.2 mm apart) or dense (hairs

<0.1 mm apart), sessile glands 8-64 mm 2 ;

transition from leaves to floral bracts abrupt

or gradual. Floral bracts ovate, 7-15 mm long,

3-10 mm wide, not consistently exceeding

verticils. Verticils (inflorescence clusters) all

overlapping, forming continuous terminal

inflorescence or overlapping near apex, cymes

once dichasial at base then monochasial or

twice dichasial (± globose), with 3-5 flowers

per monochasium, peduncles 0-10 mm long

Bean, Revision of Anisomeles

357

on lowermost cluster; bracteoles spathulate

or linear, 27-4.2 mm long, 0.2-0.5 mm

wide. Corolla tube longer than calyx, or

same length as calyx; annulus 3-4 mm from

base of corolla, annulus hairs 0.2-0.3 mm

long; upper lip elliptical, 4.9-6.4 mm long,

with eglandular hairs on outer surface or

glabrous; lower lip 4.4-7.5 mm long to end of

lateral lobes, 10.2-11.8 mm long overall, with

more than 100 eglandular hairs on platform.

Longest stamens 13.5-17 mm long from base

of corolla tube; filament hairs 0.9-1.5 mm

long, mainly along middle part. Style 14-18

mm long; longer stigma lobe 0.6-0.9 mm

long, shorter stigma lobe 0.4-0.8 mm long.

Fruiting calyces 1-2 mm apart on rachilla;

fruiting calyx obconical, 7.2-10.8 mm long,

5-7.5 mm wide at lobe apices, 1.3-2 times

longer than wide, exterior surface with hairs

of different sizes or types or with all hairs

same size and type, hairs glandular or hairs

eglandular, 0.6-2 mm long, sessile glands

16-96 mm 2 ; lobes attenuate or acute, 3-5 mm

long. Fruiting calyx fringe hairs about the

same length throughout, 0.15-0.3 mm long

at apical end, 0.15-0.3 mm long at sinus end,

sinus hairs absent, inner surface of tube with

dense ring of long hairs in medial section or

with sparse long hairs in medial section or

glabrous. Nutlets 1.7-1.9 mm long. Figs. 4D,

7G, 9E.

Additional selected specimens examined : Caribbean.

Seamens Valley, Portland, Jamaica, Feb 1920, Maxon

& Killip 72 (A). Mauritius. lie de France, undated,

Commerson 263 (L). Pakistan, below the Lowari Pass,

Sep 1895, Gatacre 17411 (BM). India. Mawryngkneng,

Khasi Hills, Assam, Oct 1951, Chand 5035 (L); Dehra

Dun and vicinity, Nov 1927, Singh 287 (NY). Sri

Lanka. Pitiduwa, Galle district, Oct 1971, Cramer

3423 (L). Maldives. Hitaddu Islet, Sep 1964, Sigee

116 (BM). Nepal. SE of Thagaon, above Bhotekoshi

Nadi, Sep 2011, Watson EKS1N14 et al. (E). Thailand.

Huay Bankau, Nov 1971, Beusekom 3559 et al. (L);

Ban Bing Khong, Chiang Mai province, Oct 1987,

Maxwell 87-1309 (L). Laos. Along path to Ban Silia,

Khammouan, Oct 2005, Newman et al. LA0486 (E).

China. Pak Shik Ling and vicinity, Oct 1932, Lei 110

(NY); Dinghu Mountains, Oct 1963, Ting & Shih 1078

(L). Japan. Miyako Island, Ryukyus, Jan 1940, Naiko

s.n. (L). Philippines. Mindanao: Zamboanga, Feb 1904,

Hallier 4619 (L). Fiji. Levuka, May 1923, Greenwood

583 (BRI). Indonesia. Java: Buitenzorg, Oct 1922,

Bakhuizen van den Brink 1939 (L). Sulawesi: Minanga,

Dec 1895, Koorders 17361B (L). Lesser Sunda Islands:

Kada, Timor, Jul 1970, Kooy 751 (L); Komodo Island,

Jun 1982, Verheijen 4921 (L). Papua: N of Andjai village,

Kebar Valley, Nov 1954, van Royen 5023 (L). Australia.

Christmas Island, Stubbings Point, W side of South

Point, Jun 1984, Mitchell 32 (AD, CANB).

Distribution and habitat : Anisomeles indica

is a widespread and common species. It

is indigenous in southern Asia, including

Pakistan, India, Sri Lanka, Maldives, Nepal,

Bhutan, Bangladesh, Burma, Thailand, Laos,

Malaysia, Vietnam, China, Taiwan, Japan,

Philippines and Indonesia (including Papua).

From its eastern extent, the Ryukyu Islands

of southern Japan, it extends west to Pakistan,

north to the foothills of the Himalayas, and

south to Java and Christmas Island, the latter

being an external territory of Australia (Map

4). It grows on a wide range of habitats from

steep mountain slopes to alluvial flats, and on

sandy or clayey soils.

The African Plants Database (APD

2013) states that A. indica is naturalized

in Madagascar. Hedge (1998) cited two

specimens for Madagascar, commenting that

“Perhaps A. indica is present in Madagascar,

but this has never been confirmed, even as

an introduced species”. Baker (1877) stated

that the occurrences of A. ovata (=A. indica)

on nearby Mauritius were naturalisations.

Naturalised populations also occur at Fiji,

Samoa, Trinidad and Jamaica.

Phenology : Flowers and fruits have been

recorded from every month of the year.

Nomenclature : Govaerts et al. (2013) have

listed Anisomeles cuneata as a synonym

of A. malabarica. However, the very good

illustration in the protologue undoubtedly

depicts A. indica. The protologue states that

the plant originated in India, and seeds (sent

from England) were raised at Vienna botanical

garden. No specimen matching the protologue

is present at W (A. Lockher, pers. comm.

July 2013). In the absence of a specimen, the

illustration in the protologue is considered the

type of the name. Fenzl’s name appears after

the descriptive paragraph, indicating that he

wrote the validating description, though he

credits Jacquin with the species epithet.

The Rheede syntype of Nepeta amboinica

is referable to Anisochilus carnosus (L.f.)

Wall. (Suddee & Paton 2009).

358

Notes : Anisomeles indica is a very

widespread species and is accordingly very

variable. It is most readily recognised by its

obconical fruiting calyces, which are usually

considerably less than twice as long as they

are wide. All other species have fruiting

calyces that are cylindrical or at most narrowly

campanulate. The calyx fringe hairs of A.

indica are very short throughout, in contrast

to all other Asian species.

The syntypes of A. indica var. mollissima

possess a much denser indumentum and

more numerous leaf lobes than the type of

A. indica , and on that basis it is tempting

to recognise it as a distinct species, but it is

possible to find A. indica specimens with

a range of indumentum densities and leaf

lobe numbers, so that recognition even at

varietal rank seems unwise. Furthermore, the

tomentose specimens seem to have no ‘core’

distribution; they occur in Nepal, eastern

India and Sumatra.

From my study of herbarium specimens,

1 conclude that the variation in fruiting calyx

size, leaf size and shape, flower size, and

indumentum density for A. indica appears to

be continuous, and hence no infraspecific taxa

are proposed. However, a careful field-based

study may yet find that distinct taxa exist.

Conservation status : Least Concern.

13. Anisomeles inodora R.Br., Prodr. FI.

Nov. Holl. 503 (1810); Anisomeles salviifolia

var. denudata Domin, Repert. Spec. Nov.

Regni Veg. 12: 98 (1913), nom. illeg ;; A.

salviifolia var. inodora (R.Br.) Domin,

Biblioth. Bot. 89: 567 (1928); Epimeredi

inodorus (R.Br.) Rothm., Repert. Spec. Nov.

Regni Veg. 53: 12 (1944). Type: [Australia:

Northern Territory], Arnhem North Bay

[Melville Bay], 14 February 1803, R. Brown

s.n. [Bennett Number 2356] (lecto: BM

001041066 [here designated]; isolecto: BM

001041067, E 00649578).

Procumbent or erect to spreading shrub, 0.6-

2 m high. Upper stems and rachises without

patent hispid hairs, or rarely with patent hispid

hairs; short curved hairs retrorse, sparse or

moderately dense; stalked glandular hairs

absent; sessile glands 48-112 mm 2 Cauline

Austrobaileya 9(3): 321-381 (2015)

leaves 38-115 mm long, 12-43 mm wide,

I. 9-3.4 times longer than wide, base obtuse or

broadly cuneate (> 60°) or narrowly cuneate

(< 60°) or attenuate; marginal lobes crenate

or serrate, irregular or regular, 7-24 on each

side, obtuse, 0.5-3 mm deep; petioles 7-42

mm long, 20-60% of lamina length. Lamina

upper surface indumentum erect or curved,

eglandular, 0.15-0.25 mm long, confined to

midrib or sparse (hairs > 0.2 mm apart) or

moderately dense (hairs 0.1-0.2 mm apart),

sessile glands 8-48 mm 2 ; lower surface

indumentum erect or curved, eglandular,

0.15-0.25 mm long, confined to veins or

sparse (hairs >0.2 mm apart) or moderately

dense (hairs 0.1-0.2 mm apart), sessile glands

80-144 mm 2 ; transition from leaves to floral

bracts abrupt. Floral bracts lanceolate or

elliptical, 9-31 mm long, 3-11 mm wide, not

consistently exceeding verticils. Verticils

(inflorescence clusters) all separated on

rachis, cymes entirely monochasial or once

dichasial at base then monochasial, with 3-14

flowers per monochasium, peduncles 0-19

mm long on lowermost cluster; bracteoles

spathulate or linear, 27-5.8 mm long, 0.3-1.1

mm wide. Corolla tube longer than calyx, or

same length as calyx; annulus 2.3-3.3 mm

from base of corolla, annulus hairs 0.25-0.3

mm long; upper lip elliptical, 4.8-5.8 mm

long, with glandular hairs on outer surface or

with eglandular hairs on outer surface; lower

lip purple or pink, 6.3-76 mm long to end

of lateral lobes, 11.7-13.6 mm long overall,

glabrous on platform. Longest stamens

II. 5-12.5 mm long from base of corolla tube;

filament hairs 0.8-1.5 mm long, mainly at

distal end. Style 12.5-13.5 mm long; longer

stigma lobe 07-0.8 mm long, shorter stigma

lobe 0.45-0.55 mm long. Fruiting calyces 1.3-

3.8(—12) mm apart on rachilla; fruiting calyx

cylindrical, 7-9.3 mm long, 3.1-3.5 mm wide

at lobe apices, 2.1-2.9 times longer than wide,

exterior surface with hairs of different sizes

or types or with all hairs same size and type,

hairs glandular or hairs eglandular, 0.1-0.9

mm long, sessile glands 112-208 mm 2 ; lobes

acute, 1.5-2.8 mm long. Fruiting calyx fringe

hairs longer at sinus end than at apical end,

0.15-0.3 mm long at apical end, 0.6-1.1 mm

long at sinus end, sinus hairs absent, inner

Bean, Revision of Anisomeles

359

surface of tube with dense ring of long hairs

in medial section or with sparse long hairs in

medial section or glabrous. Nutlets 1.8-2.2

mm long. Figs. 4C, 9F.

Additional selected specimens examined : Australia:

Western Australia. Mitchell Plateau, N of mining

camp, Aug 1978, Beauglehole 59057 & Errey (PERTH);

Mt Barnett, Jun 1905, Fitzgerald 1093 (PERTH); 17 km

from camp towards Pt Warrender, Mitchell Plateau,

Apr 1982, Keighery 4824 (PERTH); Camp Creek, near

bauxite crusher, Mitchell Plateau, Jan 1982, Kenneally

7987 (PERTH); N end of Bougainville Peninsula

separating Admiralty Gulf and Vansittart Bay, Apr

1982, Kenneally 8091 (AD, K, PERTH); 21 km N of

mining camp, Mitchell Plateau, May 1982, Kenneally

8194 (PERTH); Cape Anjo, Jul 1973, Wilson 11296

(PERTH). Northern Territory. Nhulunbuy, Gove

Peninsula, Jun 1982, Him 134 (DNA); Cape Arnhem,

Mar 1995, Barritt 1729 (DNA, MEL); 5 km NNE of

Gwapilina Point, Port Bradshaw, Feb 1994, Brennan

2478 (DNA); Truant Island, English Company Islands,

Jul 1992, Leach 3012 (BRI, DNA); Black Point, Cobourg

Peninsula, May 1983, Wightman 402 (BRI, CANB,

DNA, MEL); Latram River, NE Arnhem Land, Feb

1988, Wightman 4163 (BRI, DNA, MEL); Giddy River

crossing, Nhulunbuy area. May 1989, Wightman 4706

(BRI, DNA). Queensland. Cook District: near Little

Laura River, SSW of Laura, Jul 1990, Bean 1681 (BRI);

Newcastle Bay, 2.5 miles [4 km] S of Somerset, May

1948, Brass 18756 (A, BRI, L); 10.2 km S of Batavia

Downs on the Peninsula Development Road, Apr 1990,

Clarkson 8432 & Neldner (BRI); Cape Melville NP, 8

km SSW of Cape Melville, Jul 1993, Fell DGF3364 &

Stanton (BRI, CANB); 102 km NW of Coen, boundary

of Archer Bend NP and Merluna Holding, Jun 1994,

Fell DGF4413 & Buck (BRI); Batavia Downs, 7.2 km

W by road of Bromley HS, towards Moreton Telegraph

Station, Jun 2007, Forster PIF32757 & McDonald (BRI,

NSW); Normanby Holding, Duffers Creek catchment, N

of Battle Camp Road, Jun 2007, Forster PIF32933 (BRI);

Snake Creek, Gulf of Carpentaria, 35 km NNW of Delta

Downs HS, Jun 2003, Fox 1DF1791 (BRI); Hammond

Island, Jul 1974, Heatwole 207 & Cameron (BRI); c. 18

km W of Cholmondeley Creek crossing, on Telegraph

Line road. Mar 1992, Johnson 5162 (BRI, NSW); Middle

Creek ridge, Errk Oykangand NP, Jun 2010, McDonald

KRM9440 & Little (BRI); Running Creek Nature Refuge.

Scrubby Lagoon, Jun 2013, McDonald KRM14501 (AD,

BRI); Pormpuraaw, Chapman Point camping ground, 1.9

km S of township. May 2009, McKenna SGM512 (BRI,

CANB, CNS); Weipa, edge of Lake Patricia vine scrub.

Mar 1980, Morton AM680 (BRI); Chuula Outstation,

Kaanju Nation, central Cape York, May 2005, Smith

4862 & Claudie (BRI); Roberts Creek, Squatter camp,

Weipa, Apr 2008, Stephensen KFS38 (BRI, CANB);

Rutland Plains, near mouth of Mitchell River, Jun 1943,

Whitehouse s.n. (BRI [AQ160159]).

Distribution and habitat : Anisomeles

inodora occurs in the northern Kimberley area

of Western Australia, in coastal parts of the

‘Top End’, Northern Territory, and throughout

Cape York Peninsula in Queensland (Map

7). It inhabits sandy soils in open eucalypt

woodland and forest, and occasionally on

sandstone outcrops.

Phenology : Flowers from February to

October; fruits from April to October.

Notes : This species is allied to Anisomeles

moschata, but differs by the shorter leaf hairs,

the sparser indumentum on the lower leaf

surface, the longer lower lip of the corolla,

the glabrescent stems and calyces glabrous or

with a few hairs along the ribs.

Specimens from the Faura region of

Queensland have particularly widely spaced

flowers and fruits and quite narrow leaves, but

from herbarium study, I am unsure whether

there is a discontinuity between this form

and typical A. inodora. Specimens from the

Kimberley region of Western Australia also

have widely spaced flowers and fruits and

narrow leaves, and they have shorter calyx

fringe-hairs than the typical form. Further

study may reveal that a taxonomic distinction

is warranted for these populations.

Conservation status : Least Concern.

14. Anisomeles languida A.R.Bean

sp. nov. habitu prostrato, praesentia

pilorum hispidorum patentium in caulibus

superioribus, pilis 1-2.4 mm longitudine foliis

calycibusque insidentibus, in pagina superiore

folii appressis, calycibus fructificantibus late

dispositis et labio inferiore corollae glabro

distinguitur. Typus: Australia: Queensland.

Cook District: Isabella Falls, Battlecamp

Road, NW of Cooktown, 7 June 2013, K.R.

McDonald KRM14297 (holo: BRI; iso: MEF,

NSW).

Prostrate shrub, 0.1-0.2 m high. Upper stems

and rachises with patent hispid hairs; short

curved hairs retrorse, sparse or moderately

dense; stalked glandular hairs absent; sessile

glands 8-64 mm 2 . Cauline leaves 55-78 mm

long, 27-37 mm wide, 1.9-2.4 times longer

than wide, base obtuse or broadly cuneate (>

60°) or narrowly cuneate (< 60°) or attenuate;

marginal lobes crenate, irregular or regular,

12-16 on each side, acute or obtuse, 0.6-1.8

360

mm deep; petioles 11-30 mm long, 18-45%

of lamina length. Lamina upper surface

indumentum appressed, eglandular, 1-2

mm long, sparse (hairs > 0.2 mm apart) or

moderately dense (hairs 0.1-0.2 mm apart),

sessile glands 32-96 mm 2 ; lower surface

indumentum erect or curved, eglandular or

appressed, eglandular 1-1.6 mm long, sparse

(hairs > 0.2 mm apart) or moderately dense

(hairs 0.1-0.2 mm apart) or dense (hairs <

0.1 mm apart), sessile glands 64-128 mm 2 ;

transition from leaves to floral bracts abrupt.

Floral bracts elliptical or ovate, 6-27 mm

long, 2.5-10 mm wide, not consistently

exceeding verticils. Verticils (inflorescence

clusters) all separated on rachis, cymes

entirely monochasial or once dichasial at

base then monochasial, with 4-9 flowers per

monochasium, peduncles 0-5.5 mm long

on lowermost cluster; bracteoles spathulate,

2.5- 67 mm long, 0.3-0.8 mm wide. Corolla

tube longer than calyx, or same length as

calyx; annulus 2.5-3.2 mm from base of

corolla, annulus hairs 0.15-0.2 mm long;

upper lip elliptical, 4-5.1 mm long, with

eglandular hairs on outer surface; lower

lip purple or pink, 5.6-6.9 mm long to end

of lateral lobes, 11-14.5 mm long overall,

glabrous on platform. Longest stamens

11.5- 13.5 mm long from base of corolla tube;

filament hairs 1.1-1.5 mm long, mainly along

middle part or mainly at distal end. Style

11.5- 14 mm long; longer stigma lobe 0.6-0.8

mm long, shorter stigma lobe 0.4-0.45 mm

long. Fruiting calyces 1.7-4.4 mm apart on

rachilla; fruiting calyx narrowly campanulate

or cylindrical, 6.9-9 mm long, 3.2-4.3 mm

wide at lobe apices, 2.1-2.4 times longer than

wide, exterior surface with all hairs same

size and type, hairs eglandular, 1.1-2.4 mm

long, sessile glands 16-96 mm 2 ; lobes acute,

1.8-3.1 mm long. Fruiting calyx fringe hairs

about the same length throughout, 0.15-0.25

mm long at apical end, 0.15-0.4 mm long at

sinus end, sinus hairs absent, inner surface of

tube with sparse long hairs in medial section

or glabrous. Nutlets 2.1-2.4 mm long. Figs.

1A, IB, 2A, 4E, 9G.

Additional specimens examined : Australia:

Queensland. Cook District: Isabella Falls, near

Cooktown, May 1970, Blake 23444 (BRI, CANB, L,

MEL, PERTH); Lockerbie, 10 miles [16 km] WSW of

Austrobaileya 9(3): 321-381 (2015)

Somerset, Apr 1948, Brass 18404 (BRI, CANB, L);

Bridge Creek Holding (proposed NP), upper Bridge

Creek catchment, NW of Cooktown, Apr 2010,

Forster PIF36462 & Thomas (BRI); Bridge Creek

Holding (proposed NP), S of Battle Camp road, NW

of Cooktown, May 2010, Forster P1F36860 & Thomas

(BRI); Cape Melville NP, Nookai Creek, May 2014,

McDonald KRM15738 (BRI, NSW); Big Fish Camp

waterhole. Silver Plains station, Jun 2014, McDonald

KRM16018 (BRI, DNA, MEL).

Distribution and habitat : Anisomeles

languida is endemic to Queensland. It occurs

in four disjunct areas of Cape York Peninsula,

viz. Battle Camp road near Cooktown, Cape

Melville, Silver Plains, and Lockerbie (Map

2). It grows in riparian woodland, or its margin

with Eucalyptus woodland on sandstone or

granite hillslopes.

Phenology : Flowers have been recorded from

April to June; fruits in May and June.

Notes: Anisomeles languida is distinguished

by the prostrate habit, the presence of patent

hispid hairs on the upper stems, the hairs

1-2.4 mm long on leaves and calyces, the

hairs appressed on the upper leaf surface,

the widely spaced fruiting calyces, and the

glabrous platform of the corolla.

Conservation status : Least Concern.

Etymology: From the Latin languidus,

meaning listless or weary. This is an oblique

reference to the prostrate habit of the plant; it

seems too weary to battle against gravity.

15. Anisomeles lappa A.R.Bean sp. nov.

habitu prostrato vel procumbente, pilis

hispidis crebris caulibus insidentibus, foliis

basalibus paribus 4-10 loborum marginalium

praeditis, monochasiis paucifloris et bracteis

floralibus longitudine verticillos superantibus

distinguitur. Typus: Australia: Queensland.

Cook District: 1 km S of Lappa on Mt Garnet

Road, 12 April 2005, P.I. Forster PIF30742 &

K.R. McDonald (holo: BRI; iso: MEL, NSW).

Anisomeles salviifolia var. tomentoso-hirsuta

Domin, Biblioth. Bot. 89: 567 (1928). Type:

Australia: Queensland. Cook District: Walsh

River near Chillagoe, February 1910, K.

Domin s.n. (holo: PR 530817).

Prostrate or procumbent shrub, 0.15-0.5

m high. Upper stems and rachises with

Bean, Revision of Anisomeles

361

patent hispid hairs; short curved hairs

retrorse, moderately dense or dense; stalked

glandular hairs absent; sessile glands 8-80

mm 2 Cauline leaves 26-49 mm long, 12-25

mm wide, 2-3.3 times longer than wide,

base narrowly cuneate (< 60°) or attenuate;

marginal lobes dentate or serrate, irregular or

regular, 4-10 on each side, acute or obtuse,

0.9-1.5 mm deep; petioles 4-13 mm long, 11-

33% of lamina length. Lamina upper surface

indumentum erect or curved, eglandular,

0.4-1 mm long, sparse (hairs < 0.2 mm apart)

or moderately dense (hairs 0.1-0.2 mm apart),

sessile glands 8-64 mm 2 ; lower surface

indumentum erect or curved, eglandular,

0.4-07 mm long, confined to veins or

moderately dense (hairs 0.1-0.2 mm apart),

sessile glands 64-112 mm 2 ; transition from

leaves to floral bracts gradual. Floral bracts

16-46 mm long, 8-17 mm wide, consistently

exceeding verticils. Verticils (inflorescence

clusters) all separated on rachis, cymes once

dichasial at base then monochasial, with 3-7

flowers per monochasium, peduncles 1-6 mm

long on lowermost cluster; bracteoles obovate

or spathulate, 4-13 mm long, 0.7-5 mm wide.

Corolla tube longer than calyx, or same

length as calyx; annulus 2.2-3 mm from base

of corolla, annulus hairs 0.1-0.3 mm long;

upper lip ovate or elliptical, 4.1-6.2 mm long,

with glandular hairs on outer surface; lower

lip 6.2-77 mm long to end of lateral lobes,

14-15.5 mm long overall, glabrous or with

1-20 eglandular hairs on platform. Longest

stamens 13-15 mm long from base of corolla

tube; filament hairs 0.9-1.3 mm long, mainly

at distal end. Style 13-15 mm long; longer

stigma lobe 0.65-0.8 mm long, shorter stigma

lobe 0.45-0.5 mm long. Fruiting calyces

1-2 mm apart on rachilla; fruiting calyx

cylindrical, 7-10 mm long, 4.3-5.5 mm wide

at lobe apices, 17-2.5 times longer than wide,

exterior surface with hairs of different sizes

or types or with all hairs same size and type,

hairs eglandular, 0.4-1.9 mm long, sessile

glands 32-96 mm 2 ; lobes acute, 2-3 mm

long. Fruiting calyx fringe hairs about the

same length throughout, 0.15-0.3 mm long at

apical end, 0.15-0.25 mm long at sinus end,

sinus hairs absent, inner surface of tube with

dense ring of long hairs in medial section or

with sparse long hairs in medial section or

glabrous. Nutlets 2.1-2.5 mm long. Figs. 4F,

7E, 9H.

Additional selected specimens examined: Australia:

Queensland. Cook District: Alma-den, undated, Bick

176 (BRI); Springmount Station, c. 13 km from the

Mareeba - Dimbulah Road, on the road to Collins Weir,

Apr 1983, Clarkson 4664 (BRI, CNS, DNA, K, MEL,

MO, PERTH); 10 km from Alma-den on road to Mt

Surprise, Jan 1992, Forster PIF9593 (BRI); 16 km from

Mt Garnet on road to Lappa, 1 km past Mt Cardwell

turnoff, Jan 1993, Forster PIF12777 & Bean (BRI, CNS,

MEL); Pannikin Springs area, 29 km W of Mungana,

Jan 1993, Forster P1F13013 & Bean (BRI, DNA);

Copperfield River, Kidston Goldmine water supply dam,

Gilbert Range, Feb 1994, Forster PIF14902 & Bean

(BRI, CNS, MEL); Donkey Spring Creek, BulleringaNP,

80 km NW of Mt Surprise, Apr 1998, Forster PIF22414

& Booth (BRI); Stannary Hills, 11 km S of Mutchilba,

Portion 603, May 2006, Forster PIF31593 & McDonald

(BRI); Chillagoe, Jan 1931, Hubbard 6815 & Winders

(BRI); Kent Holdings, W of road from Mt Garnet to

Minnamoolka, May 1999, Jago 5239 & Wannan (BRI);

1 km from Walsh River crossing, N of Chillagoe, Mar

2000, McDonald KRM329 (BRI); near Barwidgi Road

turnoff, S of Mt Garnet, Apr 2004, McDonald KRM2176

& Covacevich (BRI); 51 km along Alma-den road from

junction with Gulf Development Road, near Mt Surprise,

May 2004, McDonald KRM2596 (BRI); Undara NP, 5.1

km along northern boundary firebreak with St Ronan’s

station, Dec 2006, McDonald KRM6032 (BRI); Gorge

Creek, Mareeba, Apr 1962, McKee 9257 (CANB);

Pinnacle Springs Road, 4 km W from Kennedy Highway,

Apr 2004, Purdie 5924 (BRI, CANB); 2 km downstream

of junction of Graves and Fulford Creeks, Burlington

Station, N of Mt Surprise, Jun 1999, Thompson SLT2467

& Newton (BRI); 100 mile swamps, ‘Rosella Plains’, Jul

1981, Williams 81134 (BRI). North Kennedy District:

12 km along Deadmans Road, off Silver Valley Road,

Feb 1996, Forster P1F18415 & Ryan (BRI, CNS, NSW);

c. 1.2 km ESE of Hidden Valley township, Paluma Road,

Feb 2000, Pollock ABP853 & Edginton (BRI).

Distribution and habitat : Anisomeles lappa

is endemic to Queensland. It is found in the

north-east of the state, including the western

parts of the Atherton Tableland, Bulleringa

NP, Alma-den, and Undara NP (Map 2).

It grows in eucalypt woodland. Almost all

records are from granite hills or sandy soils

adjacent to granite outcrops.

Phenology : Flowers and fruits are recorded

from December to July inclusive.

Notes: Anisomeles lappa can be distinguished

by the prostrate or procumbent habit, the

abundant patent hispid hairs on flowering

stems, the short cauline leaves, and the long

362

lower lip of the corolla. It may be distinguished

from the geographically close species A.

moschata by the prostrate to procumbent

habit, the floral bracts exceeding the verticils,

and the calyx fringe hairs only 0.15-0.25 mm

long at sinus end.

Conservation status : Least Concern.

Etymology : The epithet refers to the locality

of Lappa, a railway siding near the township

of Petford, near where the type was collected.

It is used as a noun in apposition.

16. Anisomeles leucotr icha A. R. Bean sp. nov.

foliis ellipticis praeditis tomento brevi albo,

verticillis omnino vel partim superpositis,

pilis marginalibus longis loborum calycis,

pilis 1-20 labio inferiori corollae insidentibus

et bracteis floralibus parvis distinguitur.

Typus: Australia: Northern Territory. 3.4 km

S of Goodparla airstrip (abandoned), Kakadu

National Park, 29 April 1990, A. V Slee & L.A.

Craven 3034 (holo: DNA; iso: CANB).

Erect or spreading shrub, 07-1.5 m high.

Upper stems and rachises without patent

hispid hairs; short curved hairs retrorse,

sparse or moderately dense; stalked glandular

hairs absent; sessile glands 16-112 mm 2

Cauline leaves 48-97 mm long, 13-43

mm wide, 2.2-37 times longer than wide,

base narrowly cuneate (< 60°) or attenuate;

marginal lobes crenate, regular, 15-37 on each

side, obtuse, 0.6-1.5 mm deep; petioles 10-26

mm long, 12-29% of lamina length. Lamina

upper surface indumentum erect or curved,

eglandular, 0.15-0.3 mm long, sparse (hairs

> 0.2 mm apart), sessile glands 8-32 mm 2 ;

lower surface indumentum erect or curved,

eglandular, 0.2-0.3 mm long, dense (hairs <

0.1 mm apart), sessile glands 64-112 mm 2 ;

transition from leaves to floral bracts abrupt.

Floral bracts elliptical, 3.5-25 mm long,

1-9 mm wide, not consistently exceeding

verticils. Verticils (inflorescence clusters) all

overlapping, forming continuous terminal

inflorescence or overlapping near apex, cymes

once dichasial at base then monochasial or

twice dichasial (± globose), with 2-9 flowers

per monochasium, peduncles 0-12 mm long

on lowermost cluster; bracteoles linear, 2.8-

4.5 mm long, 0.2-0.4 mm wide. Corolla tube

Austrobaileya 9(3): 321-381 (2015)

longer than calyx, or same length as calyx,

or shorter than calyx; annulus 2.5-3.2 mm

from base of corolla, annulus hairs 0.2-0.3

mm long; upper lip elliptical, 3.9-5.2 mm

long, with glandular hairs on outer surface

or with eglandular hairs on outer surface or

glabrous; lower lip 5.6-6.5 mm long to end

of lateral lobes, 8.8—12.3 mm long overall,

with 1-20 eglandular hairs on platform.

Longest stamens 11-12 mm long from base

of corolla tube; filament hairs 0.9-1.5 mm

long, mainly at distal end. Style 11-12 mm

long; longer stigma lobe 0.75-0.85 mm long,

shorter stigma lobe 0.4-0.5 mm long. Fruiting

calyces 0.8-1.2 mm apart on rachilla; fruiting

calyx cylindrical, 7.2-9.3 mm long, 3.1-3.6

mm wide at lobe apices, 2.1-2.9 times longer

than wide, exterior surface with all hairs same

size and type, hairs eglandular, 0.25-0.5 mm

long, sessile glands 80-128 mm 2 ; lobes acute,

2.5-37 mm long. Fruiting calyx fringe hairs

about the same length throughout, 07-1.4

mm long at apical end, 1-1.5 mm long at sinus

end, sinus hairs present, inner surface of tube

glabrous. Nutlets 1.9-2.3 mm long. Figs. 4G,

10A.

Additional selected specimens examined: Australia:

Northern Territory. Bukbuk Hill, near Arnhem

Highway, 4-5 km E of Kapalga turnoff, Feb 1991,

Brennan Bre995 (DNA); Kakadu NP, fire plot 202,

Jabiluka Outlier, Mar 1999, Brennan 3752 (DNA);

Litchfield Road, 2 miles [3 km] from HS, Apr 1967,

Byrnes N237 (DNA, NT); 3 miles [5 km] W of, and 8

miles [13 km] S of, Adelaide River township, Jan 1972,

Byrnes & McKean B259 (DNA); Kakadu Highway,

Nellie Creek Range, Mar 1987, Clark 882 (CANB,

DNA); Tipperary Station, Jun 1990, Clark 2347 (DNA);

13 miles [21 km] SE of Darwin, May 1958, Chippendale

4443 (BRI, DNA); 13 miles [21 km] W of Burrundie,

Mar 1961, Chippendale 7629 (DNA, MEL); Port

Essington, S of Wangewanga Cove, Apr 1993, Cowie

3356 (DNA); Port Essington, in 1819, Cunningham 270

(MEL); Melville Island, SE coast, Apr 1992, Fensham

1280 (DNA); Shoal Bay, 34 kmNE of Darwin, Apr 1992,

Halford Q1101 (BRI, DNA); Port Darwin, undated,

Holtze 54 (MEL); Mary River Station, Mar 2001, Liddle

2605 (DNA); Bathurst Island, Nguiu Forestry Suburb,

May 1998, Michel! & Risler 1520 (DNA); 0.5 miles [0.8

km] N of CSIRO block, Tipperary, May 1963, Muspratt

SSO610 (DNA); 2.5 miles [4.0 km] W of Mud Spring,

May 1963, Muspratt SS0542 (DNA); 10 miles [16 km]

out from Daly River, Feb 1964, Robinson R79 (DNA,

NT); Koolpinyah Station, 1 km S of Banka’s Jungle, Mar

1990, Taylor SMT12 (DNA); Jindare Station, Stray Creek

catchment, Jun 2010, Westaway 3251 (B, DNA, MO);

White Cliffs beach, west of airstrip, Croker Island, Mar

Bean, Revision of Anisomeles

363

2013, Westaway JOW4142 (BRI, CANB, DNA); Bynoe

Harbour, c. 200 m from MacKenzie Arm boat ramp off

Barramundi Drive, Mar 2006, Wirf247 (DNA).

Distribution and habitat: Anisomeles

leucotricha is endemic to the Northern

Territory, from Melville Island to Kakadu

NP and Litchfield NP (Map 3). It grows

on a variety of habitats, including vine

thicket on foreshore, open woodland with

Corymbia foelscheana (F.Muell.) K.D.Hill

& L. A. S. Johnson and/or Erythrophleum

chlorostachys (F.Muell.) Baill. on hills.

Substrate is quaternary sand or sandstone.

Phenology : Flowers are recorded from

January to July; fruits between April and July.

Notes: Anisomeles leucotricha is

distinguished by the elliptical leaves with an

attenuate base and a short even indumentum,

the exceptionally long calyx fringe hairs, the

closely arranged fruiting calyces (0.8-1.2

mm apart), the adjacent verticils sometimes

overlapping to form a dense cluster, and

the tiny floral bracts. The leaves have many

crenate lobes. The lower leaf surfaces have

short antrorse hairs, very often conspicuously

white, especially along the veins.

This species possibly intergrades with

A. brevipilosa to the east of (and west of)

Katherine, as some specimens from these

areas are difficult to allocate to either species.

Conservation status: Least Concern (IUCN

2012 ).

Etymology: The epithet is from the Greek

leucos - white, and trichos - hair or trichome.

In this species, the hairs on the underside of

the leaf are frequently conspicuously white,

especially along the veins.

17. Anisomeles macdonaldii A.R.Bean sp.

nov. abundantia pilorum glandularium et

absentia pilorum eglandularium retrorsorum

caulibus insidentibus, foliis basalibus latis

basi obtusa et petiolo longo (longitudine 32-

54% laminae aequante) praeditis distinguitur.

Typus: Australia: Queensland. Cook District:

c. 8 km by road E of Chillagoe, 21 April 2013,

K.R. McDonald KRM14083 & G.P. Guymer

(holo: BRI; iso: MEL, distribuendi).

Anisomeles salviifolia var. hispida Domin,

Biblioth. Bot. 89: 567 (1928). Type: Australia:

Queensland. Cook District: Smelling Bluff

near Chillagoe, February 1910, K. Domin

s.n. (syn: PR 530815); limestone hill near

Chillagoe, February 1910, K. Domin s.n. (syn:

PR 530816).

Erect or spreading shrub, 0.4-2 m high.

Upper stems and rachises with patent hispid

hairs; short curved hairs absent or retrorse,

sparse to moderately dense; stalked glandular

hairs abundant; glandular hairs extending

to lower stems; sessile glands 8-80 mm 2

Cauline leaves 51-91 mm long, 32-63 mm

wide, 1.2-1.9 times longer than wide, base

obtuse or broadly cuneate (> 60°); marginal

lobes crenate or dentate, regular, 10-18 on

each side, acute or obtuse, 0.9-2.4 mm deep;

petioles 19-40 mm long, 32-54% of lamina

length. Lamina upper surface indumentum

of erect glandular and curved eglandular

hairs, 0.15-1.3 mm long, moderately dense

(hairs 0.1-0.2 mm apart) or dense (hairs <

0.1 mm apart), sessile glands 8-48 mm 2 ;

lower surface indumentum of erect glandular

and curved eglandular hairs, 0.15-1.3 mm

long, moderately dense (hairs 0.1-0.2 mm

apart) or dense (hairs < 0.1 mm apart), sessile

glands 8-80 mm 2 ; transition from leaves to

floral bracts gradual. Floral bracts ovate or

broadly ovate, 14-47 mm long, 11-28 mm

wide, not consistently exceeding verticils or

consistently exceeding verticils. Verticils

(inflorescence clusters) all separated on

rachis, cymes entirely monochasial or once

dichasial at base then monochasial, with 4-11

flowers per monochasium, peduncles 2-28

mm long on lowermost cluster; bracteoles

spathulate or linear, 2.5-5.5 mm long, 0.4-1.1

mm wide. Corolla tube longer than calyx, or

same length as calyx; annulus 3-3.4 mm from

base of corolla, annulus hairs 0.2-0.3 mm

long; upper lip ovate or elliptical, 5.8-67 mm

long, with glandular hairs on outer surface

or with eglandular hairs on outer surface;

lower lip pink, 8.2-9 mm long to end of

lateral lobes, 14-16.5 mm long overall, with

1-20 or 20-100 eglandular hairs on platform.

Longest stamens 12.8-13.5 mm long from

base of corolla tube; filament hairs 1.3-1.6

mm long, mainly at distal end. Style 12.8-14

364

mm long; longer stigma lobe 0.65-0.75 mm

long, shorter stigma lobe 0.4-0.5 mm long.

Fruiting calyces 1-7 mm apart on rachilla;

fruiting calyx cylindrical, 7.4-10 mm long,

3-4.8 mm wide at lobe apices, 2.1-2.7 times

longer than wide, exterior surface with hairs

of different sizes or types, hairs glandular

and eglandular, 0.8-1.3 mm long, sessile

glands 8-48 mm 2 ; lobes acute, 1.8-3.2 mm

long. Fruiting calyx fringe hairs about the

same length throughout, 0.15-0.25 mm long

at apical end, 0.15-0.3 mm long at sinus end,

sinus hairs absent, inner surface of tube with

dense ring of long hairs in medial section or

with sparse long hairs in medial section or

glabrous. Nutlets 1.8-2.3 mm long. Figs. ID,

4H, 10B.

Additional specimens examined: Australia:

Queensland. Cook District: McDonald Creek area, 42

km from Mt Surprise township, Mt Surprise Gemfield,

Apr 1985, Champion 135 (BRI); Mt Mulligan, between

the abandoned mine site and the falls on Richards Creek

along the water supply pipeline, Apr 1984, Clarkson

5272 (BRI, MEL); Royal Arch Tower, c. 5 km SW of

Chillagoe, Mar 1987, Clarkson 6832 & McDonald

(BRI, DNA, L, NSW, PERTH); Near Royal Arch Cave,

Chillagoe, Jun 1992, Fensham 243 (BRI); on Chillagoe

Road, 14 km from Almaden, Mar 1990, Forster PIF6524

(BRI, CNS, DNA); Mt Pinnacle, SSW of Dimbulah, Jan

1993, Forster PIF12952 & Bean (BRI); Bridle Logging

Area, SF 607 Dinden, Jul 1995, Forster PIF17353 et al.

(BRI); NE slopes of Hann Tableland above the Southedge

HS, Jul 2002, Fox 1DF1648 (BRI); on a bench alongside

the water-supply pipeline between Ngarrabullgan and

the Mt Mulligan HS, Jul 2003, Fox IDF2072 (BRI);

Mungana, Aug 1995, Hyland 15355 (BRI); c. 1.9 km

ENE of Mt Carbine, c. 1.7 km along Mt Spurgeon

road, Apr 2013, Jensen 2801 & Kemp (BRI, CANB);

Pinchgut Hill, NE of Chillagoe, Mar 2005, McDonald

KRM3899 & Little (BRI); Chillagoe, Jan 1918, Michael

289 (BRI); Mt Mulligan, 40 km NW of Dimbulah, Apr

1989, Neldner 2758 (BRI). North Kennedy District:

NW of Pear Rock, Mt Stewart Range, Allandale’, May

1995, Forster PIF16624 & Figg (BRI); 9.5 km W of

Homestead, 83 km SW of Charters Towers, Aug 1992,

Thompson CHA196 & Sharpe (BRI).

Distribution and habitat: Anisomeles

macdonaldii is endemic to Queensland.

It is found in several disjunct populations

from Mt Carbine to Chillagoe, with disjunct

occurrences near Mt Surprise and Pentland.

It appears to be especially prevalent around

limestone outcrops near Chillagoe (Map 1).

It grows in low eucalypt woodland or on the

margins of vine thicket. The substrate can be

granite, limestone or sandstone.

Austrobaileya 9(3): 321-381 (2015)

Phenology : Flowers and fruits are recorded

from January to August inclusive.

Notes: Anisomeles macdonaldii is

distinguished by the broad cauline leaves,

the abundant stalked glandular hairs on the

stems, and the abundant patent hispid hairs on

the stems.

Conservation status: Least Concern.

Etymology: Named for Keith Raymond

McDonald (b. 1950), an expert on the

taxonomy and identification of many faunal

groups, especially frogs, and for the last

15 years, an avid plant collector. He has

increased our knowledge of Queensland plant

species considerably through his many high-

quality specimens, and has discovered or

rediscovered a number of species.

18. Anisomeles malabarica (L.) R.Br. in

Sims, Bot. Mag. 46: t. 2071 (1819); Nepeta

malabarica L., Mant. PI. Altera 566 (1771);

Ajuga fruticosa Roxb., Hort. Bengal. 44

(1814), nom. illeg .; Epimeredi malabaricus

(L.) Rothm., Repert. Spec. Nov. Regni Veg.

53: 12 (1944). Type: Herb. Linn. no. 726.26

(lecto: LINN^sfe Cramer 1981).

Nepeta pallida Salisb., Prodr. Stirp. Chap.

Allerton 78 (1796), nom. illeg. {Nepeta

malabarica L. cited in synonymy).

Stachys mauritianus Pers., Syn. PI. [Persoon]

2(1): 123 (1806); Craniotome mauritianum

(Pers.) Bojer, Hortus Maurit. 249 (1837).

Types: Isle de France, undated [June 1769],

P. Commerson s.n. (syn: P 00541421; P

00541422; P 04359606).

Erect or spreading shrub, 0.9-1.5 m high.

Upper stems and rachises without patent

hispid hairs; short curved hairs retrorse,

very dense, obscuring stem surface at x40

magnification; stalked glandular hairs absent.

Cauline leaves 47-90 mm long, 13.5-32

mm wide, 2.8-3.6 times longer than wide,

base narrowly cuneate (< 60°) or attenuate;

marginal lobes serrate, regular, 14-29 on

each side, acute or obtuse, 0.2-0.9 mm deep;

petioles 9-13 mm long, 14-21% of lamina

length. Lamina upper surface indumentum

erect or curved, eglandular, 0.4-1.2 mm long,

moderately dense (hairs 0.1-0.2 mm apart) or

Bean, Revision of Anisomeles

365

dense (hairs < 0.1 mm apart), sessile glands

8-48 mm 2 ; lower surface indumentum lanate,

tangled or erect or curved, eglandular, 0.9-1.2

mm long, very dense, obscuring surface at

x40 magnification; transition from leaves to

floral bracts abrupt. Floral bracts lanceolate

or elliptical, 6-17 mm long, 2-5 mm wide,

not consistently exceeding verticils. Verticils

(inflorescence clusters) all overlapping,

forming continuous terminal inflorescence or

overlapping near apex, cymes once dichasial

at base then monochasial or twice dichasial (±

globose), with 5-14 flowers per monochasium,

peduncles 0-1 mm long on lowermost cluster;

bracteoles linear, 4.2-6.5 mm long, 0.3-0.5

mm wide. Corolla tube shorter than calyx;

annulus 3.5-4.2 mm from base of corolla,

annulus hairs 0.1-0.15 mm long; upper lip

elliptical, 4.5-5.2 mm long, with eglandular

hairs on outer surface; lower lip 5-5.4 mm

long to end of lateral lobes, 8.5-10.7 mm

long overall, with 20-100 eglandular hairs

on platform. Longest stamens 13-14 mm

long from base of corolla tube; filament hairs

0.4-0.6 mm long, mainly along middle part.

Style 13-13.5 mm long; longer stigma lobe

0.7-1 mm long, shorter stigma lobe 0.3-0.5

mm long. Fruiting calyces 1.5-2.2 mm apart

on rachilla; fruiting calyx cylindrical, 8-12.2

mm long, 2.8-37 mm wide at lobe apices,

3-4.3 times longer than wide, exterior surface

with hairs of different sizes or types, hairs

glandular and eglandular, 1.2-1.5 mm long,

sessile glands 8-16 mm 2 ; lobes attenuate,

3.1-5.3 mm long. Fruiting calyx fringe hairs

about the same length throughout or longer

at sinus end than at apical end, 0.3-0.6 mm

long at apical end, 0.5-0.7 mm long at sinus

end, sinus hairs present, inner surface of

tube with dense ring of long hairs in medial

section. Nutlets 1.9-2.2 mm long. Figs. 41,

10C. Malabar catmint.

Additional specimens examined : India. San Thome, in

1856, Cleghorn s.n. (E); Kulasegaram, S. Travancore,

Feb 1934, Erlanson 5378 (NY); Pondicherry, May

1837, Gaudichand s.n. (P); Cuddalore, Nov 1959,

Govindarajalu 3699 (L); Bounds of Vedanthangal

Sanctuary, Tamil Nadu, Jan 1976, Henry 47027

(MH); Tiruchi, Nov 1978, Matthew et al. RHT19392

(L); Banavar - Arsikere road, Jan 1970, Saldanha

16079 (E); Kanjamalai, Apr 1944, Sinclair 3464 (E);

Papanasam project area, Madras State, Jun 1964,

Subramanian 1653 (L); Perur, Noyal riverbank, Jun

1956, Subramanyam 45 (MH); way to Pachakumatchi,

Jun 1959, Subramanyam 8259 (L); Susindram, Tamil

Nadu, Dec 1980, Swaminathan 68985 (MH); Walayar

Dam, outlet canal, Kerala State, May 1964, Vajravelu

19064 (MH); Peninsula Ind. orientalis, undated, Wight

2173 (NY); Pulicat, Madras, Mar 1837, Wight 2173 (E).

Sri Lanka. Kaddaikadu, Mar 1973, Bernardi 14265 (L);

beside Trincomalee - Kinniya Road, Sep 1974, Cramer

4325 (E); Palaiyutta, Mar 1994, Cramer 6960 et al.

(E); 9 miles [14 km] NNE of Jaffa, along road to Palai,

Dec 1974, Davidse 9090 & Sumithraarachchi (K, L);

Trincomalee, Oct 1976, Fosberg 56390 (E, NY, P); Foul

Point, Trincomalee district, Feb 1972, Jayasuriya 651 et

al. (E); Talankuda, Batticaloa district, Apr 1973, Stone

11188 (L); Just N of Trincomalee, Mar 1973, Townsend

73/245 (E); Kinniyai, Trincomalee district, Sep 1974,

Waas & Tirvengadum 809 (L, NY). Malaysia. Penang,

in 1822, Wallich 2037/2 (NY). Western Indian Ocean.

Mauritius, undated, Sieber s.n. (NY).

Distribution and habitat : Anisomeles

malabarica is native to Sri Lanka and

southern India (Map 6). In addition, it is (or

was) naturalised on Penang Island, Malaysia,

and on the island of Mauritius. It inhabits

open sunny areas; sandy flats on dunes near

the coast, stream banks, and “waste ground”.

Phenology : Flowers are recorded for every

month of the year; fruits are recorded in

January, April, September and December.

Nomenclature : The authorship fox Anisomeles

malabarica has often been given as “(L.) R.Br.

ex Sims”, but the notation ‘Brown mss’ after

the description in the protologue indicates

that the author was Brown and not Sims.

Hence the correct authorship is “(L.) R.Br. in

Sims” or, where the plant name stands alone,

just “(L.) R.Br”.

Notes: Anisomeles malabarica is a very

distinctive species because of its very densely

tomentose stems, relatively narrow leaves,

long attenuate calyx lobes and long hairs on

the external surface of the calyx.

Baker (1877) accepted A. malabarica as

being native on Mauritius, but it seems far

more likely that it was transported there by

man, as specimens of the Mauritian plant are

identical to those from India, and the species

has no means of long-range seed dispersal.

366

Conservation status : Least Concern.

19. Anisomeles moschata R.Br., Prodr. FI.

Nov. Holl. 503 (1810); Anisomeles salviifolia

var. moschata (R.Br.) Domin, Biblioth. Bot.

89: 567 (1928); Epimeredi moschatus (R.Br.)

Rothm., Repert. Spec. Nov. Regni Veg. 53: 12

(1944). Type: [Australia: Queensland. Port

Curtis District:] Keppel Bay, 9 August 1802,

R. Brown [Bennett Number 2357] (lecto: BM

001041069 [here designated]; isolecto: CANB

278977, E 00649579, P 04358989).

Anisomeles salviifolia var. subtomentosa

Domin, Biblioth. Bot. 89: 567 (1928), pro

parte. Type: Australia: Queensland. [?North

Kennedy District]: Rockingham Bay,

undated,/. Dallachys.n. (syn: K).

Procumbent, erect or spreading shrub,

0.3-2.5 m high. Upper stems and rachises

without patent hispid hairs; short curved hairs

retrorse, sparse to dense; stalked glandular

hairs absent; sessile glands 8-112 mm 2

Cauline leaves 11-100 mm long, 6-43 mm

wide, 1.7-2.8 times longer than wide, base

obtuse or broadly cuneate (> 60°) or narrowly

cuneate (< 60°) or attenuate; marginal lobes

crenate, dentate or serrate, irregular or

regular, 3-18 on each side, acute or obtuse,

0.3-2 mm deep; petioles 4-22 mm long, 21-

38% of lamina length. Lamina upper surface

indumentum erect or curved, eglandular,

0.25-0.7 mm long, sparse (hairs > 0.2 mm

apart) or moderately dense (hairs 0.1-0.2 mm

apart), sessile glands 8-96 mm 2 ; lower surface

indumentum erect or curved, eglandular,

0.25-0.6 mm long, moderately dense (hairs

0.1-0.2 mm apart) or dense (hairs <0.1 mm

apart), sessile glands 48-160 mm 2 ; transition

from leaves to floral bracts abrupt, or gradual.

Floral bracts elliptical or ovate, 8-22 mm long,

3-9 mm wide, not consistently exceeding

verticils or consistently exceeding verticils.

Verticils (inflorescence clusters) all separated

on rachis, cymes entirely monochasial or once

dichasial at base then monochasial or twice

dichasial (± globose), with 2-10 flowers per

monochasium, peduncles 0-11 mm long on

lowermost cluster; bracteoles spathulate or

linear, 2.8-47 mm long, 0.2-1.1 mm wide.

Corolla tube longer than calyx, or same

length as calyx, or shorter than calyx; annulus

Austrobaileya 9(3): 321-381 (2015)

2.3-3.5 mm from base of corolla, annulus

hairs 0.15-0.3 mm long; upper lip ovate or

elliptical, 3.6-5.3 mm long, with glandular

hairs on outer surface or with eglandular hairs

on outer surface; lower lip purple, 47-6.6

mm long to end of lateral lobes, 8.6-12 mm

long overall, glabrous on platform. Longest

stamens 10.5-12.5 mm long from base of

corolla tube; filament hairs 1.1-1.9 mm long,

mainly at distal end. Style 10.5-14 mm long;

longer stigma lobe 0.55-0.7 mm long, shorter

stigma lobe 0.3-0.45 mm long. Fruiting

calyces 1.2-2.8 mm apart on rachilla; fruiting

calyx cylindrical, 6.2-8.6 mm long, 2.5-4.5

mm wide at lobe apices, 1.7-27 times longer

than wide, exterior surface with all hairs same

size and type, hairs eglandular, 0.2-0.6 mm

long, sessile glands 64-176 mm 2 ; lobes acute,

1.5-2.5 mm long. Fruiting calyx fringe hairs

longer at sinus end than at apical end, 0.2-0.4

mm long at apical end, 0.5-1.1 mm long at

sinus end, sinus hairs absent, inner surface of

tube with dense ring of long hairs in medial

section or with sparse long hairs in medial

section or glabrous. Nutlets 1.8-2.1 mm long.

Figs. 2B, 4J, 5A, 5B, 7A, 7B, 7C, 7D, 7F,

7H, 10D.

Additional selected specimens examined : Australia:

Queensland. Cook District: between Cairns and

Kuranda, May 1952, Everist 5139 (BRI, CANB);

Daintree NP, Orania Creek off Little Daintree River

head. May 1998, Forster PIF22858 et al. (AD, BRI,

CNS, MEL); Luster Creek, Aug 1995, Jago 3571 (BRI,

MEL); Cairns, Koombal, Jun 1993, Lyons 142 (BRI);

Mt Misery Road, Turnoff, S of Shiptons Flat, Jul 2001,

McDonaldKRM938 (BRI). North Kennedy District: Mt

Abbot, 50 km W of Bowen, Aug 1992, Bean 4833 (BRI);

Adjacent to Proserpine Dam, c. 25 km W of Proserpine,

Jul 2007, Bean 26462 (BRI); Sinclair Bay, Cape

Gloucester, Jul 1992, Batianoff 9209206 (BRI); NW of

Pentland, near ‘Lowholm’, Jul 1954, Blake 19380 (BRI,

CANB, MEL); SF 605 Koombooloomba, Tully Falls,

Mar 2002, Forster PIF28444 (BRI, K, MEL, NSW).

South Kennedy District: St Bees Island, 38 km NE of

Mackay, Mar 1989, Batianoff 11152 (AD, BRI, PNH);

Rabbit Island, 6 km N of Seaforth, Jun 1994, Batianoff

940683 & Price (BRI). Leichhardt District: slopes of

Ropers Peak, NE of Capella, May 1987, Bean 577 (BRI);

Ropers Peak, E of Capella, Jan 2006, Fensham 5361 &

Butler (BRI); eastern slopes of Dilly Pinnacle, 7 km

NNE of Springsure, Oct 1998, Bean 14179 (BRI, NSW);

Consuelo Tableland, upper tributary of Rocky Creek,

Mar 2006, Eddie CPE1511 (BRI, NSW). Port Curtis

District: Mt Slopeaway, 5 km from Marlborough, May

1991, Batianoff MS9105121 & Franks (BRI); Mt Archer,

Rockhampton, Feb 1980, Stanley 592 (BRI); Middle

Bean, Revision of Anisomeles

367

Percy Island, 87 miles [140 km] SE of Mackay, Apr

1956, Lazarides 5604 (BRI, CANB, PERTH); 5 miles

[8 km] NE of Tynan HS, Jun 1963, Lazarides 6889

(BRI, CANB, L). Wide Bay District: Biggenden, May

1931, White 7716 (BRI). Moreton District: SW end of

Macleay Island, Jun 1976, Elsol 9 (BRI); Mt Greville,

c. 20 km W of Boonah, Apr 1975, Sharpe 1211 & Saul

(BRI); Wilson’s Peak, Apr 1949, White 13006 (BRI,

CANB).

Distribution and habitat : Anisomeles

moschata is endemic to Queensland where

it is widely distributed in eastern areas,

from just north of Cooktown to within a few

kilometres of the New South Wales border,

and extending up to 400 km from the east

coast (Map 3). It inhabits eucalypt forests and

woodlands on wide range of topographies and

geologies, where the soils are well drained

and frosts are absent or few.

Phenology : Flowers and fruits have been

recorded from every month of the year.

Notes : This is a very widespread and variable

species. Specimens have been assigned to this

species on the basis of their glabrous corolla

platform, the calyx fringe hairs longer at the

sinus end than at the apical end, the petioles

21-38% of lamina length, and the moderately

dense to dense tomentum on the lower leaf

surface. Three or four regional forms exist

- for instance, many specimens from the

Leichhardt pastoral district of Queensland

are consistently small-leaved and have

verticils with short monochasia; specimens

from Cairns and surrounding areas have

large leaves with numerous marginal lobes.

Because of apparently intergrading forms and

lack of data on floral characteristics, I have

not erected any new species from A. moschata

sens. lat. However, it seems highly likely that

such a division will be possible if further

studies are undertaken.

The syntypes of Anisomeles salviifolia

var. subtomentosa at K comprise a mixture

of A. moschata and A. dallachyi. It is the

K specimens that Domin examined before

describing that variety (Orchard 1999); he did

not see material at MEL or P.

Conservation status : Least Concern.

20. Anisomeles ornans A.R.Bean sp. nov.

affinis A. eriodi sed foliis praeditis lobis

marginalibus paucioribus, floribus paucioribus

in monochasiis, praesentia pilorum in labio

inferiore corollae et calycibus fructificantibus

brevioribus differens. Typus: Australia:

Queensland. Cook District: Simpson’s Gully,

Agate Creek fossicking area, 30 April 2006,

K.R. McDonald KRM5243 (holo: BRI; iso:

CNS, MEL, distribuendi).

Erect or spreading shrub, 0.3-0.6 m high.

Upper stems and rachises without patent

hispid hairs; short curved hairs no fixed

direction, very dense, obscuring stem surface

at x40 magnification; stalked glandular hairs

absent. Cauline leaves 29-68 mm long, 11-

32 mm wide, 2-3.6 times longer than wide,

base narrowly cuneate (< 60°) or attenuate;

marginal lobes dentate or serrate, irregular

or regular, 7-14 on each side, acute or obtuse,

0.5-1 mm deep; petioles 6-13 mm long, 16-

27 % of lamina length. Lamina upper surface

indumentum lanate, tangled, 0.4-0.6 mm

long, dense (hairs <0.1 mm apart), sessile

glands 16-48 mm 2 ; lower surface indumentum

lanate, tangled, 0.4-0.7 mm long, dense (hairs

< 0.1 mm apart) or very dense, obscuring

surface at x40 magnification; transition from

leaves to floral bracts abrupt. Floral bracts

lanceolate or elliptical, 6-26 mm long, 2-8

mm wide, not consistently exceeding verticils.

Verticils (inflorescence clusters) all separated

on rachis, cymes entirely monochasial or once

dichasial at base then monochasial, with 3-6

flowers per monochasium, peduncles 0-2

mm long on lowermost cluster; bracteoles

linear, 2-6.6 mm long, 0.2-0.4 mm wide.

Corolla tube same length as calyx; annulus

2-2.5 mm from base of corolla, annulus

hairs 0.25-0.4 mm long; upper lip elliptical,

4.3-5 mm long, with glandular hairs on outer

surface; lower lip purple or pink, 6.3-77 mm

long to end of lateral lobes, 11-13.6 mm long

overall, with 1-20 or 20-100 eglandular hairs

on platform. Longest stamens 10.5-11.5 mm

long from base of corolla tube; filament hairs

0.6-0.9 mm long, mainly at distal end. Style

11.5-12 mm long; longer stigma lobe 0.7-0.8

mm long, shorter stigma lobe 0.45-0.55 mm

long. Fruiting calyces 1-1.9 mm apart on

rachilla; fruiting calyx narrowly campanulate

or cylindrical, 6.7-8.1 mm long, 2.4-3.9 mm

wide at lobe apices, 2.1-2.8 times longer than

368

wide, exterior surface with all hairs same size

and type, hairs eglandular, 0.5-0.9 mm long;

lobes acute, 2-2.5 mm long. Fruiting calyx

fringe hairs longer at sinus end than at apical

end, 0.2-0.4 mm long at apical end, 0.7-1.1

mm long at sinus end, sinus hairs absent,

inner surface of tube with dense ring of long

hairs in medial section. Nutlets 2-2.1 mm

long. Figs. 5C, 10E.

Additional specimens examined: Australia:

Queensland. Burke District: The Crater, Blackbraes

NP, N of Hughenden, Apr 2002, Bean 18853 (BRI). Cook

District: Cave Creek, Gilbert River, undated, Daintree

s.n. (MEL 684767); Chadshunt, Gilbert River, May

1954, Everist 5401 (BRI, MEL); Flat Creek, c. 50 km S

of Georgetown, May 2011, Mathieson MTM1076 (BRI,

NSW); 5 km along Blacksoil Creek Road, Agate Creek

fossicking area, Apr 2006, McDonald KRM5202 (BRI,

NSW); Rungulla Station HS area, S of Georgetown,

Sep 2013, McDonald KRM14725 & Little (BRI, CANB);

Gilbert River Flolding, about 300 m along track, Apr

2014, McDonald KRM15524 (BRI, CANB); Gilbert

River, Feb 1922, White 1425 (BRI). North Kennedy

District: c. 25 km N of Wando Vale HS, Broken River,

Poley Cow Creek, Apr 1988, FellDF882 (BRI); Gregory

Springs Station, N of Hughenden, Feb 1931, Hubbard

7710 & Winders (BRI, K).

Distribution and habitat: Anisomeles ornans

is endemic to Queensland. It is found from

‘Wando Vale’ (N of Pentland) to Gilbert River

(E of Croydon) (Map 7). It grows on sandstone

escarpments and basalt hills, with a variety

of species including Eucalyptus chartaboma

D.Nicolle and Corymbia erythrophloia

(Blakely) K.D.Hill & L.A.S.Johnson. Soils

may be clayey, sandy or skeletal.

Phenology: Flowers recorded from February

to September; fruits from April to September.

Notes: Anisomeles ornans is allied to A.

eriodes, but differs by the leaves with fewer

marginal lobes (7—14), the more widely

separated verticils, the fewer flowers (3-6)

in the monochasia, the corolla tube the same

length as the calyx; the corolla platform with

1-20 hairs, and the shorter fruiting calyces.

Conservation status: Least Concern.

Etymology: Derived from the Latin ornatus

meaning ornate or embellished. It refers to the

pleasing appearance of the silvery stems and

leaves in this species.

Austrobaileya 9(3): 321-381 (2015)

21. Anisomeles papuana A.R.Bean sp. nov.

foliis lobis marginalibus praeditis, labio infero

breviore corollae, calycibus fructificantibus

brevissimis et nuculis tantum 1.5-1.7 mm

longis distinguitur. Typus: Papua New

Guinea. Western Province: Near Morehead

Patrol Post, 30 August 1967, R. Pullen 7199

(holo: BRI; iso: A, CANB, K n.v., L, LAE

n.v).

Erect or spreading shrub, 1.2-2 m high. Upper

stems and rachises without patent hispid hairs;

short curved hairs retrorse, moderately dense

or dense; stalked glandular hairs absent; sessile

glands 8-64 mm 2 Cauline leaves 62-112 mm

long, 26-60 mm wide, 1.8-2.8 times longer

than wide, base obtuse or broadly cuneate (>

60°) or narrowly cuneate (< 60°) or attenuate;

marginal lobes crenate or dentate, irregular or

regular, 19-33 on each side, acute or obtuse,

0.7-3.5 mm deep; petioles 8-24 mm long, 13-

25% of lamina length. Lamina upper surface

indumentum erect or curved, eglandular, 0.3-

0.6 mm long, sparse (hairs > 0.2 mm apart)

or moderately dense (hairs 0.1-0.2 mm apart),

sessile glands 8-48 mm 2 ; lower surface

indumentum erect or curved, eglandular, 0.3-

0.6 mm long, moderately dense (hairs 0.1-0.2

mm apart) or dense (hairs < 0.1 mm apart),

sessile glands 16-144 mm 2 ; transition from

leaves to floral bracts abrupt. Floral bracts

elliptical or ovate, 4.5-14 mm long, 3-5 mm

wide, not consistently exceeding verticils.

Verticils (inflorescence clusters) overlapping

near apex or all separated on rachis, cymes

once dichasial at base then monochasial or

twice dichasial (± globose), with 5-14 flowers

per monochasium, peduncles 0-5 mm long

on lowermost cluster; bracteoles spathulate or

linear, 1.8-4.2 mm long, 0.15-0.45 mm wide.

Corolla tube same length as calyx; annulus

2.1-3 mm from base of corolla, annulus hairs

0.15-0.25 mm long; upper lip elliptical, 4-4.2

mm long, with eglandular hairs on outer

surface; lower lip 5-5.9 mm long to end of

lateral lobes, 7.5-9.3 mm long overall, with

20-100 eglandular hairs on platform. Longest

stamens 10.5-11.5 mm long from base of

corolla tube; filament hairs 1-1.4 mm long,

mainly at distal end. Style 10-12 mm long;

longer stigma lobe 0.45-0.7 mm long, shorter

stigma lobe 0.25-0.4 mm long. Fruiting

Bean, Revision of Anisomeles

369

calyces 1-1.6 mm apart on rachilla; fruiting

calyx cylindrical, 6.2-8.3 mm long, 3.1-3.5

mm wide at lobe apices, 2.1-2.5 times longer

than wide, exterior surface with all hairs same

size and type, hairs eglandular, 0.2-0.6 mm

long, sessile glands 8-160 mm 2 ; lobes acute,

1.4-2.7 mm long. Fruiting calyx fringe hairs

longer at sinus end than at apical end, 0.1-

0.25 mm long at apical end, 0.45-1.1 mm long

at sinus end, sinus hairs absent, inner surface

of tube with dense ring of long hairs in medial

section or glabrous. Nutlets 1.5-1.7 mm long.

Figs. 5D, 10F.

Additional specimens examined: Indonesia. Moluccas:

Yamdena Island, road from Saumlakki to Olilit, Mar

1938, Buwalda 4043 (L). Papua New Guinea. Central

Province: Baroka, Mekeo district, Apr 1933, Brass 3715

(BRI); Roger’s airstrip, c. 8 miles [13 km] W of Kanosia

Plantation, Jul 1962, Darbyshire 653 (BRI, CANB,

L); Nebiri Quarry, Port Moresby sub-district, Apr

1970, Gebo 369 (CANB); near Jackson’s Airport, Port

Moresby, Aug 1963, Heyligers 1006 (CANB); Tovobada

Hills, 12 miles [19 km] N of Port Moresby, May 1965,

Heyligers 1180 (BRI, CANB, L); S coast near Kwikila,

Abau subdistrict, Jun 1969, Paijmans 767 (CANB);

Rubulogo Creek, c. 18 miles [29 km] N of Port Moresby,

Apr 1967, Pullen 6689 (CANB); north vicinity of Rigo,

Aug 1962, Schodde 2700 (BRI, CANB); Port Moresby,

Jul 1918, White 61 (BRI). Gulf Province: Near Malalaua,

Mar 1966, Craven & Schodde 904 (CANB); Malalaua,

lower Tauri River area, Feb 1966, Pullen 6528 (CANB).

Morobe Province: Bulolo, Feb 1950, Fryar 3948 (BRI,

CANB); Bulolo, Wau subdistrict, Jul 1970, Streimann

& Kairo NGF47869 (BRI, CANB). Northern Province:

Between Dabora and Wabubu, Cape Vogel Peninsula,

Apr 1953, Brass 21876 (CANB). Western Highlands

Province: Baiyer River, Nov 1954, Floyd & Womersley

6826 (BRI). Western Province: Near Rouku, Morehead

subdistrict, Jul 1973, Henty NGF49713 (BRI, CANB,

L). Australia: Queensland. Cook District: Thursday

Island, Jun 1897, Bailey s.n. (BRI [AQ160154]); Darnley

Island, Torres Strait, Apr 2005, Hacks LAH285 (BRI,

CANB); Yorke Island, Torres Strait, Oct 1971, Lawrie

s.n. (BRI [AQ3927]); Mabuyag Island, Torres Strait, Aug

2008, McKenna SGM222 (BRI, CANB); Gabba island,

Torres Strait, Feb 2000, Waterhouse BMW5655 (BRI,

CANB).

Distribution and habitat: Anisomeles

papuana is widely distributed in lowland New

Guinea, also Torres Strait, Queensland, and

the southern Moluccas Islands of Indonesia

(Map 5). It inhabits savannah woodland at

altitudes from 0-100 metres and mostly below

30m (except the three specimens mentioned

below).

Phenology: Flowers and fruits are recorded

from February to November inclusive.

Notes: Anisomeles papuana is distinguished

by the many lobes per leaf, the relatively short

petioles, the consistently short cylindrical

calyces and the nutlets only 1.5-1.7 mm long.

Three specimens {Floyd & Womersley

6826, Fryar 3948 , Streimann & Kairo

NGF47869 ) are known from relatively high

altitude (from 750 to 1150 metres) in Papua

New Guinea. These specimens have a more

compact infructescence, and the floral bracts

exceed the verticils; they may represent an

additional undescribed taxon.

Conservation status : Least Concern.

Etymology : The epithet refers to Papua

New Guinea, where the great majority of

collections have been made.

22. Anisomeles principis A.R.Bean sp.

nov. pilis antrorsis retrorsisve plusminusve

patentibusve, eglandularibus, densis in

caulibus, bracteolis latis, pilis 1-20 labio

inferiori corollae insidentibus et pilis

marginalibus brevibus loborum calycis

distinguitur. Typus: Australia: Western

Australia. South-west Osborne Island,

Bonaparte Archipelago, 29 June 1973, P.G.

Wilson 11152 (holo: PERTH).

Erect or spreading shrub, 0.5-1.5 m high.

Upper stems and rachises without patent

hispid hairs, or with patent hispid hairs; short

curved hairs retrorse or antrorse, sparse to

dense; stalked glandular hairs absent; sessile

glands 16-160 mm 2 Cauline leaves 63-104

mm long, 24-42 mm wide, 2-2.7 times

longer than wide, base obtuse or broadly

cuneate (> 60°) or narrowly cuneate (< 60°) or

attenuate; marginal lobes crenate or serrate,

regular, 10-21 on each side, acute or obtuse,

0.9-2.1 mm deep; petioles 16-19 mm long,

18-25% of lamina length. Lamina upper

surface indumentum erect, glandular or erect

or curved, eglandular, 0.25-0.8 mm long,

sparse (hairs > 0.2 mm apart) or moderately

dense (hairs 0.1-0.2 mm apart), sessile glands

16-80 mm 2 ; lower surface indumentum erect,

glandular or erect or curved, eglandular,

0.25-0.7 mm long, moderately dense (hairs

370

0.1-0.2 mm apart) or dense (hairs < 0.1 mm

apart), sessile glands 48-176 mm 2 ; transition

from leaves to floral bracts abrupt, or gradual.

Floral bracts elliptical or ovate, 7-30 mm

long, 3.5-18 mm wide, not consistently

exceeding verticils. Verticils (inflorescence

clusters) all separated on rachis, cymes

entirely monochasial or once dichasial at

base then monochasial, with 5-14 flowers

per monochasium, peduncles 0-15 mm long

on lowermost cluster; bracteoles obovate or

spathulate or linear, 4.8-7.8 mm long, 0.5-2.5

mm wide. Corolla tube longer than calyx, or

same length as calyx; annulus 2.5-3 mm from

base of corolla, annulus hairs 0.2-0.3 mm

long; upper lip elliptical, 3.6-47 mm long,

with eglandular hairs on outer surface; lower

lip 4-5.8 mm long to end of lateral lobes,

8-10.5 mm long overall, with 1-20 eglandular

hairs on platform. Longest stamens 12-14 mm

long from base of corolla tube; filament hairs

07-1.2 mm long, mainly at distal end. Style

12-14 mm long; longer stigma lobe 0.65-0.7

mm long, shorter stigma lobe 0.4-0.55 mm

long. Fruiting calyces 0.8-1.5 mm apart on

rachilla; fruiting calyx cylindrical, 8.2-11

mm long, 3.4-4.8 mm wide at lobe apices,

2-2.6 times longer than wide, exterior surface

with hairs of different sizes or types or with

all hairs same size and type, hairs glandular

or hairs eglandular, 0.4-1 mm long, sessile

glands 80-144 mm 2 ; lobes acute, 2.3-3 mm

long. Fruiting calyx fringe hairs about the

same length throughout, 0.15-0.25 mm long

at apical end, 0.15-0.25 mm long at sinus end,

sinus hairs absent, inner surface of tube with

dense ring of long hairs in medial section or

glabrous. Nutlets 1.9-2.3 mm long. Figs. 5E,

10G

Additional specimens examined : Indonesia. Lesser

Sunda Islands: Timor, undated, herb. Drake (P); Timor,

Apr 1821, Reinwardt s.n. (L). Australia: Western

Australia. 8.5 km ENE of Mt Agnes, near headwaters

on N side of Prince Regent River, Jan 2001, Barrett &

O ’Connor RLB1588 (PERTH); Secure Bay, side arm on

W side of bay, Yampi Peninsula, Mar 2001, Barrett &

Handasyde RLB2047 (PERTH); Ellenbrae Station, Jul

2004, Brennan & Done 6328 (DNA, PERTH); slopes

overlooking campsite, Margaret Island, Buccaneer

Archipelago, Jun 1982, Hopkins BA0475 (PERTH);

Boongarie Island, 18.3 km NE of Mt Knight, Jun 1988,

Keighery & Alford 1817 (PERTH); E side of Mindjau

Creek, Port Warrender, Admiralty Gulf, Jan 1982,

Kenneally 7764 (PERTH); Campsite on unnamed

Austrobaileya 9(3): 321-381 (2015)

tributary of Prince Regent River, arising 19 km SE of the

mouth, north-west Kimberley, Jun 1984, Kenneally 8922

(MEL, PERTH); unnamed island in Prince Frederick

Harbour at entrance to Hunter River, Jun 1984, Kenneally

8978 (PERTH); junction of unnamed creek and Sale

River, 30 km ESE of mouth. May 1986, Kenneally

9603 (MEL, PERTH); Heywood Island (South island),

Naturalist Island, E side, directly opposite ‘Ninepin’,

or the Sentinel, Prince Frederick Harbour, May 1987,

Kenneally 9963 (PERTH); Boomerang Bay, Bigge

Island, Jun 1972, Marchant 72/60 (PERTH); adjacent

to creek running out of Koolan townsite. May 1993,

Mitchell 3112 (PERTH); c. 5 km SW of Crystal Head on E

side of major river and adjacent to W.A. Water Authority

gauging station. Mar 1994, Mitchell 3403 (PERTH);

junction of two creeks where G. Grey had his main base

camp in 1837, 15° 22’ 04”S 124° 45’ 19”E, Mar 1994,

Mitchell 3465 (NSW, PERTH); beach on Hanover Bay,

north Kimberley, May 1998, Mitchell 5401 (PERTH);

Bachsten Creek, Jul 2001, Riissell-Smith & Handasyde

TH01-171 (DNA, PERTH); Mallam Spring, SE side of

Augustus Island, Jul 1990, Willing 187 (PERTH); above

Moran River Gorge, Prince Regent Nature Reserve, May

1991, Willing 420 (PERTH); Bonaparte Archipelago,

May 1972, Wilson 10896 (CANB, PERTH).

Distribution and habitat : Anisomeles

principis is found in the Kimberley region of

Western Australia and on the island of Timor

(Map 5). In Australia, it reportedly grows

on sandstone hills and gullies, in Triodia

hummock grassland, eucalypt woodland or on

the edges of vine thicket. Two specimen labels

mention basalt as the substrate.

Phenology : Flowers collected from January

to July; fruits from May to July.

Notes: Anisomeles principis is variable

with regard to indumentum types present

on the stems. Several specimens (including

Kenneally 8922, Wilson 10896) have dense

antrorse hairs, and one {Kenneally 9603 )

has abundant spreading hispid hairs. These

may prove to be taxonomically distinct.

Distinguishing features of the species as

currently circumscribed are: monochasial or

once-dichasial cymes of the verticils; short

petioles (16-19 mm); short calyx fringe hairs,

often pedunculate lower verticils; crowded

flowers and fruits; and broad bracteoles.

Bean, Revision of Anisomeles

371

Conservation status : Least Concern (IUCN

2012 ).

Etymology : From the Latin principis - of

the prince. This is in reference to the Prince

Regent River, where the distribution of this

species is centred.

23. Anisomeles salviifolia R.Br., Prodr. FI.

Nov. Holl:. 503 (1810); Anisomeles salviifolia

var. typica Domin, Biblioth. Bot. 89: 567

(1928), nom. illeg .; Epimeredi salviifolius

(R.Br.) Rothm., Repert. Spec. Nov. Regni Veg.

53: 12 (1944). Type: [Australia: Northern

Territory] Maria Island, Gulf of Carpentaria,

1 January 1803, R. Brown s.n. [Bennett

Number 2355] (lecto [here designated]: BM

001041064; isolecto: BM 001041065; CANB

278978).

Erect or spreading shrub, 0.8-2 m high.

Upper stems and rachises without patent

hispid hairs; short curved hairs retrorse or

antrorse or no fixed direction, very dense,

obscuring stem surface at *40 magnification;

stalked glandular hairs absent. Cauline

leaves 39-100 mm long, 11-36 mm wide,

1.7-5.4 times longer than wide, base obtuse or

broadly cuneate (> 60°) or narrowly cuneate

(< 60°) or attenuate; marginal lobes crenate

or serrate, irregular or regular, 12-20 on

each side, acute or obtuse, 0.3-1.8 mm deep;

petioles 9-19 mm long, 14-40% of lamina

length. Lamina upper surface indumentum

erect or curved, eglandular, 0.3-0.6 mm long,

moderately dense (hairs 0.1-0.2 mm apart) or

dense (hairs < 0.1 mm apart), sessile glands

8-48 mm 2 ; lower surface indumentum lanate,

tangled, 0.5-0.8 mm long, dense (hairs < 0.1

mm apart) or very dense, obscuring surface

at x40 magnification, sessile glands 16-64

mm 2 ; transition from leaves to floral bracts

abrupt. Floral bracts elliptical or ovate, 7-25

mm long, 2-12 mm wide, not consistently

exceeding verticils. Verticils (inflorescence

clusters) overlapping near apex or all

separated on rachis, cymes once dichasial at

base then monochasial or twice dichasial (±

globose), with 4-7 flowers per monochasium,

peduncles 0-12 mm long on lowermost cluster;

bracteoles spathulate, 4.8-7.5 mm long, 0.9-

1.2 mm wide. Corolla tube longer than calyx,

or same length as calyx; annulus 3.3-3.4 mm

from base of corolla, annulus hairs 0.2-0.3

mm long; upper lip ovate or elliptical, 4.2-4.8

mm long, with glandular or eglandular hairs

on outer surface; lower lip 5.1-6.1 mm long to

end of lateral lobes, 9.7-11 mm long overall,

with 20-100 eglandular hairs on platform.

Longest stamens 10.5-12 mm long from base

of corolla tube; filament hairs 0.9-1.1 mm

long, mainly at distal end. Style 10.5-11.5 mm

long; longer stigma lobe 0.75-0.9 mm long,

shorter stigma lobe 0.4-0.5 mm long. Fruiting

calyces 1.5-2 mm apart on rachilla; fruiting

calyx cylindrical, 7.2-10.5 mm long, 3.3-4.8

mm wide at lobe apices, 1.6-2.5 times longer

than wide, exterior surface with all hairs same

size and type, hairs eglandular, 0.5-0.7 mm

long, sessile glands 16-80 mm 2 ; lobes acute,

1.8-2.9 mm long. Fruiting calyx fringe hairs

about the same length throughout, 0.5-1 mm

long at apical end (or 0.15-0.3), 0.6-1 mm

long at sinus end (or 0.15-0.3), sinus hairs

absent, inner surface of tube with dense ring

of long hairs in medial section or with sparse

long hairs in medial section. Nutlets 2-2.4

mm long. Figs. 5F, 10H.

Additional specimens examined : Australia: Northern

Territory, west side of South West Island, Sir Edward

Pellew group, Feb 1976, Craven 3725 (BRI, CANB,

DNA); Maria Island, Jul 1972, Dunlop 2747 (DNA,

NT); Bing Bong Station, Jun 1971, Henry 94 (BRI,

DNA, NSW); 6 km SSE of Lake Eames, Vanderlin

Island, Aug 1988, Latz 11067 (DNA, NT); Buchanan

Bay, May 1977, McKey 125 (DNA, NT); point at the

N end of Eagle Bay, Maria Island, Mar 2008, Mitchell

8789 (CANB, DNA, NSW); northern end. West Island,

Jul 1988, Parsons 63 (DNA, NT); Camp Beach, Centre

Island, Jun 2010, Randell 821 (DNA, K, NT); South West

Island, Sir Edward Pellew group, Feb 1976, Rice 2392

(CANB); Watson Island, Sir Edward Pellew group, Jul

1988, Thomson 2536 (DNA, NT); South West Island, Sir

Edward Pellew group, Jul 1984, Wightman 1577 (CANB,

DNA).

Distribution and habitat : Anisomeles

salviifolia is endemic to the Northern

Territory. It is confined to the south-western

Gulf of Carpentaria, including Maria Island,

the islands of the Sir Edward Pellew group,

and on the mainland at Bing Bong Station

(Map 2). It grows in dune swales never far

from the ocean, usually with grasses and

woodland species, but sometimes on the edge

of vine thicket. Soils are sandy and infertile.

372

Phenology : Flowers are recorded from

February to August; fruits are recorded from

May to August.

Notes : Specimens of Anisomeles salviifolia

seen from Maria Island differ consistently

from those collected in the Sir Edward Pellew

islands or Bing Bong station by the narrower

leaves. Two collections from Maria Island

have long calyx fringe hairs, contrasting with

the short calyx fringe hairs in specimens

from other locations. However, the type of

A. sahiifolia has short calyx fringe hairs.

Because the calyx fringe hairs difference

is not consistent and only leaf shape seems

consistently different, no taxonomic

distinction has been made.

Conservation status : Least Concern.

24. Anisomeles viscidula A.R.Bean sp.

nov. pilis glandularibus abundantibus et

hispidis sparsis usque crebris insidentibus

caulibus, foliis comparate angustis et carpis

late dispositis in cymis distinguitur. Typus:

Australia: Western Australia. Cave Spring

Gap on Kununurra to Legune road, 26 April

1977, H. Eichler 22501 (holo: DNA; iso:

CANB, L, MEL, MO, PERTH).

Procumbent shrub, or erect or spreading

shrub, 0.45-1.5 m high. Upper stems and

rachises with patent hispid hairs; short curved

hairs absent or retrorse, sparse to moderately

dense; stalked glandular hairs abundant, or

occasional; glandular hairs extending to lower

stems; sessile glands 8-48 mm 2 Cauline

leaves 71-110 mm long, 19-42 mm wide,

2.4-3.8 times longer than wide, base obtuse or

broadly cuneate (> 60°) or narrowly cuneate

(< 60°) or attenuate; marginal lobes serrate,

irregular or regular, 15-30 on each side, acute

or obtuse, 0.6-1 mm deep; petioles 14-43 mm

long, 19-43% of lamina length. Lamina upper

surface indumentum of erect glandular and

curved eglandular hairs, 0.1-0.7 mm long,

moderately dense (hairs 0.1-0.2 mm apart) or

dense (hairs < 0.1 mm apart), sessile glands

8-48 mm 2 ; lower surface indumentum of erect

glandular and curved eglandular hairs, 0.25-

0.4 mm long, moderately dense (hairs 0.1-0.2

mm apart) or dense (hairs < 0.1 mm apart),

sessile glands 8-128 mm 2 ; transition from

Austrobaileya 9(3): 321-381 (2015)

leaves to floral bracts abrupt or gradual. Floral

bracts lanceolate or elliptical, 5-44 mm long,

1-16 mm wide, not consistently exceeding

verticils or consistently exceeding verticils.

Verticils (inflorescence clusters) all separated

on rachis, cymes entirely monochasial or once

dichasial at base then monochasial, with 3-11

flowers per monochasium, peduncles 0-13

mm long on lowermost cluster; bracteoles

spathulate or linear, 4-7.5 mm long, 0.4-1.2

mm wide. Corolla tube longer than calyx, or

same length as calyx; annulus 2.5-3.4 mm

from base of corolla, annulus hairs 0.2-0.25

mm long; upper lip elliptical, 3.8-57 mm

long, with eglandular hairs on outer surface;

lower lip 7-9 mm long to end of lateral

lobes, 13.5—16 mm long overall, with 1-20

eglandular hairs on platform. Longest stamens

12.5- 14 mm long from base of corolla tube;

filament hairs 0.6-1.2 mm long, mainly at

distal end. Style 11.5-13.5 mm long; longer

stigma lobe 0.7-0.85 mm long, shorter stigma

lobe 0.5-0.65 mm long. Fruiting calyces

1.5- 4.8 mm apart on rachilla; fruiting calyx

cylindrical, 8.9-12.5 mm long, 3.5-5.2 mm

wide at lobe apices, 1.9-2.6 times longer than

wide, exterior surface with hairs of different

sizes or types, hairs glandular and eglandular,

longest hairs 0.4-1.3 mm long, sessile glands

8-128 mm 2 ; lobes acute, 2.3-37 mm long.

Fruiting calyx fringe hairs about the same

length throughout, 0.15-0.4 mm long at

apical end, 0.15-0.4 mm long at sinus end,

sinus hairs absent, inner surface of tube with

dense ring of long hairs in medial section or

with sparse long hairs in medial section or

glabrous. Nutlets 1.9-2.2 mm long. Figs. 2C,

5G, 11A

Additional specimens examined : Australia: Western

Australia. SW side of Mt Page, S of Walcott Inlet,

Yampi Peninsula, Mar 2001, Barrett & Start RLB2339

(PERTH); Point Springs, 40 km N of Kununurra, May

1988, Kenneally 10692 (NSW, PERTH); head of Tin Can

Gully, Mornington Peninsula, May 2005, Legge 976

(PERTH); 2 km E of junction of Charnley and Calder

Rivers, Eastern Walcott Inlet, May 1983, Milewski 205

(PERTH); Foot of Cave Spring Range, 30 km NNE

of Kununurra, Mar 1978, Paijmans 2543 (CANB).

Northern Territory. 10 km from Jim Jim Falls, Sep

1984, Brennan 2675 (DNA); Katherine Gorge NP, Apr

1968, Byrnes NB618 (BRI, DNA); c. 450 m from Top Car

Park, on track to Upper Falls, Edith Falls, Nitmiluk NP,

May 1993, Conn 3703 & Doust (DNA, NSW); McArthur

River area, near the Glyde River, Jan 1976, Craven 3567

Bean, Revision of Anisomeles

373

(BRI, CANB, DNA); Mt McMinns Station, 11.8 km E of

station turnoff on road to Roper Bar, Mar 2002, Dixon

& Harwood 1016 (DNA); Fitzmaurice River basin. May

1994, Dunlop 9922 & Latz (DNA, MEL); Keep River NP,

Jarrarm area, Jun 1995, Egan 5038 (DNA); NitmilukNP,

Feb 1991, Evans 3639 (BRI, DNA, K); Wearyan River

headwaters, Jan 1989, Latz 11262 (DNA, MEL, NT); 28

km S of Nathan River HS, Sep 1995, Latz 14580 (DNA,

NT); 2.5 km ESE of Katherine Gorge ranger station. May

2009, Latz 24344 & Quarmby (DNA, NT); Yamburran

Range, 10 km NNE of Mt Millikmonmir, May 1994,

Leach 4555 & Albrecht (DNA, MEL); Nitmiluk NP, Feb

2001, Michell 3348 (B, DNA); Hayes Creek, May 1963,

Muspratt SS0603 (DNA); Edith Falls, Katherine Gorge

NP, Apr 1983, Thompson 249 (CANB); Bullo River

Station, Mar 2009, Westaway 2792 (DNA, MO).

Distribution and habitat : Anisomeles

viscidula is endemic to Australia and is

distributed from the north-west Kimberley

region of Western Australia, to the north¬

eastern Northern Territory (Map 7). It grows

in open woodland dominated by species such

as Erythrophleum chlorostachys. Eucalyptus

miniata A.Cunn. ex Schauer or E. phoenicea

F.Muell. It is apparently confined to quartzose

sandstone plateaux and outcrops on sandy or

skeletal soil.

Phenology : Flowers are recorded from

January to June. There is also a single

flowering record from September, but that

specimen was collected from a spring with

permanent water supply. Fruits are recorded

from April to September.

Notes: Anisomeles viscidula is characterised

by the relatively lax inflorescences exceeded

by the floral bracts on the lower verticils, the

relatively narrow leaves, and the presence of

stalked glandular hairs and hispid hairs on

the stems and leaves. Eastern populations

(adjacent to the Gulf of Carpentaria) have

relatively few glandular hairs, but in all other

populations, they are abundant. It differs from

A. macdonaldii by the narrower cauline leaves

with serrate marginal lobes, and the mostly

longer fruiting calyces.

Conservation status : Least Concern.

Etymology: From the diminutive form of

the Latin word viscidus meaning sticky or

clammy. This is in reference to the densely

glandular leaves and stems of this species.

25. Anisomeles vulpina A.R.Bean sp. nov.

pilis densis retrorsis in caule, foliis latis basi

obtusa, in pagina superiore folii pilis 1.0-1.3

mm longitudine et pilis marginalibus loborum

calycis ubique 0.3-0.6 mm longitudine

distinguitur. Typus: Australia: Queensland.

North Kennedy District: c. 50 metres from

the top of Mt Fox crater, SW of Ingham, 4

October 2014, R. Jensen 3350 & J.E. Kemp

(holo: BRI; iso: CANB, CNS, E, MEL, NSW,

NY, distribuendi).

Erect or spreading shrub, 0.3-0.4 m high.

Upper stems and rachises without patent

hispid hairs; short curved hairs retrorse,

moderately dense or dense; stalked glandular

hairs absent; sessile glands 8-48 mm 2

Cauline leaves 27-50 mm long, 16-30 mm

wide, 1.6-2.1 times longer than wide, base

obtuse or broadly cuneate (> 60°) or narrowly

cuneate (< 60°) or attenuate; marginal lobes

crenate or dentate or serrate, regular, 9-14 on

each side, acute or obtuse, 0.4-1.8 mm deep;

petioles 3.5-20 mm long, 13-44% of lamina

length. Lamina upper surface indumentum

erect or curved, eglandular, 0.9-1.3 mm long,

moderately dense (hairs 0.1-0.2 mm apart) or

dense (hairs < 0.1 mm apart), sessile glands

8-32 mm 2 ; lower surface indumentum lanate,

tangled or erect or curved, eglandular, 1-1.2

mm long, dense (hairs < 0.1 mm apart), sessile

glands 16-64 mm 2 ; transition from leaves to

floral bracts gradual. Floral bracts elliptical

or ovate or broadly ovate, 4.5-20 mm long,

3-14 mm wide, not consistently exceeding

verticils. Verticils (inflorescence clusters)

overlapping near apex or all separated on

rachis, cymes entirely monochasial, with 3-6

flowers per monochasium, peduncles 0-1

mm long on lowermost cluster; bracteoles

spathulate or linear, 2.4-3.5 mm long, 0.3-0.5

mm wide. Corolla tube same length as calyx,

or shorter than calyx; annulus 2.7-3 mm from

base of corolla, annulus hairs 0.25-0.4 mm

long; upper lip elliptical, 3.4-4.9 mm long,

with eglandular hairs on outer surface; lower

lip purple or pink, 5.3-6.4 mm long to end

of lateral lobes, 9.5-12.1 mm long overall,

glabrous or with 1-20 eglandular hairs on

platform. Longest stamens 11.5-12 mm long

from base of corolla tube; filament hairs

0.9-1.9 mm long, mainly at distal end. Style

374

12-13 mm long; longer stigma lobe 0.6-0.65

mm long, shorter stigma lobe 0.4-0.45 mm

long. Fruiting calyces 0.9-1.5 mm apart on

rachilla; fruiting calyx cylindrical, 7.3-8 mm

long, 3.8-4.4 mm wide at lobe apices, 1.7-2.1

times longer than wide, exterior surface with

all hairs same size and type, hairs eglandular,

0.6-1 mm long, sessile glands 64-112 mm 2 ;

lobes acute, 1.8-2.4 mm long. Fruiting calyx

fringe hairs about the same length throughout,

0.3-0.5 mm long at apical end, 0.3-0.5 mm

long at sinus end, sinus hairs absent, inner

surface of tube with dense ring of long hairs

in medial section or with sparse long hairs

in medial section. Nutlets 1.9-2.1 mm long.

Figs. 5H, 11B.

Additional specimens examined : Australia:

Queensland. North Kennedy District: Mt Fox crater,

Seaview Range, Apr 1985, Rodd 4462 & Hardie (BRI,

NSW).

Distribution and habitat : Anisomeles

vulpina is endemic to Queensland where it is

known only from Mt Fox, a volcanic crater

south-west of Ingham (Map 5). It grows in

fertile volcanic soil amongst basalt rocks and

boulders, in an open grassland habitat.

Phenology : Flowers are recorded in April and

October; fruits in October.

Notes : Anisomeles vulpina differs from A.

moschata by the much longer hairs on both

the upper and lower leaf surfaces, and by the

shorter calyx fringe hairs.

Conservation status : A status of Vulnerable,

criteria D1 and D2, is recommended.

Etymology : From the Latin vulpes, meaning

“fox”. This is in reference to the type locality,

Mount Fox.

26. Anisomeles xerophila A.R.Bean sp.

nov. absentia pilorum glandularium, pagina

interiore labii inferioris corollae longa,

calycibus fructificantibus brevibus (5.2-7.1

mm longitudine) et pilis folii 0.3-0.5 mm

longitudine distinguitur. Typus: Australia:

Northern Territory. North Hayward Creek on

Stuart Highway, 58 km N of Tennant Creek,

19 April 1983, R.M. Barker 186 (holo: DNA;

iso: AD).

Austrobaileya 9(3): 321-381 (2015)

Erect or spreading shrub, 0.3-1.5 m high.

Upper stems and rachises with or without

patent hispid hairs; short curved hairs

retrorse, moderately dense or dense; stalked

glandular hairs absent; sessile glands 48-96

mm 2 . Cauline leaves 24-78 mm long, 9-23

mm wide, 2.1-3.9 times longer than wide,

base narrowly cuneate (< 60°) or attenuate;

marginal lobes crenate or dentate, irregular

or regular, 7-17 on each side, acute or

obtuse, 0.4-2.1 mm deep; petioles 3.5-11

mm long, 13-22% of lamina length. Lamina

upper surface indumentum erect or curved,

eglandular, 0.25-0.4 mm long, sparse (hairs >

0.2 mm apart) or moderately dense (hairs 0.1-

0.2 mm apart), sessile glands 32-112 mm 2 ;

lower surface indumentum erect or curved,

eglandular, 0.25-0.4 mm long, moderately

dense (hairs 0.1-0.2 mm apart) or dense (hairs

< 0.1 mm apart), sessile glands 16-128 mm 2 ;

transition from leaves to floral bracts abrupt.

Floral bracts lanceolate or elliptical, 6-27

mm long, 1.5-7 mm wide, not consistently

exceeding verticils. Verticils (inflorescence

clusters) all separated on rachis, cymes once

dichasial at base then monochasial or twice

dichasial (± globose), with 3-7 flowers per

monochasium, peduncles 0-12 mm long on

lowermost cluster; bracteoles spathulate or

linear, 2.8-5 mm long, 0.3-0.8 mm wide.

Corolla tube same length as calyx; annulus

2.3-3 mm from base of corolla, annulus

hairs 0.25-0.3 mm long; upper lip ovate or

elliptical, 4-5.1 mm long, with glandular or

eglandular hairs on outer surface; lower lip

4-6 mm long to end of lateral lobes, 6.5-10.3

mm long overall, with 20-100 eglandular hairs

on platform. Longest stamens 10-12.8 mm

long from base of corolla tube; filament hairs

0.7-1.3 mm long, mainly at distal end. Style

10.5-13 mm long; longer stigma lobe 0.65-0.7

mm long, shorter stigma lobe 0.45-0.6 mm

long. Fruiting calyces 1.2-2.2 mm apart on

rachilla; fruiting calyx cylindrical, 5.8-7.9

mm long, 2.4-3.9 mm wide at lobe apices,

1.8-2.9 times longer than wide, exterior

surface with all hairs same size and type, hairs

eglandular, 0.3-0.5 mm long, sessile glands

128-160 mm 2 ; lobes acute, 1.7-2.4 mm long.

Fruiting calyx fringe hairs longer at sinus end

than at apical end, 0.1-0.3 mm long at apical

Bean, Revision of Anisomeles

375

end, 0.7-1 mm long at sinus end, sinus hairs

absent, inner surface of tube with dense ring

of long hairs in medial section. Nutlets 2-2.2

mm long. Figs. 5J, 11C.

Additional specimens examined : Australia: Northern

Territory. 2 miles [3 km] N of Katherine, Apr 1967,

Adams 1755 (CANB, L, NT); Attack Creek, W of Stuart

Hwy, c. 66 km N of Tennant Creek, May 1996, Albrecht

7633 & Latz (NT); ‘Nickson’s Place’, across river from

Katherine, Apr 1956, Burbidge 5111 (BRI, CANB,

PERTH); Seven Emu Station, 34.3 km (by road) from

junction with main road to Wollogorang, May 1993,

Conn 3767 & Doust (DNA, NSW); Robinson River

Station, May 1983, Cowen 6 (CANB, DNA); Settlement

Creek (below jump-up) on road between Redbank Mine

and Wollogorang, May 1984, Halford 845123 (NT); c. 30

miles [48 km] E of Borroloola, Jun 1973, Henry 836 (NT);

Whittington Range, 70 km N of Tennant Creek, Oct

2006, Latz & Albrecht 22238 (MEL, NT); Wollogorang

Station, Branch Creek gorge, Jul 1998, Michell & Risler

1640 (DNA); Nitmiluk NP, Mar 2001, Michell 3349 &

Deichmann (DNA); Gibson Creek, 35 miles [56 km] N

of Tennant Creek, Jul 1968, Must 209 (MEL, NT); 15

miles [24 km] S of McArthur River station, Jul 1948,

Perry 1735 (MEL, NT, PERTH); Hayward Creek, W

of highway, Phillip Creek Station, Feb 1984, Strong 45

(NT); Attack Creek, Banka Banka station, Jul 1983,

Thomson 353 (NT). Queensland. Burke District:

Cloncurry, Nov 1935, Blake 10133 (BRI); Westmoreland

Station, just W of Hells Gate, 15 km SE of HS, May

2005, Booth 4293 (BRI, NSW); Norfolk Station, on

remote station track, Jun 2006, Booth 4619 & Kelman

(BRI); Cabbage Tree Creek, Little Eva crossing, Apr

1962, Cole 223 & Provan (BRI); Westmoreland, Lagoon

Creek, off track to Camp Ridgeway, May 1997, Forster

PIF21036 & Booth (BRI); 49 km from Mt Isa on Duchess

road. May 1997, Forster P1F21172 & Booth (BRI, DNA,

MEL); 12 km S of Mt Isa, on road to Duchess, Jun 1991,

Halford Q519 (BRI); Lawn Hill Station, Jul 1971, Latz

1607 (CANB, MEL, NSW, NT); 2 miles [3 km] S of Mt

Isa township. Mar 1954, Lazarides 4377 (BRI, CANB,

MEL, NT); Mt Isa, Aug 1928, MacGillivray 2213 (BRI).

Distribution and habitat: Anisomeles

xerophila is widely distributed from Katherine

south to Tennant Creek in the Northern

Territory and to Cloncurry in north-western

Queensland (Map 1). In the more inland

areas, it is confined to creek banks, but closer

to the coast, it inhabits eucalypt woodland on

flats or sandstone scree slopes; occasionally it

sometimes grows in limestone areas.

Phenology: Flowers mostly recorded from

January to July, but with a single record from

November; fruits from January to October.

Notes: Anisomeles xerophila is closely related

to A. leucotricha , but differs by the leaves

broadest towards the base, leaves with fewer

marginal lobes, the longer hairs of the lower

leaf surface, the well separated verticils

(verticils overlapping for A. leucotricha ),

and the more widely spaced fruiting calyces.

A. xerophila is also closely related to A.

brevipilosa , and possibly intergrades with it in

the Wollogorang - Borroloola area, but differs

by the mostly broader leaves, the longer hairs

on the upper and lower leaf surfaces, and the

shorter lower lip of the corolla.

This species usually has verticils that

are short and compact, twice dichasially

branched, but occasionally (e.g. Must 209,

Booth 4293, Burbidge 5111 ) they are once-

dichasial and elongate with a rachis up to

25mm long.

Conservation status: Least Concern.

Etymology: From the Greek xeros (dry) and

philus (loving). This species alone extends

to the semi-arid parts of Australia, where the

annual rainfall is about 450 mm.

Excluded names

Anisomeles australis Spreng., Syst. Veg. (ed.

16) [Sprengel] 4(2, Cur. Post.) 226 (1827). =

Teucrium corymbosum R.Br.

Anisomeles furcata (Link) Sweet, Hort. Brit.

[Loudon] 232 (1830). = Craniotome furcata

(Link) Kuntze.

Anisomeles glabrata Benth. ex Wall., Numer.

List n. 2041 (1829), nomen nudum.

Anisomeles indica f. albiflora Kuntze, Rev is.

Gen. PI. 2: 512 (1891), nomen nudum.

Anisomeles indica f. rubicunda Kuntze,

Revis. Gen. PI. 2: 512 (1891), nomen nudum.

Anisomeles intermedia Wight ex Benth.,

Labiat. Gen. Spec. 703 (1835). Type: India.

Madurai [Madura], undated, R. Wight 2172/42

(lecto: MM, fide Cramer (1981), n.v., isolecto:

E, K).

Bentham (1835) speculated that Anisomeles

intermedia could be a hybrid between^, ovata

(= A. indica) and A. malabarica, and I think

that this must be the case. Its morphological

characteristics are intermediate between

A. indica and A. malabarica, and both

376

of the original localities (Madura, India

and Peradeniya, Sri Lanka) are within the

geographical overlap zone of A. indica and A.

malabarica. Of the more than 200 Indian and

Sri Lankan Anisomeles specimens seen by the

present author, none matches the type of A.

intermedia , further supporting the idea that it

is an infrequent hybrid. Aluri (1992) reported

the existence of a single plant of a presumed

hybrid between these two species, occurring

at Turimella, India.

Anisomeles nepalensis Spreng., Syst. Veg. (ed.

16) [Sprengel] 2: 706 (1825). = Craniotome

furcata (Link) Kuntze

Anisomeles secunda f. albiflora Kuntze,

Revis. Gen. PI. 2: 512 (1891), nomen nudum.

Anisomeles secunda f. rubicunda Kuntze,

Revis. Gen. PI. 2: 512 (1891), nomen nudum.

Acknowledgements

I am grateful to the Directors of A, BKF,

BM, CANB, DNA, E, G, GH, K, L, MEL,

MH, NT, P, PERTH, PR, SING and W for the

loan of specimens or provision of specimen

images. Sincere thanks to Odile Poncy (P)

who undertook a prolonged and painstaking

search for the original material of Epimeredi.

My thanks are due to Keith McDonald

for his numerous excellent herbarium

collections, including spirit material, and

his photographic images of flowers and plant

habit; Rigel Jensen and Jeanette Kemp for

their collection of A. vulpina ; Ian Cowie for

sending photographs of wild Anisomeles ;

John McNeill for nomenclatural advice; Peter

Bostock for helping to interpret some difficult

Latin phrases, and for providing the Latin

diagnoses; Jian Wang for interpreting labels

of some Chinese specimens; Gordon Guymer

for his support and encouragement; and Will

Smith for the many illustrations and digital

images.

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Map 1. Distribution of Anisomeles spp. A. ajugacea ▼, A. dallachyi ▲, A. macdonaldii ★, A. xerophila •. National

Parks and other conservation reserves in grey.

Bean, Revision of Anisomeles

379

Map 2. Distribution of Anisomeles spp. A. antrorsa A. bundeyensis T, A. farinacea •, A. languida M,A. lappa •,

A. salviifolia ▲. National Parks and other conservation reserves in grey.

Map 3. Distribution of Anisomeles spp. A. brevipilosa ▲. A. leucotricha ■. A. moschata •. National Parks and other

conservation reserves in grey.

380

Austrobaileya 9(3): 321-381 (2015)

Map 4. Distribution of Anisomeles spp. A. candicans A, A. indica •.

Map 5. Distribution of Anisomeles spp. A. carpentarica •, A. eriodes A, A. papuana X, A. principis T-T,

A. vulpina M. National Parks and other conservation reserves in grey.

Bean, Revision of Anisomeles

381

Map 6. Distribution of Anisomeles spp. A. heyneana A, A. malabarica •.

Map 7. Distribution of Anisomeles spp. A. grandibractea w, A. inodora •, A. ornans ■ . A. viscidula A. National

Parks and other conservation reserves in grey.

Three new species of Taeniophyllum Blume

(Orchidaceae) from northern Queensland

B. Gray

Summary

Gray, B. (2015). Three new species of Taeniophyllum Blume (Orchidaceae) from northern Queensland.

Austrobaileya 9(3): 382-392. Three diminutive, leafless orchids of the genus Taeniophyllum are

described as new from northern Queensland, increasing the total species enumerated from mainland

Australia to eight, namely T. clementsii (D.L.Jones & B.Gray) Kocyan & Schuit., T. confertum

B.Gray & D.L.Jones, T. epacridicola B.Gray sp. nov., T. explanatum B.Gray sp. nov., T. lobatum

Dockrill, T. malianum Schltr., T. muelleri Lindl. ex Benth. and T. triquetroradix B.Gray sp. nov. A

key to the mainland Australian Taeniophyllum species is provided. All inflorescences of the mainland

Australian taxa are also illustrated and provided here for comparison purposes and compliment the

key. Line drawings, photographs and distribution maps for the three newly described species are

given.

Key Words: Orchidaceae, Taeniophyllum , Taeniophyllum epacridicola , Taeniophyllum explanatum ,

Taeniophyllum triquetroradix , Australia flora, Queensland flora, new species, taxonomy,

identification key

B. Gray, Australian Tropical Herbarium, James Cook University, Cairns Campus, McGregor Road,

Smithfield, Queensland 4878, Australia.

Introduction

Taeniophyllum Blume is a diverse group of

small, epiphytic orchids with 229 species

widely distributed from Africa through

India to Japan southwards through the Malay

Archipelago to New Guinea, Australia and

extending east to New Caledonia and Tahiti,

with the centre of distribution in New Guinea

(Schlechter 1982; Seidenfaden & Wood

1992; Margonska & Szlachetko 2010). In

Australia, six species are enumerated, namely

T. confertum B.Gray & D.L. Jones, T. lobatum

Dockrill, T. malianum Schltr., T. muelleri

Lindl. ex Benth., T. norfolkianum D.L. Jones,

B.Gray & M.A.Clements and T. pusillum

(Willd.) Seidenf. & Ormerod (formerly T.

obtusum Blume; see Seidenfaden 1995)

(Gray & Jones 1984; Lavarack & Gray 1985;

Clements 1989; Dockrill 1992; Ormerod 1994;

Jones 2006; Jones et al. 2006). However, two

of the six species recorded occur on off shore

islands, namely T. norfolkianum on Norfolk

Island (Jones et al. 2006) and T. pusillum on

Accepted for publication 24 August 2015

Christmas Island (Clements 1989) and are not

included in this paper.

Based on recent molecular findings,

Kocyan & Schuiteman (2014) combined

Microtatorchis Schltr. with Taeniophyllum ,

and in doing so, Microtatorchis clementsii

D.L.Jones & B.Gray was transferred to

Taeniophyllum clementsii (D.L. Jones &

B.Gray) Kocyan & Schuit., bringing the

number of Taeniophyllum species recognised

for mainland Australia up to five.

Generally, all the Australian

Taeniophyllum species are small, leafless or

with minute scale like leaves covering the

stem. They have diminutive flowers, usually

bearing only one, rarely two, at a time on a

peduncle. The inflorescence is a raceme, often

with continuous growth, producing flowers

over a long period.

Observations have been carried out by the

author over a period of some years, and have

led to the discovery of these three novelties

described here as T. epacridicola B.Gray sp.

nov., T. explanatum B.Gray sp. nov. and T.

triquetroradix B.Gray sp. nov. Two of these,

Gray, New species of Taeniophyllum from Queensland

namely T. explanatum and T. triquetroradix,

occur within the Wet Tropics of north-eastern

Queensland (Townsville to Cooktown),

while T. epacridicola is known only from far

northern Cape York Peninsula. All three new

species are known only from a small number

of collections.

Materials and methods

This study is based on examination of living

plants in the field, cultivated material, and

preserved spirit collections deposited in

BRI, CANB and CNS (herbaria acronyms

383

follow Thiers (continuously updated)).

Measurements and illustrations were made

based on living root and stem morphology,

whereas inflorescence structure and floral

characters were studied from live plants and

preserved materials in spirit. Length x width

measurements are indicated as measurement

x measurement mm.

An illustration depicting the inflorescences

of all mainland Australian Taeniophyllum

taxa, including the three newtaxa, is provided

here for comparison purposes (Fig. 1) and to

compliment the key.

Taxonomy

Key to mainland Australian Taeniophyllum species

1 Sepals and petals fused near the base forming a tube.2

1. Sepals and petals free to the base not forming a tube.5

2 Roots terete in cross section.T. muelleri

2. Roots triangular or flattened in cross section.3

3 Roots triangular in cross section (having a raised longitudinal ridge) . . T. triquetroradix

3. Roots flattened in cross section .4

4 Roots 1.5-2 mm broad; peduncle filiform, 12-25 mm long;

rachis filiform; floral bracts small, alternating, c. 0.5 mm apart,

all in one plane; flowers c. 2.5 mm long.T. explanatum

4. Roots 1 mm or less broad; peduncle filiform, 2-5 mm long; rachis not

filiform, fleshy, parallel sided, twice as wide as peduncle; floral

bracts alternating, <0.5 mm apart; flowers < 2 mm long.T. clementsii

5 Roots 2-3 mm broad; peduncle 2-3 mm long; floral bracts overlapping,

hiding the rachis.T. confertum

5. Roots 1.5-4 mm broad; peduncle and rachis not as above.6

6 Peduncle, rachis and ovary sparsely covered with erect short-bristly

hairs; flowers green, turning yellow with age.T. lobatum

6. Peduncle, rachis and ovary glabrous.7

7 Young roots green with rows of white elongate spots; roots 1.5-2.5 mm

broad, mostly hanging free from the host, some appressed; peduncle

filiform, 20-50(-60) mm long; floral bracts overlapping; flower 7-11 mm

wide.T. malianum

7. Young roots green; roots 2-3.5(-4) mm broad, appressed to the host;

peduncle and rachis zig-zag from base, not filiform, 5-10 mm long;

floral bracts alternating 1-1.5 mm apart; flower 4.5-5 mm

wide

T. epacridicola

384

Austrobaileya 9(3): 382-392 (2015)

10mm

E

Fig. 1 . Lateral views of inflorescences from all mainland Australian Taeniophyllum species. A. T. malianum

cultivated plant (no voucher). B. T. confertum from Walker s.n. (CNS). C. T. muelleri from Gray BG9661 (CNS). D.

T. lobatum from Gray BG8584 (CNS). E. T. triquetroradix from Gray BG4129 (CNS). F. T. epacridicola from Gray

BG5235 (CNS). G. T. explanatum from Gray BG9674 (CNS). H. T. clementsii from Gray BG8508 (CNS). Scale as

indicated. Del. B. Gray.

Gray, New species of Taeniophyllum from Queensland

1. Taeniophyllum epacridicola B.Gray

sp. nov. Similar to T. mangifera Schltr. but

differs in having dorsal sepal and petals

clavate, lateral sepals obovate with a sharp

apical point and a much larger stigmatic

opening. Typus: Queensland. Cook District:

Atherton, cultivated (ex situ from Wasp

Creek), 11 September 1990, B. Gray BG5235

(holo: BRI; iso: CNS).

Plant epiphytic. Roots usually several,

flattened, creeping, 50-150 mm long, 2-3.5(-

4) mm broad, greyish green. Inflorescences

one to several, scape 4-6 mm long. Rachis

rough, zig-zag, 5-10 mm long, reddish-green.

Floral bracts alternating triangular 1-1.5 mm

apart, reddish green. Flowers lasting one day,

4.5- 5 mm across, pale creamy yellow. Sepals

spreading widely free to the base. Dorsal

sepal concave, elliptic to obovate, c. 2.8 x

1.6- 1.8 mm. Lateral sepals broadly elliptic,

acute at the apex, c. 2.6 x 1.6 mm, keeled on

the back. Petals spathulate, c. 2 x 1.4 mm.

Labellum thick and fleshy, deeply concave

with raised sides and obtuse at the apex, c. 2.2

x 1.8 mm. Spur 1.4-1 mm, in line with the

labellum. Column short and stout, c. 1.2 x 1

mm, purplish towards the front. Anther cap

c. 0.8 x 1 mm, beaked at the front. Pollinia 4

in unequal pairs. Capsule 19-20 x c. 4.5 mm,

with slight longitudinal ridges. Figs. 2-4.

Additional specimens examined : Queensland. Cook

District: Wasp Creek north of Bamaga, Sep 1989, Gray

BG5120 (CNS); Usher Point, Cape York, Jan 2010, Gray

BG9681 & Bourne (CNS).

Distribution and habitat : Taeniophyllum

epacridicola is endemic to northern Cape

York where it is known from a few localities

north of the Jardine River (Map 1). Specimens

examined in the field grow on twigs and small

branches primarily in epacrid dominated

shrubland at elevations below 100 m. T.

epacridicola has also been rarely observed in

the margin of the rainforest.

Phenology : Flowering and fruiting has been

recorded between July and January.

Notes : Taeniophyllum epacridicola is in the

section Taeniophyllum (synonym: section

Trachyrhachis Schltr.), and this represents the

first record of the section for Australia. The

first specimen of T. epacridicola examined

385

was on a fallen branch in the edge of rainforest

near Lockerby homestead north of Bamaga in

September 1979. However, efforts to locate

more material for detailed examination were

unsuccessful until a second collection (sterile)

was made at Wasp Creek in September

1989 (Gray BG5120 ), and was successfully

flowered in cultivation in September 1990

(Gray BG5235). In 2008 another population

was located at Usher Point by David Baume.

This population was revisited in January of

2010 to enable voucher preparation (Gray

BG9681 & Baume). A single plant (tentatively

identified as this species), has been located

close to Punsand Bay; however, no fertile

specimen has been available to date.

Etymology : Taeniophyllum epacridicola is

named for its seeming preference to grow

in epacrid dominated shrubland consisting

of Leucopogon ruscifolius R.Br. and L.

yorkensis Pedley (Ericaceae).

2. Taeniophyllum explanatum B.Gray sp.

nov. Similar to T. muelleri Lindl. ex Benth.

but differs in having roots flattened in cross-

section. Typus: Queensland. Cook District:

Bridle Creek on power line access road, 24

October 2002, B. Gray BG8339 (holo: BRI;

iso: CNS).

Plant epiphytic forming small clumps with

5-30 roots. Roots ± flat in cross section, 1.5-

2 mm across, up to 10 cm long, dull green.

Inflorescences filiform, peduncle 12-25

mm x c. 0.2 mm with 1-3 bracts. Rachis

increasing in length as flowering progresses

producing 10-30 flowers one at a time; buds,

flowers and capsules can be present at the

same time. Floral bracts acute, alternate,

0.5-0.6 mm long, c. 0.5 mm apart and all

in one plane. Flowers opening singly, c. 2.5

mm long including the spur and c. 2.5 mm

across when open, green. Sepals and petals

connate at the base into a tube c. 0.8 mm

long, then spreading. Dorsal sepal linear

to narrowly lanceolate, c. 2 x 0.6-0.7 mm,

incurved. Lateral sepals linear, c. 2 x 0.6

mm. Petals ovate, c. 1.8 x 0.6 mm, incurved,

apex acuminate. Labellum cymbiform,

narrowly triangular, 1.8-2 mm x c. 0.5 mm

with low erect lobes at the base, apex acute

with an inflexed spur c. 0.5 mm long. Spur

386

Austrobaileya 9(3): 382-392 (2015)

Fig. 2. Taeniophyllum epacridicola. A. habit of a mature flowering plant. B. face view of flower. C. lateral view of

flower. D. lateral view of column and labellum. E. longitudinal section of flower. F. face view of anther. G. lateral view

of anther. H. pollinia. I. transverse section of mature root. All from Gray BG5235 (CNS). Scale as indicated. Del. B.

Gray.

Gray, New species of Taeniophyllum from Queensland

387

Fig. 3. Taeniophyllum epacridicola. A fruiting specimen showing flat roots closely appressed to a branch of an

understorey tree {Gray BG9681, CNS). Photo: B. Gray.

subglobose, c. 0.9 x 0.7 mm. Column domed,

c. 0.7 x 0.4 mm, wings forward facing and c.

0.3 x 0.1 mm. Anther cap c. 0.4 x 0.3 mm,

with 2 prominent humps. Pollinia 4 in two

pairs, pale yellow. Capsule not seen. Figs. 5

& 6 .

Additional specimens examined : Queensland. Cook

District: Robson Creek, Danbulla NP, Aug 2014, Ford

6332 (CNS); Bridle Creek on power line access Road,

Aug 2009, Gray BG9312 (CNS); Windsor Tableland NP,

Oct 2014, Gray BG9674, Baume & Walker (CNS); Mt

Windsor Tableland, May 1989, Jones 4377 & Clements

(CANB).

Distribution and habitat : Taeniophyllum

explanatum occurs within the Wet Tropics of

Queensland from Mount Windsor Tableland

south to Innisfail (Map 2) at elevations from

500 to 1000 m, in rainforest, usually on small

trees and vines.

Phenology : Flowering has been recorded

between May and January.

Fig. 4 .Taeniophyllum epacridicola. Close-up view of Notes'. Taeniophyllum explanatum has only

an open flower {Gray BG5235, CNS). Photo: B. Gray. been collected on a few occasions possibly

388

Austrobcdleya 9(3): 382-392 (2015)

Fig. 5. Taeniophyllum explanatum. A. habit of a mature flowering and fruiting plant. B. lateral view of flower. C. face

view of flower. D. transverse section of mature root. E. face view of column. F. lateral view of column. G. face view

of labellum. H. pollinia. I. lateral view of anther. J. face view of anther. K. lateral view of labellum and column. L.

longitudinal section through flower. M. dorsal sepal. N. lateral sepal. O. petal. All from Gray BG8339 (CNS). Scale as

indicated. Del. B. Gray.

Gray, New species of Taeniophyllum from Queensland

389

Fig. 6. Taeniophyllum explanatum. Plant showing inflorescences bearing flower buds and open flowers (Gray BG9674,

CNS). Photo: B. Gray.

because this small species is easily overlooked

in the rainforest. Most of the collections made

are from the more accessible understorey;

however, a single collection ( Ford 6332 ) is

from the rainforest canopy.

Etymology : The specific epithet refers to the

roots being flat.

3. Taeniophyllum triquetro radix B.Gray sp.

nov. Similar to Taeniophyllum muelleri Lindl.

ex Benth. but differs in having roots triangular

in cross-section. Typus: Queensland. Cook

District: Ridge above Tinaroo Dam perimeter

road, 8 September 1990, B. Gray BG5238

(holo: BRI; iso: CNS).

Taeniophyllum sp.; Ormerod (1994).

Illustrations : Jones (2006: 454), as T.

confertum B.Gray & D.L.Jones.

Plants epiphytic, spider like in appearance

sometimes proliferating from root tips

and forming small colonies. Roots 4-10,

appressed to the host, ± triangular in cross

section, 10-60 mm x 1.5-2 mm, glaucous to

greyish green, strongly ridged on the upper

surface. Inflorescences 1—3(—4); peduncle

5-12 mm long. Rachis extending as flowering

progresses; 5-20 or more flowered with one or

sometimes two open together; buds, flowers

and capsules can be present at the same

time. Floral bracts projecting 0.5-0.6 mm,

somewhat fleshy and all in one plane. Flowers

c. 4.5 mm long including the spur and 3 mm

in diameter, green, aging to yellowish green.

Sepals and petals fused at the base into a tube

c. 1.4 mm long. Sepals spreading, somewhat

ovate to narrowly triangular, c. 1.5 x 0.9 mm,

acute at the apex. Petals spreading, ovate, c.

1.4 x 0.8-0.9 mm, acute at the apex. Labellum

fleshy, cymbidiform c. 1.9 mm long, obscurely

three lobed near the base, lateral lobes c. 0.3

mm high, midlobe narrowly triangular, acute

at the apex with an inflexed spine c. 0.6 mm

long at the tip. Spur elongate-globose, c. 1.9

x 0.9 mm. Column domed c. 0.7 mm high,

390

Austrobaileya 9(3): 382-392 (2015)

Fig. 7. Taeniophyllum triquetroradix. A. habit of a mature flowering plant. B. lateral view of flower. C. face view of

flower. D. face view of labellum. E. face view of column. F. lateral view of column. G. longitudinal section of labellum

and column. H. lateral view of anther. I. face view of anther. J. face view of pollinia. K. lateral view of pollinia. L.

transverse section of mature root. M. lateral view of labellum and column. N. dorsal sepal. O. petal. P. lateral sepal. All

from Gray BG5238 (BRI). Scale as indicated. Del. B. Gray.

Gray, New species of Taeniophyllum from Queensland

column wings projecting forward, c. 0.4 mm

long. Anther Cap c. 0.5 x 0.5 mm, with two

distinct humps. Pollinarium c. 0.5 mm long.

Pollinia 4, ovoid, in 2 equal sized pairs,

yellow. Capsule 11.5-12 x 2-3 mm with 5-6

longitudinal ridges. Figs. 7 & 8.

Additional specimens examined : Queensland. Cook

District: Euluma Creek Road, Julatten, Jan 2003, Gray

BG8424 (CNS); Bushy Creek, Julatten, Aug 1985, Gray

BG4129 (CNS); Cairns Inlet near White Rock, Jul 2013,

Ormerod008( CNS).

Fig. 8. Taeniophyllum triquetroradix. Close-up view

of a flowering specimen in cultivation showing (i)

triangular roots with a clear dorsal median ridge, and (ii)

an inflorescence bearing a flower bud and an open flower

(Gray BG8424, CNS). Photo: B. Gray.

Distribution and habitat : Taeniophyllum

triquetroradix occurs within the Wet Tropics

of Queensland between Mossman and

Innisfail (Map 2) from sea level to 400 m.

Plants have been most commonly found in

poorly developed, open rainforest, either in

391

the understorey or the upper canopy based on

specimens found on fallen branches; however,

it has also been recorded at the edge of

mangroves. Some populations have also been

observed on trees in open paddocks.

Phenology : Flowers have been observed

between July and January.

Notes: Taeniophyllum triquetroradix is

probably more common than the number of

collections indicates because plants are small

and often overlooked.

Etymology : The specific epithet refers to the

roots which are triangular in cross section.

Acknowledgements

I am grateful to David Baume for his

kind help with locating new populations

of Taeniophyllum epacridicola and T.

explanatum , and who also assisted me in

numerous field trips; James Walker for good

company in the field, testing and commenting

on the taxonomic key, and his constructive

criticism with the manuscript. Special thanks

to Yee Wen Low (Singapore Botanic Gardens)

and Paul Ormerod (Cairns, Queensland) for

help with the initial stage of the manuscript.

Professor Darren Crayn and Frank Zich kindly

provided me with access to the Australian

Tropical Herbarium (CNS) to examine the

Taeniophyllum collection there; Paul Forster

and the curators of the Queensland herbarium

(BRI) for making available relevant materials

on loan for this study.

References

Clements, M.A. (1989). Catalogue of Australian

Orchidaceae. Australian Orchid Research 1:

1-160. (Taeniophyllum, p. 136).

Dockrill, A.W. (1992). Australian Indigenous Orchids ,

2 nd edition, 2: 874-886. Surrey Beatty &

Sons, in association with Society for Growing

Australian Plants: New South Wales.

Gray, B. & Jones, D.L. (1984). Miscellaneous notes

on the orchids of North Eastern Queensland.

1 - A new species of Bulbophyllum and

Taeniophyllum. Orchadian 8: 40-43.

Jones, D.L. (2006). A Complete Guide to Native Orchids

of Australia, including the Island Territories.

Reed New Holland: Sydney. ( Taeniophyllum , p.

453-455).

392

Jones, D.L., Gray, B. & Clements, M.A. (2006).

Taeniophyllum norfolkianum (Orchidaceae) a

new species from Norfolk Island. Orchadian

15: 156-158.

Kocyan, A. & S chuiteman, A. (2014). New combinations

in Aeridinae (Orchidaceae). Phytotaxa 161:

61-85.

Lavarack, P.S. & Gray, B. (1985). Tropical Orchids

of Australia. Thomas Nelson: Melbourne.

{Taeniophyllum, p. 140-143)

Margonska, H.B. & Szlachetko, D.L. (2010).

Orchidaceae of Tahiti, French Polynesia.

Gdansk University Press: Gdansk.

{Taeniophyllum, p. 86-90)

Ormerod, P. (1994). Orchids of Trinity Inlet. Orchadian

11: 149.

Map 1. Distribution of Taeniophyllum epacridicola,

grey shaded areas are conservation reserves.

Austrobaileya 9(3): 382-392 (2015)

Schlechter, R. (1982). The Orchidaceae of German

New Guinea. Australian Orchid Foundation:

Melbourne. {Taeniophyllum, p. 1077-1096)

Seidenfaden, G. (1995). The Descriptions

Epidendrorum of J.G. Konig 1791. Olsen &

Olsen: Fredensborg. {Taeniophyllum pusillum,

p. 23-24)

Seidenfaden, G. & Wood, J.J. (1992). The Orchids of

Peninsular Malaysia and Singapore. Olsen &

Olsen: Fredensborg. {Taeniophyllum, p. 571—

581)

Thiers, B. (continuously updated). Index Herbariorum:

A global directory of public herbaria and

associated staff New York Botanical Garden’s

Virtual Herbarium, http://sweetgum.nybg.org/

ih/, accessed November 2014.

Map 2. Distribution of Taeniophyllum explanatum

A and T. triquetroradix CD, grey shaded areas are

conservation reserves.

A taxonomic revision of Cynometra L. (Fabaceae) in

Australia with a new species from the Wet Tropics of

Queensland and a range extension to the mainland

W.E.Cooper

Summary

Cooper, W.E. (2015). A taxonomic revision of Cynometra L. (Fabaceae) in Australia with a new

species from the Wet Tropics of Queensland and a range extension to the mainland. Austrobaileya

9(3): 393-403. Cynometra comprises three species in Australia. In addition to C. iripa Kostel., the

new species C. roseiflora W.E.Cooper is described, illustrated and distinguished from related species.

C. ramiflora L. is newly recorded as occurring on the Australian mainland in north Queensland. All

species are described with notes provided on typification, distribution and habitat. An identification

key to the species of Cynometra in Australia is presented.

Key Words: Fabaceae, Leguminosae, Cynometra, Cynometra iripa, Cynometra ramiflora, Cynometra

roseiflora, new species, taxonomy, Australia flora, Queensland flora, identification key

W.E. Cooper, Australian Tropical Herbarium, James Cook University, Cairns Campus, McGregor

Road, Smithfield, Queensland 4878, Australia.

Introduction

Cynometra L. has been classified in the

leguminous tribe Detarieae (within the

Caesalpiniaceae) that broadly corresponds to

the ‘Detarieae clade’ recovered from multiple

genetic sequence data (Bruneau et al. 2008).

There appears to be a close relationship with

Maniltoa based on the available genetic

evidence; however, taxon sampling has been

limited to date (Bruneau et al. 2008), hence

the traditional circumscription of Cynometra

based on floral morphology (Knapp-van

Meeuwen 1970; Ding Hou 1996) is followed

in the current paper.

Cynometra comprises approximately

88 species when so defined (The Plant List

2013) with a pantropical distribution from

the African continent, Indian subcontinent

(including Indian Ocean Islands, India, Sri

Lanka), SE Asia, Malesia, Pacific Islands,

Australia to Central America (Mexico), the

West Indies and South America (Brazil,

Argentina and Chile), occurring in forests

from sea level to altitudes of 1300 m. The

genus has its greatest diversity on the African

continent (Cowan & Polhill 1981: 124) and

Accepted for publication 23 July 2015

is present only in small species numbers in

Australia and the western Pacific.

Historically, within Australia and in Asia,

specimens of Cynometra iripa Kostel. (in

the sense applied in this paper) have been

previously determined as C. ramiflora var.

bijuga (Bentham 1864; Bailey 1900; Knaap-

van Meeuwen 1970). Recent consensus

has been that true C. ramiflora did not

occur on mainland Australia (Knapp-van

Meeuwen 1970; Tomlinson 1986; Ding Hou

1996); although it has been subsequently

recorded from the Australian territory of

Christmas Island in the Indian Ocean (Du

Puy 1993; Claussen 2005). Despite these

published statements, Cynometra ramiflora

does indeed occur on mainland Australia

and was first collected at the Jardine River

on Cape York Peninsula in 1978 {Stirling

AIM462 BRI), although this specimen was

possibly overlooked by the second and

third groups of authors mentioned above.

Cynometra ramiflora was again collected in

1991 ( Sankowsky 1223 [BRI], this specimen

resulting in the phrase name Cynometra sp.

(Paira Homestead Rd., G. Sankowsky +1223)

at BRI) and again at Jardine River in October

2014 {Cooper, Jensen, Kemp & Zdenek 2265,

2267 .; 2268 [CNS]).

394

In 1992, Garry Sankowsky collected a

sterile specimen of Cynometra at Mossman

River. This specimen and further collections

were later determined at QRS (now CNS)

to be C. ramiflora, and as C. iripa by the

Queensland Herbarium (BRI). Habitat as

well as leaf, flower and fruit morphology

indicated that these collections were distinct

from C. ramiflora and C. iripa and required

investigation.

This morphological and ecological study

confirms that three species of Cynometra

occur in Australia: C. iripa , C.ramiflora and

the newly described C. roseiflora known

only from the Mossman River. Knaap-Van

Meeuwen (1970: 13) stated that all Indo-

Pacific species of Cynometra grow under

ever-wet conditions; however, the three

species that occur in Australia grow in

tropical climates with a distinct dry season.

C. iripa and C. ramiflora do occur in back

mangal {sensu Tomlinson 1986) areas, but C.

roseiflora occurs in rainforest with a distinct

dry season on porous granitic soil.

Materials and methods

The study is based upon the examination of

selected herbarium material from CNS, BRI,

CANB and NSW (herbarium acronyms as

per Index Herbariorum: A Global Directory

of Public Herbaria and Associated Staff

2015), combined with field observations of all

species. All specimens cited have been seen

by the author.

Measurements of the floral parts and fruits

are based on fresh material as well as material

preserved in 70% ethanol.

Taxonomy

Cynometra L., Sp. PI. 1: 382 (1753) & Gen.

PI, 5 th edn. 466 (1754). Type species: C.

cauliflora L.

Austrobaileya 9 ( 3 ): 393-403 ( 2015 )

Cynometra in Australia: Monoecious shrubs

or trees to 15 m tall, can be multistemmed

or buttressed. Bark finely fissured, lenticels

linear or round; stipules caducous and leaving

no visible scar; bracts enclosing new leaves

and inflorescences similar to the stipules,

several overlap to create a cone-like structure,

broadly reniform, semi-circular, broadly

ovate, somewhat cupular or oblong-obovate,

0.8-2.5 mm long, rust coloured, minute hairs

on dorsal surface, margin ciliate, caducous.

Leaves coriaceous, alternate, 1-2-jugate,

discolorous, new leaves green, pink or

bright red; petiole + rachis chanelled on the

upperside; petiolules thickened, wrinkled,

mostly enclosed by leaflet base; leaflets

opposite, asymmetrical, basal pair (if present)

are smaller than terminal pair; base oblique;

margin entire; venation brochidodromous.

Inflorescences axillary, terminal or

ramiflorous, racemes on a swelling; bracteoles

1 or 2 towards pedicel base, filiform or strap¬

shaped, 1.5-3.5 mm long, caducous. Flowers

bisexual; hypanthium inverted cone-shaped;

sepals 4, imbricate, unequal in width, margin

ciliate at apex, somewhat ragged or entire,

acute; petals 5, free, white or pink, margin

entire or barely fimbriate at apex, caducous;

disk absent; stamens 10; anthers orbicular-

cordate, cleft at base, bilocular, introrse,

medi-dorsifixed, dehiscing longitudinally;

ovary asymmetrically elliptical, stipe short

and inserted excentrically; ovules 1 (rarely

2 or 3); style slender, almost directly in line

with dorsal margin of ovary or excentric to

varying degrees and becoming more excentric

post anthesis; stigma capitate. Fruit an

indehiscent woody nut, asymmetrical with a

beak at apex of dorsal suture, rugose, scurfy;

seeds 1 (rarely 2 or 3). Germination epigeal.

Key to Cynometra species in Australia

1 Petals bright pink, about half as long as sepals; ovary inner wall glabrous

but for a tuft of hairs at base; new leaves bright red.3. C. roseiflora

1. Petals white or very pale pink, about same length as sepals; ovary inner

wall glabrous to generally hairy (not tufted at base); new leaves green or pink

2

395

Cooper, Cynometra in Australia

2 Leaf rachis and petiolules minutely hairy; sepal apices ciliate; ovary

inner walls pubescent; fruit laterally compressed and distinctly beaked

partway along dorsal side; new leaves green.1. C. iripa

2. Leaf rachis and petiolules glabrous on mainland Australia (minutely

hairy or glabrescent on Christmas Island); sepal apices entire

or few-toothed; ovary inner walls glabrous or with a few sparse

hairs; fruit globose and with a small beak near apex of dorsal side;

new leaves pink .2. C. ramiflora

1. Cynometra iripa Kostel., Allg. Med-

Pharm. FI. 4: 1341 (1835). Type: Rheede,

Hort. Malab. 4: t. 31 (1683).

Cynometra ramiflora var. bijuga auct. non

(Span, ex Miq.) Benth. as to type; Bentham

(1864: 296).

Cynometra ramiflora auct. non. L.; Bailey

(1900: 469).

Illustrations : Parkinson A (1768-71) as

C. ramiflora ; Lear & Turner (1977: 31) as

C. ramiflora ; Wightman (1989: 63); Ross

(1998: 168); Tomlinson (1986: 253); Cooper

& Cooper (2004: 102); Duke (2006: 136, 137

[upper image], (2013).

In Australia: Shrub or tree to 6 m, sometimes

multistemmed. Bark finely fissured, lenticels

linear or round, pale; new flush foliage green;

stipules not seen. Leaves 1-2-jugate; petioles

1-4 mm long, minutely pubescent; petiole

+ rachis 9-30 mm long, channelled on the

upperside, puberulent, lenticels pale; petiolules

up to 3 mm long, wrinkled, mostly enclosed

by leaflet base, puberulent on underside,

glabrescent on upperside; leaflets discolorous,

asymmetrical, obliquely-elliptical, obliquely-

oblong, obliquely-obovate or obliquely-ovate;

basal pair 8.5-50 x 5-30 mm; terminal pair

23-85 x 11-50 mm; coriaceous, glabrous;

base oblique, cuneate, attenuate or obtuse;

apex obtuse and emarginate; margin entire;

venation brochidodromous, primary vein

raised (more so on under-side); secondary

veins 6-10 pairs, slightly raised on both sides

of dried specimens but ± flush on fresh leaves,

angle of divergence from primary vein 50-70°,

forming loops 2-5 mm from margin; tertiary

venation reticulate. Inflorescences axillary,

terminal or ramiflorous, solitary racemes or

a fascicle of racemes on a swelling, racemes

3-9-flowered; bracteoles 1 or 2 towards pedicel

base, filiform, 1.5-3 mm long, sparsely hairy,

caducous. Flowers fragrant, erect, 6.5-8.5

x 5.5-8 mm wide; receptacle inverted cone-

shaped, c. 1.5 x 1.5 mm; sepals 4, unequal in

width, lanceolate or oblong-ovate, becoming

reflexed and often incurved at apex, 2.5-3 x

0.5-1.7 mm, white or very pale pink; margin

at apex ciliate, from mid position to base

entire or sparsely ciliate; glabrous or with

few sparse hairs on abaxial surface; petals

5, white or very pale pink, lanceolate, 2-3

x 0.5-0.7 mm, glabrous, caducous; stamens

10; filaments terete, 4-7 mm long, straight or

curved, glabrous; anthers c. lxl mm, brown;

ovary inserted excentrically on a short stipe,

c. 2.25 x 1.25 mm, pink, white pubescent

externally, appressed-pubescent internally;

ovules 1 (rarely 2); style slender, almost

directly in line with dorsal margin of ovary or

excentric to varying degrees, becoming more

excentric post anthesis, 2-4 mm long, sparsely

hairy from median section to base; stigma

capitate. Fruit on a 6-10 mm long pedicel, an

asymmetrical nut with a distinct beak at apex

of dorsal suture and partway along dorsal

side, suborbicular, laterally compressed, 30-

40 x 30-34 x 17.5-20 mm, deeply rugose,

glabrescent, scurfy, brown; seeds 1 (rarely 2),

25-29 x 17-20 mm. wrinklepod mangrove.

Figs. 1A, 2A-C.

Additional selected specimens (from 60 examined ):

Australia: Northern Territory. Ingliss Island, Dec

1987, Dunlop 7510 (CANB); Arnhem Bay, Probable

Island, Oct 2009, Westaway 3190 (NSW). Queensland.

Cook District: c. 6 km NE of Mapoon community,

Sep 2006, Wharton s.n. (BRI [AQ783524]); Cape York

Peninsula, E of Bramwell Homestead on Olive River,

Dec 1987, Kanis 2047 (CANB); Weipa Concession,

Marmoss Creek, Sep 1974, Dockrill 853 (CNS); near

Taylors Landing, Claudie River, Jul 2014, Cooper 2239,

Jensen & Venables (CNS); Nesbit River, Jun 1995,

Forster PIF17083 (CNS); Annie River, Dec 1978, Duke

AIM877 (BRI); Endeavour River Estuary, May 1991,

Le Cussan 25 (CNS); Cooktown, Keatings Lagoon

396

Fig. 1. Cynometra flowers: A. Cynometra iripa ( Cooper

2238 et al. [CNS]), B. C. ramiftora ( Cooper 2245 et al.

[CNS]), C. C. roseiflora ( Cooper 2215 et al. [CNS]).

Photos: A & B, W. Cooper; C, T. Hawkes.

Austrobaileya 9(3): 393-403 (2015)

Conservation Park, Dec 2008, Booth 5243 & Lynch

(BRI); 100 m N of Daintree River ferry crossing, Jul

1995, Gray 6247 (CNS); North bank of Mossman River

mouth, Sep 1948, Smith 3997 (CANB); Redden Island,

Machans Beach, Apr 2014, Cooper 2235, Venables &

Cooper (CNS); Holloways Beach, Jan 2015, Cooper

2273 (CNS); Holloways Beach, Jul 2014, Cooper 2238

& Venables (CNS); Russell River Road, Jul 2004, Gray

8933 (CNS); Mouth of Maria Creek near Kurramine

Beach, Jul 1994, Waterhouse 3356 (BRI); Deluge Inlet,

Hinchinbrook Island, Aug 1976, Abel AS144 (BRI).

North Kennedy District: Hayman Island, Jun 1934,

White 10120 (BRI). South Kennedy District: Mackay,

Sep 1968, Jones s.n. (BRI [AQ19340]).

Distribution and habitat : Cynometra iripa

is distributed throughout tropical southern

and south-eastern Asia (including India,

Bangladesh, Myanmar, Cambodia, Vietnam

and Thailand), Malesia (Malaysia, Indonesia,

Singapore, the Philippines and New Guinea),

Micronesia, Melanesia (Solomon Islands,

New Caledonia) and Australia, at altitudes

reportedly to 500 m (Ding Hou 1996). Within

Australia, C. iripa has been recorded in

Queensland on the mainland and off shore

islands from Cape York to the Mackay area,

as well as west to East Arnhem Land in the

Northern Territory (Map 1); altitude near sea-

level to 20 m.

In Australia, Cynometra iripa is a plant

of back mangal areas, rarely of upstream

wetlands (Keatings Lagoon near Cooktown

lBooth 5243 & Lynch]) or adjoining

monsoon forest (Arnhem Land [Westaway

3190]). In Queensland it commonly co¬

occurs with Acacia polystachya A.Cunn.

ex Benth., Acrostichum speciosum Willd.,

Aegiceras corniculatum (L.) Blanco, Ary tern

bifoliolata S.T.Reynolds, Avicennia marina

(Forssk.) Vierh., Brownlowia argentata

Kurz, Bruguiera gymnorhiza (L.) Savigny,

Cierodendrum inerme (L.) Gaertn.,

Cryptocarya exfoliata C.K.Allen, Dalbergia

candenatensis (Dennst.) Prain, Derris

trifoliata Lour., Dillenia alata (R.Br. ex DC.)

Martelli, Dysoxylum acutangulum subsp.

foveolatum (Radik.) Mabb., Excoecaria

agallocha L., Ganophyllum falcatum Blume,

Heritiera littoralis Dryand., Lumnitzera

littorea (Jack) F.Voigt, Melaleuca leucadendra

(L.) L., Thespesia populneoides (Roxb.)

Kostel., Terminalia sericocarpa F.Muell and

Xylocarpus spp. In the Northern Territory it is

Cooper, Cynometra in Australia

397

Fig. 2. Cynometra fruit all natural size: C. iripa (Cooper 2273 [CNS]), A. ventral view, B. lateral view showing beak

on dorsal side; C. ramiflora {Cooper 2265 et al. [CNS]), C. dorsal view, D. lateral view; C. roseiflora {Cooper 2271 et

al. [CNS]), E. lateral view, F. ventral view, G. dorsal view. Del. W.T. Cooper.

known to occur in back mangal communities

co-occurring with Aegiceras corniculatum

(L.) Blanco, Lumnitzera racemosa Willd.

and Flacourtia territorialis Airy Shaw, and

in adjoining monsoon forest dominated by

Peltophorum pterocarpum (DC.) Backer ex

K. Heyne.

Phenology : Flowers have been recorded from

March to October; fruits have been recorded

from January to March, June to September

and in November.

Typification: Cynometra iripa Kostel. is

solely based on Rheede’s plate in Hortus

Malabaricus (Kosteletzky 1835; Knaap-van

Meeuwen 1970). This plate was also one of

the two elements cited by Linnaeus (1753) in

his description of C. ramiflora L.; however,

this element has now been excluded from

398

typification of that name (Kosteletzky 1835;

Knaap-van Meeuwen 1970). Kosteletzky

(1835) quite clearly indicated the single

typifying elements for both his name and that

of Linnaeus (1753) and provided accounts of

both species. Although this does not equate

to a formal lectotypification in the modern

sense, subsequent authors have followed

this citation (e.g. Knaap-van Meeuwen 1970;

Ding Hou 1996: 606; Ross 1998: 169). Jarvis

(2007) goes so far as to state “Lectotype

(Knaap-van Meeuwen in Blumea 18: 23

(1970): [icon] “ Cynomorium Silvestre ” in

Rumphius, Herb. Amboin. 1: 167, t. 63. 1741”,

although Kosteletzky (1835) rather than

Knaap-van Meeuwen (1970) should probably

be regarded as making this decision: neither

state “lectotypify” as such.

Notes: Cynometra iripa has been described

as having a style not in line with the dorsal

suture of the ovary (Knaap-van Meeuwen

1970), but Australian material has styles

that are often directly in line with the dorsal

suture, although during and after anthesis

they become excentric to varying degrees.

All living specimens seen in the Cairns area,

as well as along the Claudie and Olive Rivers,

have pink ovaries with a dense covering of

white hairs, thus differing from the rust-

coloured pubescence described by Ross

(1998). Fresh specimens from other areas

were not seen for comparison.

Sepals have been described as being

rather long-hairy (Knaap-van Meeuwen

1970: 23; Ding Hou et al. 1996: 603). Sepals

of Australian specimens seen by the author

of this revision do not have a hairy surface

but do have a ciliate margin especially at the

apex.

Etymology: There has been confusion over

the etymology of the specific epithet iripa. It is

clear that Rheede’s name for this plant, Iripa ,

was based on an indigenous Malayalam (native

language of southern India) name (Rheede’s

opening statement is ‘Iripa Malabarensibus’;

Ram (2005) also indicates that Malayalam

names were employed by Rheede). This is

further confirmed by Nicolson et al. (1988),

who reported that “Irippa (sic) is still used. It

is found [in] mangrove swamps, increasingly

Austrobaileya 9(3): 393-403 (2015)

scarce in Kerala”. Hence the etymology for

iripa is derived from Irippa, the Malayalam

name for the plant.

2. Cynometra ramiflora L., Sp. PI. 382

(1753). Type: based on Cynomorium sylvestre

Rumph., Herb. Amboin. 1: 167, t. 63 (1741),

fide Knaap-van Meeuwen (1970: 23 “excl. syn

Iripa Rheede”).

Cynometra sp. (Paira Homestead Rd

G.Sankowsky+ 1223); Pedley (2007: 39,2010:

34).

Illustrations: Beddome (1869-1874); Pierre

(1880-1907); Kirtikar & Basu (1918);

Verdcourt (1979: 84); Whitmore (1983: 255);

Du Puy (1993: 180 D & E); Corner (1997:

399); Claussen (2005: 21); Duke (2006: 137)

as C. iripa (lower photo).

In Australia: Tree to 15 m, dbh to 60 cm,

buttressed, blaze red. Bark with numerous

lenticels, these round or elongated and often

in vertical lines; twigs with scattered to dense

round and linear lenticels; new flush foliage

pink to cream; stipules filiform, c. 1.5 mm

long; tuft of hairs at petiole apex c. 0.75 mm

long, caducous. Leaves 1 (rarely 2)-jugate;

petiole (+ rachis when present) 5-33 mm long,

shallowly grooved on upper side, glabrous

on mainland specimens and glabrescent on

Christmas Island specimens; petiolules 2-6

mm long, thickened and mostly enclosed by

leaflet base, glabrous on mainland specimens

but minutely pubescent on Christmas Island

specimens; leaflets discolorous, asymmetrical,

obliquely-ovate, obliquely-oblong, obliquely-

obovate or broadly elliptical; basal leaflets, if

present, 17-82 x 9-46 mm; terminal leaflets

63-210 x 30-98 mm; coriaceous, shiny,

glabrous; base oblique, rounded, truncate,

attenuate or cuneate; apex acute, shortly

acuminate, acuminate or emarginate; margin

entire; venation brochidodromous, primary

vein raised on upperside, secondary veins

in 8-12 pairs (6 -7 pairs on basal leaflets

if these present), angle of divergence from

primary vein 40°-50°; tertiary venation

reticulate. Inflorescences axillary, terminal

or ramiflorous, solitary or paired (sometimes

condensed) racemes, up to 20-flowered; rachis

to 20 mm long, glabrous; pedicels 6.5-12.5

Cooper, Cynometra in Australia

mm long, glabrous; bracteoles near pedicel

bases boat-shaped and keeled, 3-3.5 x c. 1.5

mm, ciliate at apex, dorsal surface with hairs

along keel line and some scattered minute

hairs may be present, caducous. Flowers

with an unpleasant odour, erect, 5-9 x c. 9

mm; receptacle inverted cone-shaped, c. 1.5 x

1.5-2.5 mm; sepals 4, oblong-ovate, c. 5 x 2.5

mm, white, apex acute and somewhat ragged

or slightly fimbriate, otherwise margin entire,

glabrous; petals 5, obovate-lanceolate, 5.5-6

x c. 1.5 mm, white, caducous, glabrous, apex

acute or mucronate and somewhat ragged or

slightly fimbriate, otherwise margin entire;

stamens 10; filaments terete, straight or

curved, 6-8 mm long, glabrous; anthers c.

1.25 mm long, cream to brown; ovary inserted

slightly off centre on its stipe, c. 1 mm long,

yellowish or pink, pubescent externally,

internal walls glabrous (or with a few sparse

hairs Cooper 2245); ovules 1-3; style in line

with dorsal suture or frequently excentric,

3-3.5 mm long, sparse hairs towards base;

stigma capitate. Fruit on a 3-10 mm long

pedicel, an asymmetrical, somewhat globose

nut, but flattened on the ventral side, 38-52 x

37-42 x 28.5-40 mm, rust brown coloured,

woody, rugose, scurfy, minutely and sparsely

pubescent, beak short, near apex of dorsal

suture, seed solitary, wrinklepod mangrove.

Figs. IB, 2C & D, 3

Additional selected specimens {from 18 examined ):

Malaysia. Pahang, Kuantan Telok Chempedak Bay, Jul

1992, Saw FRI37559 (CNS). Papua New Guinea. New

Britain Province: Cape Roebuck, west of Fullerborn

Harbour, May 1973, Womersley NGF41214 (CNS).

Western province: Morehead River, c. 8 miles inland,

Aug 1967, Pullen 7057 (CANB), 7074 (CANB); junction

of Bensbach & Tarl Rivers, Bensbach sub-district, Aug

1967, Ridsdale & GaloreNGF33711 (CANB). Australia.

Christmas Island. SE Ross Hill, Mar 2002, Holmes

CI43 & Holmes (CANB); Cultivated at National Park

Headquarters, Dec 2014, Cooper 2285 & Maple (CNS);

Cultivated at 21 Central Rehab Field, Dec 2014, Cooper

2286 & Maple (CNS). Queensland. Cook District: Paira

Homestead, Sep 1991, Sankowsky 1223 (CNS); Old Paira

Homestead Road, Cape York, Feb 1992, Sankowsky 1320

(CNS); Jardine River, Nov 1978, Stirling AIM462 (BRI);

Jardine River, Oct 2014, Cooper 2267, Jensen, Kemp &

Zdenek (CNS); Jardine River, Oct 2014, Cooper 2265,

Jensen, Kemp & Zdenek {C NS); Jardine River, Oct 2014,

Cooper 2268, Jensen, Kemp & Zdenek (CNS); ex Mew

River, Muddy Bay, Cape York (cultivated at Tolga), Oct

2005, Sankowsky 2668 & Sankowsky (BRI); ditto Joe.,

(cult, by Sankowsky at Tolga), Oct 2014, Cooper 2258,

399

Fig. 3. Cynometra ramiflora showing pink colouration

on pendulous new leaves {Sankowsky 2668 et al. [CNS]).

Photo: G. Sankowsky.

Cooper & Sankowsky (CNS); Cairns Botanic Gardens

(ex Mew River), May 2014, Cooper 2275 & Venables

(CNS); Cairns Botanic Gardens (ex Mew River), Aug

2014, Cooper 2245, Cooper & Roberts (CNS).

Distribution and habitat : Cynometra

ramiflora occurs in India, Sri Lanka,

South-east Asia (including Thailand),

Malesia (including Indonesia, Malaysia,

the Philippines, New Guinea), Melanesia

(Solomon Islands, New Caledonia) and

Australia.

On mainland Australia, it has been

recorded from two locations from north

Queensland near Cape York (Map 1). One

site is along the Jardine River in areas of

back mangal on white sand and mangrove

mud where it co-occurs with Acrostichum

speciosum Willd., Calophyllum inophyllumL.,

Cerbera manghas L., Crinum pedunculatum

R.Br., Heritiera littoralis Dryand., Melaleuca

quinquenervia (Cav.) S.T.Blake, Rhizophora

spp. and Xylocarpus granatum K.D.Koenig.

The second is near the Mew River on the east

coast adjacent to mangroves in swamp forest

dominated by Livistona benthamii F.M.Bailey.

On Christmas Island, C. ramiflora occurs

in isolated relict mangroves on an elevated

400

area at c. 300 m altitude where, at no time

since the last interglacial has sea level been

where mangroves are now found (Woodroffe

1988: 12). Christmas Island has been rapidly

uplifted during the Cainozoic pushing

tertiary limestone to 361 m above sea level

(Woodroffe 1988: 12).

Phenology : Flowers in cultivation have

been recorded in August and October; fruit

has been recorded from the Jardine River in

October and in cultivation in May.

Typification: See previous notes under

Cynometra iripa.

Notes: Previously Cynometra ramiflora

was thought not to occur in Australia, but

specimens from Cape York and Christmas

Island are confirmed to be this species.

In the past, C. ramiflora has been

distinguished from C. iripa by the glabrous

inner wall of its ovaries (Knaap-Van Meeuwen

1970: 14; Tomlinson 1986: 253) (those of C.

iripa are pubescent). However, one collection

(Cooper 2245 ) has sparse but distinct pale

hairs on the inner wall of some, but not all

ovaries.

Sepals have been variously described as

being completely hairy, with a few hairs near

their tip or glabrous (Knaap-van Meeuwen

1970: 24, Ding Hou et al. 1996: 606).

Australian material has glabrous sepals with

an entire to slightly fimbriate apex margin.

With the exception of a small tuft

of caducous hairs at the petiole apex,

all specimens seen from the Australian

mainland and New Guinea have glabrous

petioles, rachises and petiolules (including on

tender new growth), whereas material from

Christmas Island and SE Asia have glabrescent

petioles and rachises, and minutely pubescent

petiolules.

Etymology : The specific epithet, ramiflora ,

is derived from the Latin rami- (pertaining to

branches) and -florus (flowered), referring to

the ramiflorous inflorescences.

3. Cynometra roseiflora W.E.Cooper sp.

nov. Similar to Cynometra ramiflora L. but

differs in the colour of new flush foliage (red

Austrobaileya 9(3): 393-403 (2015)

versus pink); petal length (about half as long

as sepals versus of similar length); petal colour

(bright pink versus white); internal ovary

wall (glabrous but for a tuft of hairs at base

versus glabrous or sparsely hairy but lacking

a tuft of hairs at base); fruit shape (reniform

and laterally compressed versus globose

and ventrally flattened). Typus: Australia:

Queensland. Cook District: Mossman Gorge

section, Daintree National Park, north side of

the river, 10 March 2013, W. Cooper 2215 , T.

Hawkes, R. Jensen , J. Kemp & J. Leech (holo:

CNS [2 sheets + spirit]; iso: BRI, CANB, L,

K, MO distribuendi).

Cynometra iripa (in part) ( sensu Pedley 2007:

39).

Cynometra ramiflora (in part) {sensu Hyland

etal. 2003, 2010).

Shrub or tree to 15 m. Bark with round or

elongated lenticels or pustules; twigs zig¬

zag, with lenticels round and scattered;

stipules not seen. Leaves 1-jugate; petioles

4-8 mm long, not channelled, mostly

enclosed by leaflet base, glabrous; leaflets

slightly discolorous, new growth bright red

and pendulous; petiolules 1-2.5 mm long,

wrinkled, glabrous; leaflets asymmetrically

ovate, 70-175 x 20-62 mm, membranaceous-

coriaceous, glabrous, upper-side shiny, under¬

side dull; base oblique, cuneate, attenuate or

rounded; apex acuminate or drawn out with a

bluntly rounded tip, rarely emarginate; margin

entire; venation brochidodromous, primary

vein raised on both surfaces; secondary veins

8-15 per leaflet, angle of divergence from

primary vein 20-40°; tertiary veins reticulate.

Inflorescence a ramiflorous, axillary or rarely

pseudo-terminal, 1-7-flowered fascicle or

pedunculate raceme on a swelling; pedicels

3-4 mm long, glabrous; bracteoles 2 on each

pedicel, caducous, not seen but evidenced

by scars. Flowers not fragrant, erect, c. 7

x 6 mm; receptacle shortly cone-shaped c.

0.5 mm long and wide; sepals 4, lanceolate

or oblong-elliptic, reflexed and strongly

incurved at apex, 3-4 x 1-2.5 mm, bright

pink, some becoming whitish after anthesis,

glabrous, margin at apex often sparsely and

minutely ciliate; petals 5, lanceolate or strap¬

shaped, entire, 1-2 x 0.2-0.7 mm, bright pink,

Cooper, Cynometra in Australia

caducous; stamens 9-10; filaments terete,

c. 4.5 mm long; anthers c. 0.75 x 0.75 mm,

white; ovary inserted slightly excentrically on

a short stipe or sessile, c. 1.5 x l mm, pink,

pubescent externally, glabrous internally

except for a tuft of hairs at base; ovule 1; style

slender, elongate, initially in line with dorsal

suture, becoming excentric after anthesis,

c. 4.5 mm long, sparsely hairy from base to

apex, stigma minutely capitate. Fruit on a

2-5 mm long pedicel, a reniform or oblong

and laterally compressed nut, with a small

beak at apex of dorsal suture, 28-55 x 19.5—

31.5 x 13.5-23.5 mm, rugose, scurfy, sparsely

and minutely pubescent, rust brown coloured;

seeds 1 per fruit, testa thin and adhering to

mesocarp. Germination epigeal. Figs. 1C,

2E-G, 4

Additional selected specimens (from 8 examined ):

Queensland. Cook District: N bank of Mossman River,

Mossman, Oct 1992, Russell s.n. (BRI [AQ548293]);

Mossman Gorge NP, north side of the river, Jul 2013,

Fig. 4. Cynometra roseiflora showing red new leaves

(Cooper 2215 etal. [CNS]). Photo: R. Jensen.

Cooper 2223, Jensen, Jago & Russell (CNS); NPR 133,

Mossman Gorge, Jul 1995, Hyland 25906 RFK & Gray

(CNS); NPR 133, Mossman Gorge, Dec 1995, Hyland

25906 RFK & Gray (CNS); Mossman River, Silky Oaks

Resort, Jul 1992, Sankowsky 1333 (CNS); Mossman

Gorge, Silky Oaks Resort, May 1993, Sankowsky 1417

(CNS); Cultivated by G & N Sankowsky at Tolga, Nov

2014, Cooper 2271 & Sankowsky (CNS).

401

Distribution and habitat: Cynometra

roseiflora is endemic to the Wet Tropics

bioregion in north-eastern Queensland

where it is currently known to occur within

a very small area on the northern side of

the Mossman River in the Mossman Gorge

section of the Daintree National Park and

on the neighbouring property of Silky Oaks

Lodge (Map 1), altitude 20-125 m. It grows

in wet lowland rainforests (mesophyll vine

forest) on soils derived from granite. Plants

that co-occur with C. roseiflora include

Alstonia scholaris (L.) R.Br., Archidendron

ramiflorum (F.Muell.) Kosterm., Backhousia

bancroftii F.M.Bailey & F.Muell. ex

F.M.Bailey, B. hughesii C.T.White, Calamus

australis Mart., Calamus moti F.M.Bailey,

Cardwellia sublimis F.Muell., Citronella

smythii (F.Muell.) R.A.Howard, Dysoxylum

arborescens (Blume) Miq., D. papuanum

(Merr. & L.M.Perry) Mabb., D. pumilum

Mabb.; Lindsayomyrtus racemoides (Greves)

Craven, Medinilla balls-headleyi F.Muell.,

Mesua larnachiana (F.Muell.) Kosterm.,

Myristica insipida R.Br. var. insipida and

Palaquium galactoxylon (F.Muell.) H. J.Lam.

Phenology: Flowers have been recorded in

March; fruits have been recorded in June-

July.

Etymology: The specific epithet, roseiflora ,

is derived from the Latin roseus (pink) and

-florus (flowered).

Acknowledgements

I am grateful to Frank Zich and Darren Crayn

for support and access to the ATH herbarium;

Bill Cooper, Paul Forster, Tim Hawkes, Bob

Jago, Rigel Jensen, Jeanette Kemp, James

Leech, Dion Maple, Chris & Ian Parker, Tony

Roberts, Rupert Russell, Garry Sankowsky,

Brian Venables and Christina Zdenek for

valuable assistance with collections, Paul

Forster for advice and typification, Peter

Bostock for etymology and Steve Murphy for

the map. Permits to collect were issued by the

Queensland Department of Environment and

Resource Management and the Australian

Government Director of National Parks.

402

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Map 1 . Distribution of Cynometra iripa Afwithin Australia), C. ramiflora + (within mainland Australia), and C.

roseiflora •.

Olearia cuneifolia A.R.Bean & M.T.Mathieson

(Asteraceae: Astereae), a new species from Queensland

A.R. Bean & M.T. Mathieson

Summary

Bean, A.R. & Mathieson, M.T. (2015). Olearia cuneifolia A.R.Bean & M.T.Mathieson (Asteraceae:

Astereae), a new species from Queensland. Austrobaileya 9(3): 404-407. Olearia cuneifolia

A.R.Bean & M.T.Mathieson is described, illustrated and compared to related taxa. It has a restricted

distribution in the Mungallala area of southern Queensland. A conservation status of Endangered is

recommended.

Key Words: Asteraceae, Olearia , Olearia cuneifolia , endangered species, Maranoa, Queensland flora

A.R.Bean & M.T.Mathieson, Queensland Herbarium, DSITI, Brisbane Botanic Gardens, Mt Coot-

tha Road, Toowong, Queensland 4066, Australia. Email: Tony.Bean@dsiti.qld.gov.au

Introduction

Olearia Moench with 122 indigenous species

is currently Australia’s most species diverse

Asteraceae genus (APC 2015). The molecular

study by Cross et al. (2002) showed that

Olearia is polyphyletic, with some species

appearing in clades with species from other

genera of the tribe Astereae. They identified

two major taxonomic groups for Olearia ,

‘primary clade I’ and ‘primary clade II’,

to which all species could be assigned. The

study also revealed the presence of eight

robust clades within Olearia , which they

designated A-H. It is possible that the name

Olearia will, after further study, be confined

to Clade A (including the Queensland species

O. ramulosa (Labill.) Benth., O. microphylla

(Vent.) Maiden & Betche and O. nernstii

(F.Muell.) Benth.); otherwise all members of

primary clade I will retain the generic name of

Olearia. Messina et al. (2014) have provided

a revision of the species included in Olearia

sect. Asterotriche Benth., a monophyletic

subset of the Clade A of Cross et al. (2002).

Olearia cuneifolia , newly named here,

belongs to Clade B of Cross et al. (2002) and

is related to O. magniflora (F.Muell.) Benth.,

O. muelleri (Sond.) Benth. and O. calcarea

F.Muell. ex Benth.

Accepted for publication 23 July 2015

Materials and methods

This study is based on the morphological

examination of specimens held at BRI, together

with field observations. The measurements for

floral parts are based on material preserved in

70% alcohol or reconstituted with hot water;

other plant parts were measured from dried

material.

Taxonomy

Olearia cuneifolia A.R.Bean &

M.T.Mathieson sp. nov. with affinity to O.

muelleri , but differing by the cuneate leaves,

the much longer involucres, the disc florets

with hairs on the corolla tube and corolla

lobes, and the two-whorled pappus. Typus:

Queensland. Maranoa District: Nalpa

Downs, c. 16 kmNE of Mungallala, 26 March

2015, M.T. Mathieson MTM1999 (holo: BRI;

iso: CANB, K, MEL, NSW, US, distribuendi).

Erect shrub to 2 m high. Branchlets very

viscid and with sparse, erect eglandular hairs

to 0.15 mm long. Young branchlets distinctly

angular due to decurrent leaf-bases, but older

branchlets terete. Leaves alternate, sessile,

oblanceolate to cuneate, 8-15 x 2-5.2 mm, ±

glutinous, ± concolorous, venation not visible,

except midrib; glabrous or with short sparse

erect eglandular hairs, mainly along margins;

apex acute or truncate; base attenuate;

margins flat or recurved, entire or with 2

small teeth near apex. Capitula terminal,

Bean & Mathieson, Olearia cuneifolia

solitary, sessile. Involucres ellipsoidal at

anthesis, 14-16 mm long, 7-9 mm diameter;

campanulate to hemispherical at fruiting

stage. Bracts 5-7-seriate; outer bracts 3-4

x 1.8-2.5 mm, elliptical, with ± dense

eglandular hairs near distal end; margins

irregularly ciliate; inner bracts lanceolate,

6-10 x 1.5-2 mm, inner surface glabrous,

outer surface with eglandular hairs near

apex; apex obtuse, margin irregularly ciliate

near apex. Receptacle slightly to markedly

alveolate, 3-3.8 mm diameter. Ray florets

14-21, female; corolla tube linear, 6.2-6.9

mm long, with small antrorse eglandular

hairs on apical % of tube; ligule 7.5-9.5 mm

long, white; styles exserted, recurved. Disc

florets 28-42, bisexual; corolla tube linear,

6.5-7.5 mm long, with antrorse eglandular

hairs on medial section; corolla lobes 5, acute,

0.8-1.6 mm long, glabrous except for a cluster

of small eglandular hairs near apex; anthers

1.2-1.4 mm long, with sterile tip 0.8-1 mm

long, anther tails c. 0.1 mm long. Achenes

flattened-cylindric, 4-6-ribbed, 3.8-4.2 mm

long, densely silky-hairy throughout; pappus

bristles barbellate, creamy-white, in two

whorls; inner whorl 7.5-8 mm long at fruiting

stage, the outer whorl 1-1.5 mm long. Figs.

1-3

Additional specimens examined : Queensland.

Maranoa District: Lot 23, CP847082 Mitchell

8445 - Morven 8346 [Umberill Station], Sep 2003,

Baumgartner s.n. (BRI [AQ764458]); Nalpa Downs,

WNW of Mitchell, Dec 2013, Mathieson MTM1599

(BRI); Nalpa Downs, c. 40 km NW of Mitchell, Sep

2014, Mathieson MTM1790 (AD, BRI, HO, NE, NT);

Andromeda, Mar 2015, Mathieson MTM2000 (BRI,

NSW); Andromeda, N of Mungallala, May 2008, Silcock

125 (BRI); Andromeda, c. 24 km NNE of Mungallala,

Oct 2008, WangJW0170( BRI).

Distribution and habitat : Olearia cuneifolia

is endemic to Queensland. The species is

known from sites north and north-east of

Mungallala, between Roma and Charleville.

It occurs within the ecotone between open

or degraded forests dominated by Acacia

harpophylla F. M uel 1. exBenth. and Casuarina

cristata Miq. and open sclerophyll woodland

dominated by Eucalyptus crebra F.Muell.

and/or E. thozetiana (Maiden) R.T.Baker. The

soils are derived from sedimentary rocks and

405

consist of self-mulching cracking clays on flat

areas or stony clays on lower slopes of low

mesas and rises.

Phenology : Flowers and fruits are recorded

in March, May, September and October.

Affinities : Olearia cuneifolia is related to

O. muelleri, O. magniflora and O. calcarea.

All of these species have solitary and sessile

capitula, and the involucres are cylindrical to

ellipsoidal at anthesis. All have small resinous

leaves.

Olearia cuneifolia differs from O. muelleri

by the narrower, more cuneate-shaped leaves

(broadly-ovate to orbicular in O. muelleri ), the

much larger involucres, the greater number of

disc and ray florets, the presence of hairs on

the corolla tube and lobes of the disc florets

(glabrous in O. muelleri ), the longer achenes

and the pappus with two whorls of bristles

(one whorl in O. muelleri).

Olearia cuneifolia differs from O.

magniflora by the leaves without teeth or

with only two teeth (at least some leaves

with four or more teeth in O. magniflora ), the

shorter rays of the ray florets, the rays white

in colour (mauve to purple in O. magniflora ),

and the densely hairy achenes (glabrous in O.

magniflora).

Olearia cuneifolia is similarly in leaf

morphology to O. calcarea , but differs from

O. calcarea by the longer involucres, the

greater number of ray florets, the much shorter

rays of the ray florets, and the shorter achenes

and pappus. O. calcarea is often regarded as a

hybrid between O. magniflora and O. muelleri

(Walsh & Lander 1999). O. cuneifolia is not

considered to be a hybrid as other species of

Olearia in the area (i O. canescens (Benth.)

Hutch., O. elliptica DC., O. gordonii Lander,

O. subspicata (Hook.) Benth.) are not related

to it.

Conservation status: Olearia cuneifolia is

only known from three locations in a small

area (approximately 19.5 km 2 ) to the north of

Mungallala in south central Queensland. It has

not been located elsewhere in the surrounding

district despite many surveys in similar

406

Austrobaileya 9(3): 404-407 (2015)

Fig. 1. Olearia cuneifolia. A. flowering branchlet *1. B. outer involucral bract *8. C. inner involucral bract x8. D.

mature achene and pappus x8. E. disc floret ><16. F. ray floret *8. All from Mathieson MTM1999 (BRI). Del. W. Smith.

Bean & Mathieson, Olearia cuneifolia

407

Fig. 2. Lateral view of young flowering capitulum

(Mathieson MTM1999). Photo: M.T. Mathieson.

Fig. 3. Disc and ray florets on young flowering capitulum

(Mathieson MTM1999). Photo: M.T. Mathieson.

References

Apc (2015). Australian Plant Census. Council of Heads

of Australasian Herbaria, https://www.anbg.

gov.au/chah/apc/, accessed 23 May 2015.

Cross, E.W., Quinn, C.J. & Wagstaff, S.J. (2002).

Molecular evidence for the poly phyly of Olearia

(Astereae: Asteraceae). Plant Systematics and

Evolution 235: 99-120.

habitat. The total population is estimated to

be < 250 individuals occupying a total area of

less than two hectares. Applying the criteria

of the IUCN (IUCN 2012), the recommended

conservation status is Endangered (D2).

Etymology : The specific epithet is given in

reference to the leaf shape.

Acknowledgements

We are grateful to Will Smith (BRI) for the

line drawings.

Iucn (2012). IUCN Red List Categories and Criteria,

version 3.1, 2nd ed. https://portals.iucn.org/

library/efiles/documents/RL-2001-001-2nd.pdf,

accessed 29 September 2014.

Messina, A., Walsh, N.G., Hoebee, S.E. & Green,

P.T. (2014). A revision of Olearia section

Asterotriche (Asteraceae: Astereae). Australian

Systematic Botany 27: 199-240.

Walsh, N.G. & Lander, N.S. (1999). Olearia. In N.G.

Walsh & T.J. Entwisle (eds ). Flora of Victoria,

Dicotyledons Cornaceae to Asteraceae 4: 886-

912. Inkata Press: Melbourne.

Eremophila woodiae Edginton (Scrophulariaceae),

a new species from Queensland

M.A. Edginton

Summary

Edginton, M.A. (2015). Eremophila woodiae Edginton (Scrophulariaceae), a new species from

Queensland. Austrobaileya 9(3): 408-415. Eremophila woodiae is described and illustrated. Notes

on ecology, distribution and conservation status are provided. The known distribution of the new

species is restricted to the vicinity of Vergemont Station and Opalton in western central Queensland.

A conservation status of Least Concern is recommended.

Key Words: Scrophulariaceae, Myoporaceae, Eremophila , Eremophila woodiae , new species,

taxonomy, conservation status, Vergemont Station, Opalton

M.A. Edginton, Queensland Herbarium, Department of Science, Information Technology and

Innovation, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia.

E-mail: Mark. Edginton@dsiti.qld.gov. au

Introduction

Eremophila R.Br. is a genus of 215 species

and is endemic to Australia (Chinnock 2007).

The genus is widespread throughout the

arid areas of the continent (although some

species do occur outside arid areas, albeit

less commonly) with the majority of species

restricted to Western Australia. Until recently,

Eremophila was considered to be part of the

family Myoporaceae, a southern hemisphere

family. A review by Tank et al. (2006) based

on molecular studies supported the inclusion

of the traditional Myoporaceae within the

Scrophulariaceae, an inclusion which has now

been accepted at BRI and elsewhere.

A new species of Eremophila , restricted

to western central Queensland, is described

in this paper. Herbarium material was first

collected in 1986 by Neldner and Nicholson,

then in 1987 by Mitchell.

Material and methods

This study is based upon the examination

of dried herbarium material and label data

from the Queensland Herbarium. All existing

specimens were examined. No spirit material

had been collected, although parts rehydrated

Accepted for publication 27 August 2015

to facilitate drawing have subsequently been

put into spirit. Unless stated otherwise all

photographs were taken by the author.

Taxonomy

Eremophila woodiae Edginton sp. nov.

Typus: Queensland. Gregory North District:

Vergemont, W of Longreach, 1 September

2013, R. Fensham 6372 (holo: BRI).

Eremophila sp. (Opalton V. J. Neldner+ 2619);

Edginton (2014).

Small shrub to 0.5 metres; branches terete,

often exhibiting remaining basal portions of

leaves, resinous on new growth, becoming

non-resinous with age, non-tuberculate,

red-brown to yellow, becoming grey with

age; indumentum on new branches, leaves,

petioles, pedicels and sepals moderately dense

with sessile and short spreading colourless

glandular hairs 0.1-0.4 mm long, and longer

villous glandular and eglandular colourless

hairs 0.6 -1.1 mm long, branches glabrescent.

Leaves spirally arranged, densely crowded

towards the branch tips, spreading to

ascending, linear-lanceolate, conduplicate in

distal three-quarters and flattened below, 12-

22 mm long, 1.5-3 mm long, resinous, mid to

dark green; margins mainly entire apart from

distinct lobing towards apex with 3-5 lobes

per margin directed upwards, apex obtuse to

sub-acute; base narrowing to a petiole-like

Edginton, Eremophila woodiae

section 0.5-1 mm long formed by the central

vein. Flowers 1 per axil; pedicels semi-terete,

1-2 mm long, dark red-brown to yellow in

dried specimens, becoming grey with age.

Sepals 5, slightly imbricate, 12-14.5 mm long,

resinous, acute; differentiated into 1 posterior,

2 anterior and two inner ones; posterior sepal

ovate-lanceolate, 5-6 mm wide; anterior

sepals broadly triangular-lanceolate, 3-4 mm

wide; inner sepals triangular-lanceolate, 2-3

mm wide. Corolla ± zygomorphic, bilabiate,

14-19 mm long, violet to light purple,

unspotted; tube 11-14 mm long, bulbous

at base, slightly constricted above and then

dilating gradually distally; upper lip 4-5 mm

long, 2 lobed, lobes obtuse, divided for c. Vie

of length; lower lip 4-5 mm long, 3-lobed, the

lobes divided for most of their length, medial

one obtuse, slightly longer and broader than

the lateral ones which are obtuse to subacute;

outer surface of upper tube and lobes sparsely

to moderately villous, hairs colourless,

eglandular, constriction and lowermost part of

tube glabrous; inner surface of lobes glabrous;

tube prominently woolly in throat just below

lobes of upper lip, otherwise villous in upper

part, glabrous in lower part, hairs eglandular.

Stamens 4; anthers 2-2.5 mm long, lower 2

enclosed, upper two barely exserted from

throat, glabrous; filaments sparsely villous

with white eglandular hairs near base,

glabrous above; ovary conical-ovoid, slightly

to moderately laterally compressed, 2-3 mm

long, 1-1.5 mm wide, glabrous, 4-locular; 3

ovules per locule; style c. 10 mm, sparsely

villous with white eglandular hairs. Fruit

(mature) ovate-conical, 5-8 x 3.5-5.5 mm,

exocarp papery, whitish, glabrous; endocarp

woody, slightly compressed, subreticulately

ribbed distally, red-brown; immature fruit

enclosed by sepals. Figs. 1-5.

Additional specimens examined: Queensland. Gregory

North District: 18 km SW of Opalton, Nov 1986,

Neldner 2619 & Nicholson (BRI). Mitchell District:

25 km SW of Vergemont Station, W of Longreach, Sep

1989, Mitchell 463 (BRI); Vergemont Station, May 2011,

Fensham 6101 & Silcock (BRI); Vergemont Station, W

of Longreach, Sep 2013, Fensham 6376 (BRI), 6379

(BRI), 6380 (BRI).

Distribution and habitat : Eremophila

woodiae is endemic to western central

Queensland, in the vicinity of Opalton and

409

Vergemont Stations west of Longreach and

south of Winton (Map 1).

The species occurs on barren plateaux,

especially near the edges, in skeletal soil with

sparse, stunted shrubland. There is one record

from the base of a jump-up in “boree ( Acacia

tephrina) and spinifex ( Triodia sp./spp.) flats”

(Neldner 2619 & Nicholson). Associated

species commonly include Acacia shirleyi

Maiden, A. catenulata C.T.White, A. sibirica

S.Moore, Corymbia blakei K.D.Hill &

L.A.S. Johnson, Eremophila latrobei F.Muell.,

Hakea collina C.T.White, Triodia longiceps

J.M.Black and T. pungens R.Br. (Silcock

& Fensham 2014; Queensland Herbarium

HERBRECS data 2015).

Phenology : The specimens collected indicate

that flowering and fruiting occurs between

May to November.

Affinities: Eremophila woodiae bears

a superficial resemblance to E. hispida

Chinnock, which has a known distribution

which is close to, but does not quite overlap,

that of E. woodiae. Both taxa are low shrubs

with leaves which appear linear when

viewed without a microscope, and both

species have violet to purple flowers 15-20

mm long. Eremophila woodiae has leaves

which are linear-lanceolate when viewed

microscopically (linear on E. hispida ); very

crowded leaves (not crowded on E. hispida),

simple hairs only on all parts (simple hairs,

with or without some stellate hairs on

branchlets, leaves and sepals on E. hispida ),

and the top % to Vz of each leaf has lobes

(entire leaves on E. hispida). Despite the

superficial resemblance, E. woodiae and E.

hispida are unlikely to be close relations.

Eremophila woodiae can be classified

in Eremophila section Eremaeae Chinnock

based on the species description (Chinnock

pers. comm., July 2015). Chinnock further

stated that the corolla was interesting and

appeared slightly different (based on the

single image of a flower seen) in having two

obtuse upper lobes; however, as there can be

variation in the size of the flowers and the

arrangement of the lower lobes in species

of this section more flowers from different

410

Austrobaileya 9(3): 408-415 (2015)

QUEENSLAND HERBARIUM (BRI)

Flora of Queensland Mitchell

Eremophila

QUEENSLAND HERBARIUM (BRI)

Brisbane Australia

AQ 876335

Coll. R.J. Fensham 6372

1 SEP 2013

23d 30m 07s 142d48m 15s [GDA94] Depth m

(54,684224,7399756) (7450-842997) Alt. m

Vergemont, W of Longreach.

Barren ferricrete plateau-top.

Shrub to 0.5m, blue flower; about 2500 plants over 50ha of habitat.

Photo with specimen at BRI.

Queensland Herbarium (BRI)

5p . rT-rJWw

Det. 303 Myoporaceae

Dup.

* May be cited as computerised collection Number AQ 876335

(Archival Paper)

Fig. 1. Holotype of Eremophila woodiae (.Fensham 6372, BRI).

Edginton, Eremophila woodiae

411

Fig. 2. Eremophila woodiae. A. branching habit *2. B. leaf *8. C. close-up of leaf showing glandular and eglandular

hairs x24. D. front view of corolla x6. E. side view of calyx and corolla x4. F & G. inner and outer surfaces of posterior

sepal x4. H. cross-section of posterior sepal showing glandular and eglandular hairs xl6. All from Fensham 6379

(BRI). Del. W. Smith.

412

Austrobaileya 9(3): 408-415 (2015)

Fig. 3. Flower, branch and leaves of Eremophila woodiae at Vergemont Station. Photo: R. Fensham.

populations may show greater variation.

Otherwise all other aspects of the species fall

well within the boundaries of Eremophila

section Eremaeae. It may be related to E.

goodwinii F.Muell. because of the glandular

indumentum, although unlike this species it

does not have the banding on the inside lateral

walls of the corolla (Chinnock pers. comm.,

July 2015).

Notes : In one collection of Eremophila

woodiae (Mitchell 463), a flower with 6

sepals was noted (Fig. 5). All other flowers

examined during this study had 5 sepals so it

is presumed that 6 sepals is a rare exception.

None of the fruits observed had completely

shed the exocarp - though some had shed

most of it; consequently it is possible that

none of the fruits were completely mature.

Therefore 1 mm was added arbitrarily to the

highest measured values for length and width,

and these augmented values were used as the

high end of the ranges for length and width.

Conservation status : The species known

range is very restricted; however, it is

locally common in places. A species profile

(as Eremophila sp. (Opalton V.J. Neldner+

2619) by Silcock & Fensham (2014) found

that this species occurred on 6 of 10 plateaux

surveyed in the overall area of distribution.

Population sizes on each of these plateaux

varied from 1000 to 4000 plants, with a total

of about 17000 plants found. Ninety km 2 of

similar plateaux were mapped in its known

range, and of this 5 km 2 was surveyed. It was

estimated that the total population based on

known occurrences is approximately 300,000

individuals. However, Silcock & Fensham

Edginton, Eremophila woodiae

413

Fig. 4. Branch portion of Eremophila woodiae showing raised persistent leaf base remnants and glandular and

eglandular hairs.

Fig. 5. Six-sepal flower of Eremophila woodiae ( Mitchell 463 , BRI).

414

Austrobaileya 9(3): 408-415 (2015)

(2014) concluded that there is a lot of suitable

habitat to the west and south of its known

range, so the total population may be higher

than 300,000.

The habitat in which Eremophila woodiae

occurs is without domestic stock and has

low feral goat numbers; conversely grazing

by native animals is thought to be very low.

No evidence of grazing on the species was

observed. Silcock& Fensham (2014) concluded

that there are no threats to this species, that it

is abundant within its limited range and that

there are no observable fluctuations in the

number of plants over the short term. They

ascertained that the species is not eligible for

listing under any IUCN criteria. Therefore a

Least Concern conservation status under the

Queensland Nature Conservation Act 1992 is

recommended.

Etymology : The specific epithet of this

species is named for Aileen Wood, a long

term staff member at the Queensland

Herbarium. Aileen’s knowledge of cultivated

plants, especially cultivars, is unsurpassed

at the Queensland Herbarium and is greatly

appreciated by staff.

Acknowledgements

I wish to thank the following staff at the

Queensland Herbarium who assisted me in

the preparation of this paper: Rod Fensham for

collecting material to facilitate description,

and for reading an early manuscript draft

and making instructive suggestions; Ailsa

Holland for reading an early manuscript draft

and making instructive suggestions; Tony

Bean for the distribution map, methodology

advice and for reading a late manuscript draft

and making instructive suggestions; Will

Smith for the distribution map, drawings

and assistance with photographs. I also wish

to thank Bob Chinnock for reviewing the

manuscript and making useful comments.

References

Chinnock, R. (2007). Eremophila and Allied Genera: A

Monograph of the plant family Myoporaceae.

Rosenberg Publishing: Sydney.

Edginton, M.A. (2014) Scrophulariaceae. In P.D.

Bostock & A.E. Holland (eds.) (2014). Census

of the Queensland Flora 2014. Queensland

Department of Science, Information Technology

and Innovation: Brisbane, https://www.qld.

gov.au/environment/assets/documents/plants-

animals/herbarium/qld-f lora-census.pdf,

accessed 1 December 2014.

Silcock, J. & Fensham, R. (2014). Eremophila sp.

(Opalton V.J. Neldner+ 2619) species profile.

Unpublished.

Tank, D.C., Beardsley, P.M., Kelcher, S. A. & Olmstead,

R.G. (2006). L.A.S. Johnson Review No. 7.

Review of the Systematics of Scrophulariaceae

s.l. and their current disposition. Australian

Systematic Botany 19: 289-307.

Edginton, Eremophila woodiae

415

Map 1. Distribution of Eremophila woodiae , grey shaded areas are conservation reserves.

Rhynchospora croydonensis R.Booth (Cyperaceae),

a new species from northern Queensland

R. Booth

Summary

Booth, R. (2015). Rhynchospora croydonensis R.Booth (Cyperaceae), a new species from northern

Queensland. Austrobaileya 9(3): 416-420. A new species, Rhynchospora croydonensis R.Booth, is

described and illustrated and notes are provided on its distribution and habitat. A key to Queensland

species of Rhynchospora Vahl. is provided.

Key Words: Cyperaceae, Rhynchospora , Rhynchospora croydonensis, Australia flora, Queensland

flora, new species, taxonomy, identification key

R.Booth, Queensland Herbarium, Department of Science, Information Technology and Innovation,

Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email: Ron.

Booth@dsiti.qld.gov.au

Introduction

Rhynchospora Vahl. is a large, cosmopolitan

genus consisting of more than 300 species,

mostly occurring in the tropics and subtropics

of the Central and South Americas (Strong

2006). Only a few species are widely

distributed, though most of the Australian

species also occur in southeast Asia (Kern

1974). Australia has about twenty one

species, with fourteen of these occurring in

Queensland (Bostock & Holland 2014).

Bentham & Hooker (1883) separated

the genus Rhynchospora into two informal

sections, “ Haplostylae ” and “ Diplostylae ”

based on the degree of stylar branching

and this was followed by Kiikenthal (1949,

1952). Thomas et al. (2009) constructed

a preliminary molecular phylogeny of

the genus that indicated the two informal

sections in Rhynchospora are artificial. A full

phylogenetic study of the genus with greater

sampling from Australia and Asia is still

needed.

According to the classification of

Goetghebeur (1986), Rhynchospora may be

included within the subfamily Cyperoideae ,

tribe Rhynchosporeae. However, Bruhl (1995)

indicated that the tribe Rhynchosporeae is

part of the subfamily Caricoideae. Muasya

et al. (2009) found good molecular support

for the recognition of tribe Rhynchosporeae

in an isolated position near tribe Cariceae

and Scirpeae, with the Rhynchosporeae

consisting solely of the genus Rhynchospora

(including Pleurostachys Brongn.).

Sharpe (1989) provided a key and

descriptions for four species in southeast

Queensland, but the most comprehensive

set of descriptions remain those provided

by Kiikenthal (1949, 1952). Due to the

extensive field work undertaken in recent

years, particularly in north Queensland, more

Rhynchospora material has become available

for study at the Queensland Herbarium.

Critical examination of these collections by

the present author has now enabled this new

species to be formally described and named.

Materials and methods

All herbarium specimens of Rhynchospora

held at BRI have been examined.

Measurements were made from dried

material.

Accepted for publication 23 July 2015

Booth, Rhynchospora croydonensis

Taxonomy

Rhynchospora croydonensis R. Booth, sp.

nov. similar to Rhynchospora pterochaeta

F.Muell. but differs in the achene surface

that has scattered hispid hairs (versus densely

papillose), and hairs on the hypogynous

bristles at the very base of the bristles (versus

densely pilose for half their length). Typus:

Queensland. Burke District: 6.5 km along

Coralie Road from Croydon - Claraville

Road junction, 22 April 2011, K.R. McDonald

KRM10937 (holo: BRI [AQ759401]; iso:

NSW).

Rhynchospora sp. Croydon (S.L.Everist

5384); R.Booth in Bostock & Holland (2014).

Annual or short lived perennial 20-45 cm

tall; culms tufted, 3-angled, trigonous,

striate, 0.6-1.3 mm wide, glabrous. Lamina

flat or conduplicate, 1-3 mm wide with long

ciliate hairs when young, glabrescent. Ligule

membranous. Involucral bracts ciliate on

the margins, longer than the inflorescence.

Inflorescence a single dense head, 1.2-1.5 cm

long by 2-2.5 cm wide. Spikelets lanceolate or

linear-lanceolate, 7.5-10 mm long, pale brown

to brown. Perianth present as 6 hypogynous

bristles, unequal in length, longer than the nut

and style base, antrorsely scabrid, plumose at

the base, glabrous above. Glumes distichous,

6-7, narrowly ovate or ovate-lanceolate, 2-10

mm long, 1.2-1.8 mm wide, coloured straw or

light brown to brown, glabrous, keeled or with

the midrib distinct, not nerved; apex acute;

margin glabrous; rachilla wingless. Style

deciduous, entire, glabrous. Achene with a

persistent style base, not grooved, 2.4-3 mm

long, as broad or broader than the achene and

longer than, or equalling the achene; margins

winged, setulose; stramineous, or yellow-

brown. Achene obovoid, biconvex, lenticular,

or concave-convex; 2.2-3 mm long, 1.3-1.6

mm wide, not ribbed; surface smooth or with

417

a faint cell pattern, with hispid hairs mainly in

the distal half, pilose hairs only at the base of

the achene. Fig. 1.

Additional specimens examined : Queensland. Burke

District: Cothilda Station, May 2001 Turpin GPT872

(BRI); between Croydon and Gilbert River, May 1954,

Everist 5384 (BRI). Cook District: Staaten River

NP, c. 35 km W of Bulimba Homestead, 150 km W

of Chillagoe, Sep 2013, Leitch ODA004939 (BRI);

Fishermans Waterhole, Walsh River, N of Chillagoe,

Mar 2005, McDonaldKRM3936a (BRI); Bulleringa NP,

Jul 2006, Lovatt TH9249 (BRI); 79 km from Georgetown

on Croydon Road, Apr 1996, Forster PIF19012b & Ryan

(BRI); 23 miles [38.3 km] SE of Croydon township, Jul

1954, Speck 4719 (CANB).

Distribution and habitat: Rhynchospora

croydonensis is endemic to northern

Queensland where it has been found between

Croydon and the Gilbert River in the south,

Staaten National Park in the north, and west

to Bulleringa National Park (Map 1). Plants

usually grow in drainage lines in sandy soils

often with Melaleuca viridiflora Sol. ex

Gaertn. var. viridiflora.

Affinities: Rhynchospora croydonensis is

closely related to R. pterochaeta , which it

differs from most obviously in the achene

characters: R.pterochaeta being papillose over

much of its surface, while R. croydonensis

has hispid hairs, mainly in the distal part of

the achene.

Conservation status: There are only nine

collections of Rhynchospora croydonensis

and its geographic range is not large. This

sedge has been found in Bulleringa and

Staaten River National Parks and is likely to

be widespread in its seasonally ephemeral

habitat dominated by annual or geophytic

species. It is not considered to be threatened

and a Least Concern conservation status is

recommended.

Etymology: The epithet is from the town

of Croydon in the Gulf of Carpentaria near

where this species was first collected.

418

Austrobaileya 9(3): 416^-20 (2015)

Fig. 1. Rhvnchospora croydonensis. A. habit *0.4. B. inflorescence *3. C. achene and hypogynous bristles xl2. All

from McDonald KRM10937 (BRI).

Booth, Rhynchospora croydonensis 419

Key to the Queensland species of Rhynchospora

1 Inflorescence a compound panicle or clustered in loose corymbs.2

1. Inflorescence a single, dense head.4

2 Stems filiform, < 1 mm wide.R. gracillima

2. Stems stout, 2-15 mm wide.3

3 Inflorescence a large, loose, compound panicle; style base furrowed on

both sides; stout plants 70-200 cm high.R. corymbosa

3. Inflorescence of several small, irregular corymbs; style base not furrowed;

slender plants to 1 m high, usually less.R. brownii

4 Style base 0.4-0.6 mm long; bristles whitish, shorter than the achene.R. rubra

4. Style base > 0.6 mm long; bristles usually > the achene in length.5

5 Base of the achene and the bristles with no pilose

hairs (hispid hairs may be present on the bristles only).6

5. Base of the achene or at least bottom part of the bristles with pilose hairs.9

6 Achene (not including the style base) < 2 mm long.R. subtenuifolia

6. Achene > 2 mm long.7

7 Spikelets < 6 mm long.R. submarginata

7. Spikelets > 6 mm long.8

8 Style base furrowed or grooved, achene with broad triangular

based hairs at the apex; bristles much longer than the style base.R. exserta

8. Style base not furrowed, achene with rather fine hispid hairs;

bristles only just longer than the achene and style base.R. heterochaeta

9 Achene revolute, papillose on the nearly joined margins.R. leae

9. Achene biconvex or convex, not revolute.10

10 Style base grooved, curved, or winged adjacent to the achene, often

setulose on the margins. 11

10. Style base not grooved, curved or winged.13

11 Style base long-conical with a groove down the middle.R. longisetis

11. Style base as broad or broader than the achene, curved, but not grooved.12

12 Achene papillose (hairs rounded at the ends) over much of its surface,

pilose hairs covering lower half of the achene.R. pterochaeata

12. Achene with hispid hairs (pointed at the ends) mainly in the distal half,

pilose hairs only at the base of the achene.R. croydonensis

13 Achene > 3 mm long, style base at least 2 mm long.R. longisetis

13. Achene < 3 mm long, style base <1.5 mm long.14

14 Achene papillose over most of its surface, bristles unequal in length.R. wightiana

14. Achene hispid or occasionally papillose, mainly in the distal half, bristles

equal in length

R. affinis

420

Acknowledgements

I thank Keith MacDonald for his excellent

collections, Will Smith for the illustrations,

and Tony Bean for his support and helpful

suggestions.

References

Bentham, G. & Hooker, j.D. (1883). Cyperaceae. In

Genera Plantarum ad exemplaria imprimis in

herbariis Kewensibus. 3(1): 1-459. L. Reeve &

Co.: London.

Bostock, RD. & Holland, A.E. (eds.) (2014). Census

of the Queensland Flora 2014. Queensland

Herbarium, Department of Science, Information

Technology and Innovation, https://data.qld.

gov.au/dataset/census-of-the-queensland-

flora-2014, accessed 1 July 2015.

Bruhl, J. (1995). Sedge genera of the world: Relationships

and a new classification of the Cyperaceae.

Australian Systematic Botany 8: 125-305.

Goetghebeur, P. (1986). Genera Cyperacearum. Een

bijdrage tot de kermis van de morfologie,

systematiek en fylogenese van de Cyperaceae

genera. Unpublished Dr Sci. Thesis, State

University: Gent.

Kern, J.H. (1974). Cyperaceae. In C.G.G.J. van Steenis

(ed.). Flora Malesiana Series 1. 7: 435-753.

Wolters-Noordhoff Publishing: Groningen.

Austrobaileya 9 ( 3 ): 416-420 ( 2015 )

Kukenthal, G. (1949). Vorarbeiten zu einer monographic

der Rhynchosporoideae. 18. Rhynchospora

Vahl. Botanische Jahrbiicher fur Systematik,

Pflanzengeschichte und Pflanzengeographie

74: 375-509.

- (1952). Vorarbeiten zu einer monographic der

Rhynchosporoideae. Botanische Jahrbiicher

fur Systematik, Pflanzengeschichte und

Pflanzengeographie 75: 127-195, 273-314,

451-497.

Muasya, A.M., Simpson, D.A., Verboom, G.A.,

Goetghebeur, P, Naczi, R.F.C., Chase, M.W.

& Smets, E.F. (2009). Phylogeny of Cyperaceae

based on DNA sequence data: Current progress

and future prospects. The Botanical Review 75:

2 - 21 .

Sharpe, P. (1989). Cyperaceae. In T.D. Stanley & E.M.

Ross (eds.). Flora of South-Eastern Queensland

3: 277-355. Queensland Department of Primary

Industries: Brisbane.

Strong, M.T. (2006). Taxonomy and distribution of

Rhynchospora (Cyperaceae) in the Guianas,

South America. Contributions from the United

States National Herbarium 53: 1-225.

Thomas, W.W., Araujo, A.C. & Alves, M.V. (2009).

A preliminary molecular phylogeny of the

Rhynchosporeae (Cyperaceae). The Botanical

Review 75: 22-29.

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Map 1. Distribution of Rhynchospora croydonensis , grey shaded areas are conservation reserves.

First record of the Gnetales in Australia:

Gnetum gnemon L. (Gnetaceae) on Badu and

Mua islands, Torres Strait, Queensland

David G. Fell 1 , David J. Stanton, Dick Williams, Frank Loban, Frank Nona,

Troy Stow, John Wigness, Erimiah Manas & Guyai Uiduldam

Summary

Fell, D.G., Stanton, D.J., Williams, D., Loban, F., Nona, F., Stow, T., Wigness, J., Manas E., and

Uiduldam G. (2015). First record of the Gnetales in Australia: Gnetum gnemon L. (Gnetaceae) on

Badu and Mua Islands, Torres Strait, Queensland. Austrobaileya 9(3): 421-430. The gymnosperm

order Gnetales is reported for the first time in Australia, from specimens of Gnetum gnemon collected

on two continental islands in Torres Strait, Queensland. The distribution, habitat, ecology and local

conservation status of the species are discussed, and its origins in Torres Strait are considered with

reference to biogeographic and anthropogenic factors.

Key Words: Gnetaceae, Gnetum gnemon , Australia flora, Queensland flora, Badu Island, Mua Island,

Torres Strait

‘David G. Fell, P.O. Box 337, Alstonville, NSW 2477, Australia. Email: dgfell@westnet.com.au

Introduction

Gnetum L. is the sole genus within the

family Gnetaceae and order Gnetales of the

gymnosperms. The genus consists of about 40

species with 10 species in South America, one

in tropical West Africa, and 20-25 species in

tropical and subtropical Asia (Markgraf 1929,

1977; Maheshwari & Vasil 1961; Verheij &

Sukendar, 1991; Carlquist 1994, 1996; Price

1996; Wonn & Renner 2006, Hou et al. 2015).

The origins and evolutionary relationships

of the gnetophytes are considered as central

toward understanding the origin of flowers

and seed plant evolution (Markgraf 1929;

Chamberlain 1935; Price 1996; Hansen et al.

1999; Winter et al. 1999; Manner & Eleivich

2006). Recent genetic studies now recognise

that gnetophytes are more closely related to

conifers than to angiosperms (Hansen et

al. 1999; Winter et al. 1999; Becker et al.

2000), and that the major lineages of Gnetum

diverged in the Late Cretaceous (Hou et al.

2015).

Accepted for publication 21 August 2015

Gnetum gnemon L. has a wide distribution

in Malesia and the Western Pacific and is

native to Indonesia, Papua New Guinea, the

Solomons, and Vanuatu (Verheij & Sukendar

1991; Manner & Eleivich 2006; Wonn &

Renner 2006; Bourke 2010). In Papua New

Guinea it occurs in the Morobe, Eastern

Highlands, Western Highlands, Central,

East Sepik, Sandaun (West Sepik), Manus

Island, New Ireland, Western, Bougainville,

New Britain, Madang, Milne Bay, and Gulf

Provinces (Conn & Damas 2006). It is

previously known in Australia only from

cultivated trees in the vicinity of Darwin

(Atlas of Living Australia, 2012; I. Cowie

pers. comm., November 2012).

Gnetum gnemon is a dioecious evergreen

tree 15-30 m in height in the canopy or

subcanopy with a trunk diameter at breast

height to 40 cm (Peekel 1984; Conn & Damas

2006; Manner & Eleivich 2006). The trunk is

smooth and cylindrical with prominent raised

swollen bands (100-300 mm apart) caused

by the growth and fall of the opposite and

symmetrical branches (Conn & Damas 2006).

Leaves are simple and opposite, elliptic,

lanceolate or oblong-oval, 15-25 cm long and

5-9 cm wide (Manner & Eleivich 2006). As

a gymnosperm, Gnetum does not have true

422

flowers; the cones or strobili are presented at

the tip of a slender stem or axis (Chamberlain

1935; Manner & Eleivich 2006). It has yellow

single-seeded fruits which turn purplered or

orangered at maturity (Maheshwari & Vasil

1961; Manner & Eleivich 2006), ripening

between September and December (Bourke

etal. 2004).

Across its Asian, Malesian and Western

Pacific range the genus Gnetum is known

from tropical rainforest up to 1,700 m with

rainfall of 750-5,000 mm/year (Manner &

Eleivich 2006). It occurs in primary and

secondary vegetation, with cultivated trees

commonly found in home orchards and

subsistence gardens.

Wild and cultivated Gnetum gnemon is an

important natural resource in many parts of

Asia, Malesia and the Pacific where it occurs.

Documented uses include: food (leaves

for wrapping food items, young cones and

leaves cooked with meat for flavoring; seeds

ground into a flour for fried flat cakes; flowers

(stroboli) eaten; young fruits eaten raw or

cooked; timber (poles for house construction,

tool handles, burned for fuel, and pulped for

papermaking); stem bark fibre (string bags,

ropes, bowstring on musical instruments,

construction of fishing lines and fishnets,

assembling arrowheads and arrowshafts);

agroforestry and rehabilitation (intercropping

for rambutan and breadfruit, trellis for

Dioscorea yams, improving soil fertility,

windbreaks, boundary markers) (Henderson

& Hancock 1989; Verheij & Sukendar 1991;

Ohtsuksa 1983; Peekel 1984; Salim etal. 2002;

Walter & Sam 2002; Manner & Eleivich 2006;

Quartermain & Tomi 2010). Extracts derived

from Gnetum are reported useful as health

supplements (Kato et al. 2009) and the nuts

form an important home industry throughout

Indonesia (Cadiz & Florido 2001).

In Papua New Guinea it is an important

wild and cultivated food plant in many

locations (Powell 1976; Bourke 2004) and

known in Tok pisin as ‘ Tulip ’ (two leaves)

(French 1986). The Gidra people of the

Oriomo River area of southwest Papua New

Guinea adjacent to Torres Strait, eat the fruits

and leaves (Ohtsuka 1983), and the leaves and

Austrobaileya 9 ( 3 ): 421-430 ( 2015 )

fruit are identified as a source of protein across

the nation (Cordon 1970). Cultivation of trees

is achieved by propagating seed and/or from

cuttings (French 1986). In the Kiunga area of

Western Province the bark is used to make the

fibre for string bags and other products and

it is also cultivated as a food plant (leaves)

in Daru (Western Province) (B. Waterhouse

pers. comm., April 2014).

We report on the occurrence of

Gnetum gnemon on two islands in Torres

Strait, Queensland, thus recognizing the

gymnosperm order Gnetales for the first time

from Australia. The distribution, habitat and

ecology of the species at these localities are

discussed together with an assessment of

its local conservation status. Its origins in

Torres Strait are considered with reference to

biogeographic and anthropogenic factors.

Materials and methods

A survey of the vegetation of the Torres Strait

Islands, Queensland, Australia, was carried

out in 2007 (Stanton et al. 2009). The survey’s

primary objective was to map vegetation

communities at a scale of 1:25,000 and

Regional Ecosystems at a scale of 1:50,000,

and was supplemented by floristic inventory

and collections of voucher specimens for

Australian herbaria. Additional surveys on

the Badu and Mua islands between 2009

and 2015 have been carried out as part of a

biodiversity management planning program

through the Land and Sea Management Unit

of the Torres Strait Regional Authority (3D

Environmental 2011a, 2011b; Gynther et al.

2014; Reis etal 2015).

Collections of Gnetum gnemon were

made at Badu Island in October 2007 {Fell

10206 & Stanton ) (Fig. 1) and on Mua Island

in April 2011 {Fell 10803 & Stanton ) (Fig.

2), with further confirmed observations on

Mua in March 2014 and on Badu in May

2015 (Fell pers. obs.). Voucher specimens are

lodged with the Queensland Herbarium (BRI)

with duplicates to the Australian Tropical

Herbarium (CNS).

Fell et all., Gnetum gnemon in Australia

423

Fig. 1 . Leaves of Gnetum gnemon on Badu Island (Fell 10206 & Stanton). Photo: D.G. Fell

Results and discussion

Gnetum gnemon in Torres Strait

Badu Island

Badu Island is situated approximately midway

between the tip of Cape York Peninsula and

mainland New Guinea, and belongs to the

Near Western Group of Torres Strait Islands.

It is a continental island of 10,467 ha, centred

at 10° 07’ S, 142° 09’ E and located around

92 km south of the Papua New Guinea (PNG)

coastline and 70 km NW of Cape York (Map

1). The island is the homeland of the Badulgal

people with a population of 784 people as

at the 2011 census (Australian Bureau of

Statistics 2012).

The coarse grained Badu granite

dominates the landscape, forming the island’s

rugged interior of low rocky hillocks with

massive granite boulders. Granite basement

rock is overlain on its margins by younger

unconsolidated deposits including alluvial

deposits and extensive dune fields of varying

age and geomorphic expression (Willmott

& Powell 1977; Garnett & Jackes 1983; 3D

Environmental 2011a).

A total of 49 vegetation communities

within 20 broad vegetation groups and 32

Regional Ecosystems are recognised across

the island, within which 592 plant species

have been recorded. The flora comprises

560 native species with 17 ferns, one cycad,

one conifer and 572 flowering plants (3D

Environmental 2011a).

Mua Island

Mua (also known as Moa) Island lies adjacent

and immediately to the east-south east of

Badu, being separated by a narrow (2.5 km

wide) channel. Situated approximately 55 km

424

north of the tip of Cape York, Queensland and

94 km south of the New Guinea mainland

(Map 1), it has a total area of 17,001 ha,

and is the second largest island in Torres

Strait. There are two Torres Strait Islander

communities on the island: at Kubin and St

Pauls, which had populations of 163 and 258

respectively as at the 2011 census (Australian

Bureau of Statistics 2012).

Mua is formed on continental igneous

basement rocks and is topographically diverse,

dominated by the high point of Banks Peak

Austrobaileya 9(3): 421-430 (2015)

in the north-east, rising to 376 m. A rugged

east and south facing coastline features rocky

coastal headlands, and an expansive coastal

plain forms a broad enclave behind the island’s

north-eastern coastline. Vegetation is diverse

with a total of 62 vegetation communities

occurring within 23 broad vegetation groups

and 44 regional ecosystems. The flora is the

richest in the Torres Strait region with 609

native species, comprising 19 ferns, one

cycad, two conifers and 654 flowering plants

(3D Environmental 2012a).

Fig. 2. Upland habitat of Gnetum gnemon on Banks Peak, Mua Island. Photo: D.J. Stanton.

Local Habitat & Ecology

In Torres Strait Gnetum occurs in poorly

drained lowland swamp forest (Badu),

lowland riparian rainforest (Mua), and on

steep granitic hillslopes in well-developed

evergreen vineforest (Mua) (Fig. 3).

The Badu occurrence is within a closed

(swamp) forest characterised by poorly

drained soils which may be seasonally

inundated. Typical canopy tree species are

Carallia brachiata (Lour.) Merr., Deplanchea

tetraphylla (R.Br.) F.Muell., Lophostemon

suaveolens (Sol. ex Gaertn.) Peter G.Wilson

425

Fell et all., Gnetum gnemon in Australia

Fig. 3. Multi stemmed trunk of Gnetum gnemon in

lowland riparian rainforest on Mua Island. Photo: D.G.

Fell.

& J.T.Waterh., Maranthes corymbosa Blume,

Melaleuca dealbata S.T. Blake and Syzygium

angophoroides (F.Muell.) B.Hyland with a

sharp ecotone to the surrounding woodland

vegetation.

On Mua, Gnetum occurs in upland and

lowland situations. Upland habitats are on

the steep granite hillslopes and crests of

Banks Peak, the highest topographic feature

in Torres Strait. Gnetum was recorded

between approximately 250-350 m altitude

in evergreen notophyll vineforest and in

wind sheared evergreen vinethicket (Fig.

2). The evergreen vineforests are developed

on upper sheltered slopes where weathering

of the granite has produced a well-drained

and relatively fertile sandy loam soil (3D

Environmental 2011b). The rainforest type

is a unique and newly described ecosystem

for Queensland, restricted to only a few

mountain top locations in Torres Strait (3D

Environmental 2009). The canopy height

ranges from 23 m to 35 m with dominants

including Acmenosperma claviflorum (Roxb.)

Kausel., Anthocarapa nitidula (Benth.)

T.D.Penn. exMabb., Calophyllum sil Lauterb.,

Manilkara kanosiensis H.J.Lan & B.Meeuse,

Sterculia shillinglawii F.Muell. subsp.

shillinglawii and Syzygium beuttnerianum

(K.Schum.) Nied. Gnetum habitat on the

steep southern slopes of the peak features

a wind sheared canopy of Acmenosperma

claviflorum, Calophyllum sil, Podocarpus

grayae De Laub. Licuala ramsayi var. tuckeri

Barfod & Dowe and Pandanus zea H.St.

John. Riparian rainforest on alluvial sands

and silts constitute Gnetum lowland habitat

on Mua. Characteristic canopy species are

Buchanania arborescens (Blume) Blume,

Horsfieldia australiana S.T.Blake, Maranthes

corymbosa Blume, Syzygium angophoroides

and S.bamagense B.Hyland.

No uses are documented or known

for Gnetum gnemon in the Torres Strait.

Specimens were shown to Land and Sea

Rangers during our visits to Badu in

November 2007, November 2010 and May

2015, and on Mua in March 2011 and March

2014. They did not recognise the plant, and

stated that they had no name or use for it (D.

Williams pers. comm., 2015; T. Stow pers.

comm., 2007; J. Wigness pers. comm., 2011).

Natural or translocated?

Some plants and their populations in

Torres Strait are a reflection of natural

biogeographic distributions, whereas others

are anthropogenic and may indicate the

influence of people as dispersal agents

(Denham 2008; Denham etal. 2009; McNiven

2008).

The accepted hypothesis that the

pantropical range of Gnetum reflects a

Gondwanan history was established by

Markgraf (1929). However, Wonn & Renner

(2006) consider that its present range follows

a more recent radiation in the Malesian

region, and therefore is not Gondwanan.

Using analysis of fossil-calibrated molecular-

clocks, these authors investigated Gnetum

lineages now found in Africa, South America

and Southeast Asia with results suggesting the

influence of ancient long-distance dispersal of

426

seeds across seawater. Further, its distribution

throughout Asia is thought to have occurred

during radiation through Malesia where

opportunities for overland seed dispersal

coincided with times of low sea levels (Wonn

& Renner 2006).

Torres Strait was formed by rising sea

levels in the Holocene ( c . 8000-6500 years

before present) that inundated the Arafura

Plain, a low-lying land bridge that connected

Australia and New Guinea for much of

recent geological history (Jennings 1972;

Woodroffe et ah 2000). Given the occurrence

of G. gnemon in neighboring areas of central-

southern New Guinea, it is likely the Torres

Strait records are refugial populations that

reflect its biogeographic distribution across

the Malesian region.

However, the potential for the taxon to

have been introduced into Torres Strait by

human vectors should not be discounted. The

species is widely cultivated across its range,

including in New Guinea, and hence it is

possible that it was translocated to the islands.

Movements of human populations and trade

between New Guinea and Torres Strait have

greatly influenced the distribution patterns

of a number of important tropical crop plants

and has influenced cultivation practices

in the region (Haddon 1935; Barrau 1963;

Harris 1977; Barham 2000; Denham 2008;

Denham et ah 2009; McNiven et al. 2006;

McNiven 2008). Furthermore, the impact of

Torres Strait islanders upon their pre-contact

surroundings included the introduction of

new plant species from other islands (Shnukal

2004; McNiven 2008), a practice that is

ongoing in contemporary culture. The fact

that local people do not have a local name or a

use for the plant does not mean that it was not

used in the past.

Conservation Status

Across the majority of its range Gnetum

gnemon is a common tree species in the wild

that is also widely cultivated throughout

Malesia. In Torres Strait, a conservative

estimate of its population size on Badu is 500-

1000 individuals within an area of available

habitat of < 200 ha. Surveys carried out by

Austrobaileya 9(3): 421-430 (2015)

the authors in 2007,2009,2011,2014 and 2015

assessed the condition of the habitat as good,

being free of weeds although diggings by feral

pigs ( Sus scrofa) were observed throughout

the habitat. The even age of the canopy

and the species composition together with

evidence of old charred stumps indicate that

the habitat is transitional and subject to past

burning. Invasive weeds such as Singapore

daisy ( Sphagneticola trilobata (L.) Pruski),

praxelis ( Praxelis clematidea R.M.King &

H.Rob.), annual mission grass ( Pennisetum

pedicellatum Trin. subsp. pedicellatum )

and leucaena ( Leucaena leucocephala

(Lam.) de Wit) occur on the islands, but are

mostly associated with habitation and along

roadsides. Singapore daisy has successfully

invaded degraded swamp forests within and

on the margins of the Badu community and

is considered a serious threat to these habitats

across the island.

Resilience to fire of Gnetum gnemon is

not documented; however, its mesic habitat is

likely to be sensitive to fire. Fire has been an

ongoing influence on the island’s vegetation,

and fire management is an increasing focus

of the Land and Sea Ranger program (3D

Environmental 2012a, 2012b). Given the

small population size and highly localised

occurrence, there is a risk of stochastic

extinction of the species on this island.

The upland rainforest habitat of the

Mua population is similarly remote and

inaccessible, and no known direct threats

are evident. Population size is not known;

however, the spatial extent of the forest

type is mapped as < 477 ha (Stanton et ah

2009). While numerous weeds occur on the

island they are limited to the margins of the

island communities Kubin and St Pauls and

along some major tracks and roads. The

undisturbed and remote nature of the upland

habitat suggests that there would appear to

be a very low risk of stochastic extinction

of the population. Its lowland occurrence on

Mua occupies approximately 20 ha within

a linear band of riparian rainforest located

three kilometres from the coast. The ecotone

to adjacent Corymbia and Melaleuca grassy

woodland vegetation is sharp and controlled

427

Fell et all., Gnetum gnemon in Australia

by seasonal burning practices. Weeds such as

Brazilian joyweed (Alternanthera brasiliana

(L.) Kuntze) and leucaena occur nearby. Weed

incursions, and inappropriate fire management

which erode fire sensitive riparian margins,

represent potentially threatening processes to

the integrity of the lowland Gnetum habitat.

Integration of further ecological and

genetic studies are necessary to better

understand the conservation and management

requirements of Gnetum gnemon in Torres

Strait. Future assessments on the conservation

values of Gnetum gnemon in Torres Strait

should acknowledge that conservation of

peripheral populations is dependent upon the

genetic divergence from other conspecific

populations, and may be beneficial to the

protection of the evolutionary process (Lesica

& Allendorf 1995).

It is feasible that additional populations

occur on Badu and Mua and further field

assessments are required to better understand

the species ecology in terms of distribution,

population size, phenology, recruitment, and

to investigate cultural values. Similar focus

is also required for other additions to the

Australia flora recently recorded in Torres

Strait namely Cycas papuana F.Muell. and

Manilkara kanosienisis H.J.Lan & B.Meeuse

(Fell in prep.) as well as other highly disjunct

species recorded in the region (Fell & Stanton

2011). Such studies are achievable with

the support and involvement of traditional

landowners and their representative bodies

and the Torres Strait Regional Authority Land

and Sea Management Unit.

Acknowledgements

The authors acknowledge all Elders past and

present, Traditional Owners and community

members of Badu and Mua islands. Access

to Badu was provided by the Mura Badulgal

(TSI) Corporation Registered Native Title

Body Corporation (RNTBC) and to Mua by the

Mualgal (Torres Strait Islander) Corporation

RNTBC. These bodies administer the native

title rights and interests over their respective

islands. The field surveys on Badu and Mua

were funded and managed by the Land and

Sea Management Unit of the Torres Strait

Regional Authority. Field assistance on Badu

was provided by Badu Island Ranger Keijie

Bowie as well as by Mua Ranger Jackson

Ware, Warraber Trainee Ranger Young Billy

and Mer Trainee Ranger Tauki Passi. On

Mua Island, Ranger Mentor Simon Conaty

supported field work and his assistance is most

appreciated. Thanks to David Gooding who

provided GIS support to detailed vegetation

mapping over Torres Strait islands. Botanical

identifications were provided by the staff of

the Queensland Herbarium. Comments on

an earlier version of this paper from Garrick

Hitchcock, Rob Neal, Paul Forster and

Barbara Waterhouse are greatly appreciated.

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Austrobaileya 9(3): 421^-30 (2015)

Map 1. Position of Badu and Mua islands in Torres Strait indicating position between Papua New Guinea and mainland

Australia.

Plectranthus laetus P.I.Forst. and P. ventosus

P.I.Forst. (Lamiaceae), new species from

Cape York Peninsula, Queensland

Paul I. Forster

Summary

Forster, P.I. (2015). Plectranthus laetus P.I.Forst. and P. ventosus P.I.Forst. (Lamiaceae), new

species from Cape York Peninsula, Queensland. Austrobaileya 9(3): 431-438. Two new species of

Plectranthus are described from Cape York Peninsula: P. laetus P.I.Forst. from Orchid Creek Station

and P. ventosus P.I.Forst. from the Melville Range at Cape Melville National Park. Both species occur

on granite substrates within or adjacent to rainforests.

Key Words: Lamiaceae, Plectranthus , Plectranthus laetus , Plectranthus ventosus , Australia flora,

Queensland flora. Cape Melville, Orchid Creek, new species, taxonomy, distribution maps.

P.I.Forster, Queensland Herbarium, Department of Science, Information Technology and Innovation,

Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email: Paul.

For ster @dsit i. qld. gov. au

Introduction

Ongoing botanical exploration of remote

areas of Cape York Peninsula continues to

reveal vascular plants that are new to science.

A further two species of Plectranthus

L.Her. are described here, both discovered

in previously unsurveyed localities on the

tops of mountains or ridges, and in habitats

akin to the ‘sky island’ concept discussed

previously Forster (2014). In each case,

access to the localities was by helicopter.

The description of these new species brings

to 15, the number of species of Plectranthus

now recognised for the Cape York Peninsula

bioregion (https: //data. qld. gov.au/dataset/

bioegeographic-subregions-queensland). The

species previously recognised are P. apreptus

S.T.Blake, P. apricus P.I.Forst., P. arenicola

P.I.Forst., P. batianoffii P.I.Forst., P. congestus

R.Br., P. dumicola P.I.Forst., P. excelsus

P.I.Forst., P. foetidus Benth., P. megadontus

P.I.Forst., P. mirus S.T.Blake, P. pulchellus

P.I.Forst., P. scutellarioides (L.) R.Br. and P.

venustus P.I.Forst.; ten of the overall total are

endemic.

Accepted for publication 23 July 2015

Materials and methods

Fieldwork was undertaken in north

Queensland to procure fresh material

for specimens and cultivation enabling

observation of variation in morphology and

phenology. The subsequent descriptions

and observations are based on these recent

collections and other earlier collections in the

Queensland Herbarium (BRI).

Taxonomy

Plectranthus laetus P.I.Forst., sp. nov. with

affinity to P. pulchellus but differing in the

orange sessile glands (versus yellow), the

verticillasters with fewer flowers (6-10 versus

12-16), the flowers with corollas that are

strikingly blue-purple (versus light purple)

and the larger calyces. Typus: Queensland.

Cook District: Orchid Creek Station,

Coffee Scrub; SW of Lockhart River, Cape

York Peninsula, 30 April 2014, P.I. Forster

PIF41138 & S.L. Thompson (holo: BRI; iso:

CNS, MEL, NSW distribuendi ).

Perennial herb, stems erect to 50 cm high;

foliage scentless when crushed, not clammy;

non-glandular and glandular trichomes

uncoloured, non-glandular trichomes without

prominent raised bases, sessile glands

8-celled, orange. Roots fibrous, somewhat

432

fleshy and thickened. Stems square, erect

to straggling, fleshy, easily snapped, the

lower parts up to 13 mm diameter and

not noticeably thickened, pink-purple to

pink-green, upper parts with persistent

indumentum, non-glandular trichomes sparse

to dense, antrorse, 6-10-celled up to 1 mm

long, glandular trichomes absent, sessile

glands sparse. Leaves discolorous, petiolate;

petioles 7-19 x 1.5-2 mm, channelled on top,

non-glandular trichomes sparse, antrorse,

4-10-celled up to 0.5 mm long, glandular

trichomes absent, sessile glands occasional

to sparse; laminae lanceolate-ovate to ovate,

fleshy, ±flat to slightly keeled, 30-70 x 26-

50 mm, crenate with 8-14 teeth up to 4 mm

long on each margin, of similar length along

margin, secondary teeth poorly developed;

tip acute; base obtuse to rounded; upper

surface medium-green and somewhat glossy,

veins impressed and often pink-purple,

non-glandular trichomes sparse, antrorse,

4-8-celled up to 0.5 mm long, glandular

trichomes and sessile glands absent; lower

surface pale purple to purple-green and with a

light hoary silver frosting from indumentum,

veins strongly raised, non-glandular

trichomes sparse, antrorse, 4-8-celled up to

0.8 mm long, glandular trichomes absent,

sessile glands scattered. Inflorescence up to

350 mm long, usually single or with 1 or 2

side branches; axis square in cross-section,

pink-purple, non-glandular trichomes sparse,

antrorse, 4-8-celled up to 0.5 mm long,

glandular trichomes sparse and minute (< 0.2

mm long), sessile glands absent or occasional;

bracts broadly ovate, 1.5-1.8 x 7-8 mm,

ecomose, margins somewhat irregularly

crenate to crenulate due to slightly enlarged

bases of trichomes, non-glandular trichomes

sparse, antrorse, 4-6-celled up to 0.4 mm

long, glandular trichomes absent, sessile

glands occasional to scattered; verticillasters

6-10-flowered, 7-20 mm apart; pedicels

4-5 x c. 0.2 mm, non-glandular trichomes

sparse, antrorse, 2-4-celled up to 0.2 mm

long, glandular trichomes and sessile glands

absent. Flower calyces 2.5-2.8 mm long,

non-glandular trichomes sparse, antrorse,

2-4-celled up to 0.2 mm long, glandular

trichomes occasional and very short (< 0.2

Austrobaileya 9(3): 431-438 (2015)

mm long), sessile glands sparse. Corolla 9.5-

11.5 mm long, blue-purple; tube 5-6 mm long,

abruptly curved at 80-90° 2-2.4 mm from the

base, slightly inflated upwards, non-glandular

trichomes absent or occasional, 2-celled

and very short (< 0.2 mm long), glandular

trichomes and sessile glands absent; upper

lobes suborbicular, erect to weakly reflexed,

1.8-2 x 1.5-1.6 mm, non-glandular trichomes

scattered, antrorse, 2-4-celled up to 0.3 mm

long, glandular trichomes absent, sessile

glands sparse; lateral lobes rounded, c. 0.5

x 0.7-0.8 mm, glabrous; lower lobe oblong-

ovate, 5-5.2 x 4-4.2 mm, non-glandular

trichomes sparse, antrorse, 2-4-celled up to

0.3 mm long, glandular trichomes absent,

sessile glands sparse; filaments filiform, 7-8

x c. 0.2 mm, lilac, fused for 4-5 mm from the