Queensland Herbarium

// v \

^Queenslan d Government

Environmental Protection Agency

Volume 6

Number 3 2003

A Journal of Plant Systematics

Queensland Herbarium

181

Queensland Government

Environmental Protection Agency

Editorial Committee

L.W. Jessup (editor)

R.J.F. Henderson (technical advisor)

B.K. Simon (technical advisor)

Desktop Publishing

A.E. Sinclair

S. Pupavac

Yvonne Smith

Austrobaileya

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

Vol. 6, No. 2 was published on 30 September 2002

Vol. 6, No. 3 was published on 3 December 2003

Austrobaileya is published once per year.

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ISSN0155-4131

© Queensland Herbarium 2003

Austrobaileya is the journal of the Queensland Herbarium and is devoted to publication of

results of sound research and of informed discussion on plant systematics, with special empha¬

sis on Queensland plants.

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

views of the Queensland Herbarium.

Contents

A taxonomic revision of Croton F. (Euphorbiaceae) in Australia

Paul I. Forster.349

Phebalium distans P.I.Forst. (Rutaceae), a new and endangered species from

south-eastern Queensland, and reinstatement of P. longifolium S.T.Blake

Paul I. Forster.437

A synopsis of Racosperma C. Mart. (Feguminosae: Mimosoideae)

Fes Pedley.445

Studies in Euphorbiaceae A.F.Juss. sens. lat. 5

A revision of Pseudanthus Sieber ex Spreng. and Stachystemon Planch.

(Oldfieldioideae Kohler & Webster, Caletieae Mull. Arg.)

David A. Halford and Rodney J.F. Henderson.497

Backhousia oligantha (Myrtaceae), a new species from Queensland

A.R. Bean.533

Six new species of Hydrocotyle F. (Apiaceae) from Queensland

A.R. Bean & M. J. Henwood.537

Anew species of Mimulus F. (Scrophulariaceae) from Queensland, Australia

A.R. Bean.549

Polycarpelly in Idiospermum australiense (Calycanthaceae)

Stuart J. Worboys.553

Notes

Jasminum longipetalum King & Gamble (Oleaceae), and its occurrence in

Queensland, Australia

Paul I. Forster.557

Lepisanthes senegalensis (Juss. ex Poir.) Feenh. (Sapindaceae), anew

generic and specific record for Queensland

Paul I. Forster.559

Steinchisma laxa (Sw.) Zuloaga, the correct name for Clijfordiochloa

parvispiculata B.K.Simon

Bryan K. Simon.561

The puzzle of Eucalyptus hemilampra F.Muell. (Myrtaceae)

A.R. Bean.563

New combinations in Lycianthes (Dunal) Hassl. (Solanaceae) for

New Guinea and Australia

A.R. Bean.567

Megathyrsus , a new generic name for Panicum subgenus Megathyrsus

Bryan K. Simon & Surrey W.F. Jacobs.571

Obituary, John W. Parham, 1929-2002 .575

A taxonomic revision of Croton L. (Euphorbiaceae) in Australia

Paul I. Forster

Summary

Forster, Paul I. (2003). A taxonomic revision of Croton L. (Euphorbiaceae) in Australia.

Austrobaileya 6(3): 349-436. The genus Croton L. is revised for Australia. Twenty-seven native

species (all shrubs, trees or lianes) are recognised: C. acronychioides F.Muell., C. arnhemicus

Muell.Arg., C. aridus P.I.Forst. sp. nov., C. brachypus Airy Shaw, C. byrnesii Airy Shaw,

C. capitis-york Airy Shaw, C. caudatus Geisel., C. choristadenius K.Schum., C. densivestitus

C.T.White & W.D.Francis, C. dockrillii Airy Shaw, C. habrophyllus Airy Shaw, C. insularis Baill.,

C. magneticus Airy Shaw, C. mamillatus P.I.Forst. sp. nov., C. minimus P.I.Forst. sp. nov.,

C. multicaulis P.LForst. sp. nov., C. multicaulis subsp. velutinus P.I.Forst. subsp. nov., C. mutabilis

P.I.Forst. sp. nov., C. phebalioides Muell.Arg., C. rarus P.I.Forst. sp. nov., C. schultzii Benth.,

C. simulans P.I.Forst. sp. nov., C. stigmatosus F.Muell., C. stocked (Airy Shaw) Airy Shaw,

C. tomentellus Airy Shaw, C. triacros F.Muell., C. verreauxii Muell.Arg. and C. waterhouseae

P.I.Forst. sp. nov. All apart from C. caudatus, C. choristadenius and C. insularis are endemic.

Three naturalised species are recorded: C. capitatus Hook., C. glandulosus F. and C. setigerus

Hook., all being small herbaceous weeds. One species (C. armstrongii S.Moore) is of dubious origin

with the type from Australia but no subsequent collections. An identification key is provided to all

thirty-one species. All taxa are described and all native species and subspecies illustrated. Notes are

provided on distribution, habitat, typification, affinities and conservation status for each taxon.

Fectotypes are selected for the names C. acronychioides F.Muell., C. affinis Maiden & R.T.Baker,

C. arnhemicus Muell.Arg., C. stigmatosus F.Muell. and C. triacros F.Muell. The new combination

Adriana urticoides (A.Cunn.) Guymer is made for Croton urticoides A.Cunn.

Keywords: Croton - Australia; Croton aridus, Croton caudatus, Croton choristadenius, Croton

mamillatus, Croton minimus, Croton multicaulis, Croton multicaulis subsp. velutinus, Croton

mutabilis, Croton rarus, Croton schultzii, Croton simulans, Croton waterhouseae, Adriana urticoides

Paul I. Forster, Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic

Gardens Mt Coot-tha, Mt Coot-tha Road, Toowong, Queensland 4066, Australia

Introduction

The genus Croton L. was described by

Linnaeus (1753) and thirteen species were

named at that time. Since then many species

have been included in Croton, and although

some have since been transferred to other

genera, it is estimated that there are between

800 (Webster 1993) and 1200 (Berry 1999)

species in the genus. Croton is second only to

Euphorbia L. in number of species within the

family.

Croton is included in Euphorbiaceae

subfamily Crotonoideae, tribe Crotoneae with

the Old World genera Mildbraedia Pax,

Moacroton Croizat and Paracroton Miq.

(sometimes listed as the invalid Fahrenheitia

Reichb.f. & Zoll.) (Webster 1994). Occasionally

the genera Crotonopsis Michx., Eremocarpus

Benth. and Julocroton Mart, are also

recognised in this tribe (Radcliffe-Smith 2001),

although all three genera were reduced to

sections of Croton by Webster (1992). Croton

Accepted for publication 11 February 2003

is distinguished from the other genera in the

Crotoneae mainly by the filaments inflexed in

the bud and the pistillate petals being reduced

or absent (Webster 1994). The Crotoneae is

probably derived within the Crotonoideae

(Tokuoka & Tobe 1998); however, a

comprehensive phylogeny for the group is yet

to be proposed.

Species of Croton are found throughout

the tropics and subtropics in both the Old and

New Worlds, or as Hooker (1890) stated “in all

hot countries”. There are major concentrations

of species in the Neotropics (J.Mueller 1873;

Webster 1992; Berry 1999), Mexico (Webster

2001), Madagascar (Govaerts et al. 2000) and

parts of Malesia (Airy Shaw 1980a), but lesser

numbers in Africa (Radcliffe-Smith 1996,1997),

continental Asia (Hooker 1890; Chakrabarty &

Balakrishnan 1997; Philcox 1997) and Australia

(this paper). The last overall monograph of

Croton was by J.Mueller (1866), and the sheer

number of species makes the task of a modern

monograph daunting.

350

Australia is relatively “depauperate” with

twenty-seven native species, three naturalised

species and one species of doubtful origin. One

species, Croton armstrongii S.Moore, is

tentatively included, as only the type (there are

no subsequent collections) is reputedly of

Australian origin, hence it is excluded from most

discussion below. The first recording of species

for Australia that were referred to the genus

“ Croton ” was by Labillardiere (1806) who

described C. quadripartitus from Tasmania and

C. viscosus from Western Australia. These taxa

are now included in Adriana Gaudich. and

Beyeria Miq. respectively. The first Australian

species currently included in Croton was

C. verreauxii described by Baillon (1858), soon

followed by additional species in the 1860’s

(F.Mueller 1864, 1868; J.Mueller 1864, 1865,

1866). Bentham (1873) included nine species in

Croton, and some additional species and

infraspecific taxa were recognised before the

revision and conspectus by Airy Shaw (1976,

1980b,c, 1981).

Airy Shaw (1981) recognised nineteen

species of Croton for Australia, provided a

species key, bibliographic details, and notes on

distribution and habit. He did not resolve the

typification of many species, provide detailed

comparative descriptions, or adequately deal

with variation in some taxa. In the present

account several new taxa are described. Five of

these ( Croton mamillatus, C. minimus,

C. rarus, C. simulans and C. waterhouseae ) are

narrow endemics and have been discovered

subsequent to Airy Shaw’s work or were not

seen by him, whereas C. aridus, C. mutabilis

and C. multicaulis were included by Airy Shaw

within other species. Croton caudatus and

C. choristadenius are newly recorded for

Australia and occur also in Malesia. The

presence of three species in Australia, notably

Croton argyratus, C. cocchymelophyllus and

C. storckii is refuted. The three naturalised

herbaceous species Croton capitatus,

C. glandulosus and C. setigerus are also

included in this account.

The Australian species of Croton are

largely tropical and subtropical in their

distribution. The majority of species (twenty-

two) occur in rainforest communities (sensu

Webb & Tracey 1981, ranging from evergreen

Austrobaileya 6 (3): 349-436

notophyll vineforests to deciduous

vinethickets), although several taxa grow in

woodland communities and one occurs in the

arid zone on red sand-hills. Of the twenty-seven

native species, all but three are endemic. The

non-endemic native species occur elsewhere in

Malesia or Melanesia, with two of them ( Croton

caudatus and C. choristadenius ) known from

single localities in far north Queensland. Several

species are very widespread (e.g. Croton

arnhemicus, C. insularis and C. phebalioides)

and occur over 44-55 1° grid squares (Map 1).

Six endemic species ( Croton brachypus,

C. byrnesii, C. mamillatus, C. simulans,

C. stockeri and C. waterhouseae are very

restricted in occurrence with distributions in

only one or two 1° grid squares. The remaining

species fall between these two extremes, with

some such as Croton acronychioides,

C. habrophyllus and C. verreauxii being also

widely distributed (15-24 1° grid squares).

In Australia the Mcllwraith Range (grid

square 13°S, 142°E) has ten species present

(Map 1). Lesser centres of diversity (six or seven

species present) occur at Iron Range (grid

square 12°S, 142°E), the southern part of the

‘Wet Tropics’ (grid square 17°S, 145°E) and

south of Brisbane in, or adjacent to the

McPherson Range (grid square 28°S, 152°E)

(Map 1). These higher species densities are a

reflection of diverse habitats (due to rainfall and

altitude gradients, and diverse geology) being

present in these grid squares, and this pattern

is repeated in many other plant groups in eastern

Australia.

At least one of the narrow endemics

( Croton mamillatus ) can be considered as

Critically Endangered using the criteria of the

IUCN (2001). This category would also apply

to the Australian populations of Croton

caudatus and C. choristadenius. Apart from

several species that are listed as Vulnerable

{Croton magneticus ) or Rare (C. brachypus, C.

densivestitus and C. stockeri), under

Queensland Government legislation, the

majority of species are not considered

threatened.

Plant habit of Australian Crotons includes

small, wiry herbs, (the three naturalised species)

shmbs, lianes and small trees. Most of the native

Australian species are shrubs, four are trees

Forster, Croton in Australia

351

115 120 125 130 135 140 145 150 155

Map 1 . Distribution of Croton (native taxa) in Australia indicating the number of species in

each 1° degree grid square.

and one a canopy liane. Some species may be

common components in the habitats where they

occur, forming dense thickets. A useful field

indicator for species of Croton (at least in

Australia, but also in South Africa, New Guinea

and Thailand where I have encountered species)

is the colour of the fallen leaves, which are

orange. Some of the Australian species are

seasonally deciduous (e.g. Croton mutabilis,

C. rarus, C. simulans ) with the mature foliage

often being quite dissimilar to the young leaves

that are present at flowering. This process of

shedding of the old foliage just prior to

flowering, followed by a flush of new foliage at

the same time as the flowers, seems to be

widespread in some groups or species of

Euphorbiaceae (e.g. Drypetes deplanchei

(Brongn. & Gris) Merr. (Forster \991),Mallotus

surculosus P.I.Forst. (Forster 1999), Claoxylon

spp. (Forster unpubl.). Conversely, many

Australian Crotons will hold inflorescences in

an arrested state of development for months

(e.g. Croton insularis, C. magneticus,

C. phebalioides ) until sufficient moisture is

available for flower production.

All of the Australian species of Croton

appear to be monoecious with the flowers in

glomerules of one to many flowers. True dioecy

is however, relatively widespread in non-

Australian taxa (e.g. Decker & Pilson 2000). The

Australian species usually have inflorescences

with both male and female flowers, the females

usually being few and single in the glomerules

towards the base, and the males being many

and in groups of 1 to many in the glomerules

towards the apex. There are generally many more

male flowers than female flowers in any

inflorescence. It is also not unusual to observe

inflorescences where the flowers are all of one

sex. In these instances the flowers are usually

all male and are being produced during drought.

Occasionally both male and female flowers may

be present in the same glomerule.

As yet we have little information on the

reproductive biology of Australian Crotons.

Casual observations of the flowers would tend

to indicate that the female flowers towards the

base of the inflorescence open first, followed

by the males towards the top. In non-Australian

352

species the ratio between male and female may

be related to the age of the plant (Shaanker &

Ganeshaiah 1984). This pattern of non-

synchronous floral development (or temporal

dioecy) seems to be widespread in monoecious

Euphorbiaceae and would tend to favour

outcrossing (Bawa et al. 1982; Freitas et al.

2001). In many instances, however, there are

both male and female flowers open at the same

time in the one inflorescence, and certainly on

the one individual. Croton flowers seem to be

most suitable for bee pollination (Endress 1994)

and I have seen hordes of native bees (viz.

Trigona spp.) working the inflorescences of

flowering individuals. Other insects such as

various Coleoptera, Hymenoptera and

Lepidoptera have also been recorded as

pollinators of the Argentinian Croton

sarcopetalus (Freitas et al. 2001), and it is not

unreasonable to predict that such broad guilds

of insects would also visit the flowers of

Australian Crotons. The major reward for such

attention is probably both pollen and nectar,

and the small male flowers of Crotons perfectly

fit what Endress (1994) termed as ‘bowl’ flowers

that are mainly visited by diverse insects with

short proboscises. Small ants are also common

visitors to the flowers, and although they

probably mainly act as pollen and nectar

robbers, it is possible that they are more than

incidental in pollination efficiency. Interestingly

enough, for the herbaceous Croton suberosus

H.B.K. from Mexico, it has been found that floral

nectar seems mainly to attract ants that act as

herbivore predators, rather than as pollinators

(Dominguez etal. 1989).

Nearly all of the Australian Crotons have

noticeable extrafloral nectaries at the base of

the leaf lamina or distant end of the petiole and

their role as sources of attraction to ants or other

insects is unknown, although they do secrete

small amounts of nectar (Jose & Inamdar 1989;

Freitas etal. 2000,2001). Certainly the position

and gross morphological form of these organs

have proved useful as taxonomic characters in

the current account.

As with many Euphorbiaceae, in Croton

the fruit are capsular and dehiscent. The

presence of a fleshy caruncle on the seeds

undoubtedly make them attractive to ants that

may subsequently aid in dispersal. In the

Austrobaileya 6 (3): 349-436

Indian Croton bonplandianus Baill. the female

flowers bear nectar glands that only become

active during fruit maturation and attract ants

that may subsequently disperse the seeds

(Ganeshaiah & Shaanker 1988). Whether this

phenomenon is widespread in the genus is

unknown. Likewise the presence and form of

these nectaries in most species of Croton is

unknown, but it is likely that further useful

taxonomic characters could be found from their

study.

Groupings of taxa

Webster (1993a) has proposed a new

classification of Croton with forty sections. In

the present account I have not followed his

classification that is based mainly on New World

taxa. Instead the Australian species are listed

alphabetically. Some Australian species (e.g.

Croton stocked, C. arnhemicus ) appear to have

combinations of characters that transgress

Webster’s sections, many of which appear to

be artificial and do not correlate with some of

the New World taxa included. Nevertheless a

number of groups in native Australian Croton

can be inferred on the basis of morphology.

Group 1.

Lianes. Included species: C. caudatus.

Group 2.

Shrubs or trees. Foliage penninerved, not silver-

white below, indumentum generally of scattered

trichomes. Included species: Croton

acronychioides, C. brachypus, C. byrnesii,

C. choristadenius, C. dockrillii, C. habrophyllus,

C. mutabilis, C. rarus, C. triacros, C. verreauxii.

Group 3.

Shrubs or trees. Foliage palminerved, not silver-

white below, indumentum of dense trichomes

(pubescent to velutinous). Included species:

Croton aridus, C. armstrongii, C. arnhemicus,

C. minimus, C. multicaulis, C. waterhouseae.

Group 4.

Shrubs. Foliage penninerved, not silver-white

below, indumentum of dense trichomes

(pubescent to velutinous). Included species:

Croton densivestitus, C. magneticus,

C. stocked.

Forster, Croton in Australia

Group 5.

Shrubs or trees. Foliage penninerved, silver-white

below, indumentum of dense adpressed

trichomes. Included species: Croton capitis-york,

C. insularis, C. mamillatus, C. phebalioides,

C. simulans, C. stigmatosus.

Group 6 .

Shrubs or trees. Foliage palminerved, silver-

white below, indumentum of dense adpressed

trichomes. Included species: Croton schultzii,

C. tomentellus.

Materials and Methods

This revision is based on herbarium holdings

at AD, BRI, CANB (including CBG), DNA, MEL,

NSW, PERTH and QRS, selected type material

at BM and BO, photographs or microfiche of

types at BM, G-DC, K and P, and field collections

and observations by the author. In some

instances, where there is a paucity of Australian

collections for a particular taxon (e.g. Croton

capitatus, C. caudatus, C. choristadenius,

C. glandulosus and C. setigerus ), selected

non-Australian collections have also been cited

and used in formulating the descriptions.

Species are defined as groups of

populations (1-many) with discontinuities in two

or more independent character states of

morphology. Where a single character state

difference is present and the discontinuity is

geographically based, the rank of subspecies

is used. This is a species definition that is

widely used and understood (Stebbins 1950;

Cronquist 1988) but would equate to the

‘diagnostic species concept’ of Judd et al.

(2002). Characters most commonly used in the

identification key are those of habit,

indumentum type (Fig. 1), leaf lamina venation

and marginal teeth number, extrafloral nectaries

(position and form), stem stipules, flowers

(particularly stamen number) and fruit shape.

Collectors should ensure that at least both male

and female flowers are collected when making

specimens of Crotons.

Invariably my species concept is closely

tied to habitat preferences and geographic

distribution and has been largely arrived at from

extensive fieldwork in northern Australia,

particularly Queensland where twenty-four

species are found. I have been fortunate to

353

study twenty-four of the twenty-seven native

species in the field.

Floral descriptions were prepared from

material preserved in spirit (FAA or 70% alcohol

and glycerol) or reconstituted by boiling in water

and detergent. Fruit descriptions were prepared

from spirit and dried material. Foliage,

inflorescence and seed descriptions were

prepared from dried material.

Indumentum cover is described using the

terminology of Hewson (1988), except that

‘scattered’ is used instead of ‘isolated’.

Indumentum in Australian Croton species

comprises multicellular, simple trichomes or

compound trichomes (Fig. 1). The compound

trichomes include sessile stellate trichomes,

stalked stellate trichomes, peltate trichomes and

peltate scales (lepidote). Peltate scales and

stellate or peltate trichomes are usually sessile

rather than stalked. Where no indication is

given in the descriptions, it may be assumed

that they are sessile. A system for trichome

morphology in Croton has been proposed by

Webster et al. (1996), but this has not been

followed here, mainly because the trichome

types in Australian crotons are not as diverse

as the 120 species that they studied. The fruit

of Croton species commonly have sessile and/

or stalked trichomes, or rarely on a fleshy

mamillate protuberance (C. capitis-york,

C. mamillatus, C. stigmatosus ), hence it is

always specified as to the condition present.

The term ‘foliar glands’ is included in the

descriptions and this refers to the small glands

that are present at the apices of any teeth on

the leaf lamina margins. If these foliar glands

are stated to be ‘prominent’ this means that they

are discernible with the naked eye.

‘Inconspicuous’ means that they are only

discernible with a microscope. Most of the

native Australian Crotons have extrafloral

nectaries (Jose & Inamdar 1989; Freitas et al.

2000, 2001) at, or near to the base of the leaf

lamina; these may be referred to as ‘glands’ in

other works or keys.

Common abbreviations in the specimen

citations are N.P. or N.P.R. (National Park), L. A.

(Logging Area), S.F. or S.F.R. (State Forest or

State Forest Reserve) and T.R. (Timber Reserve).

Rainforest terminology follows Webb (1978).

354

Austrobaileya 6 (3): 349-436

Fig. 1. Trichome types in Australian Croton. A. simple multicellular trichome. B. sessile stellate trichome.

C. stalked stellate trichome. D. peltate trichome. E. peltate scale. Del. W. Smith.

The ‘Wet Tropics’ is defined as that area of

north-eastern Queensland which encompasses

the ‘hot, humid vine forests’ from near

Cooktown in the north to Paluma in the south

(Webb & Tracey 1981; Barlow & Hyland 1988).

Conservation codings follow those that are

listed in Queensland legislation, and are derived

from those proposed by the IUCN (Anonymous

2001 ).

The taxa are mapped in 1° grid squares.

This has enabled information to be gathered

about centres of species richness and the

restriction or otherwise of the individual taxa.

Taxonomy

Croton L., Sp. PI. 1004 (1753). Type: Croton

tiglium L. (lectotype chosen by Small

1913).

Derivation of name: From the Greek kroton

(a tick), apparently a fanciful allusion to

the ‘tick-like’ seeds.

Generic synonymy is listed in Webster (1994)

and Radcliffe-Smith (2001); however, none of

these names have been applied to Australian

taxa so they are not repeated here. The following

generic description is meant to be

comprehensive for the taxa in Australia, but

will also be largely applicable for the genus

elsewhere.

Herbs, lianes, shrubs or small trees, annual

or perennial, monoecious or dioecious,

evergreen or deciduous; stems and foliage

without latex. Indumentum of simple or

compound trichomes and scales in various

combinations, glandular trichomes absent,

Forster, Croton in Australia

stinging trichomes absent. Stipules entire or

lobed, generally inconspicuous, deciduous.

Leaves alternate to subopposite, sessile to

petiolate, simple and usually elobate, palmi- or

penninerved, entire or denticulate to crenate,

often with sessile or stipitate glands at lamina

base or on petiole. Inflorescences terminal or

axillary, racemose or spicate, solitary, uni- or

bisexual and androgynous, with flowers in

bracteate glomerules. Male flowers sessile to

pedicellate; calyx lobes 4-6, imbricate or valvate,

± equal; petals 4-6, free, usually shorter than

sepals; stamens 5-50 (-100 plus), filaments free

and attached to a slightly raised pilose

receptacle, inflexed in bud, filiform or flattened;

anthers dorsifixed, bilobate, thecae oblong and

longitudinally dehiscent; pistillodes absent.

Female flowers sessile to pedicellate; calyx lobes

(0—) 4-6 (-8), imbricate or valvate; petals usually

absent; disk annular, or of separate glands or

absent; ovary l-3(4)-locular, locules

355

uniovulate; styles shortly connate at base, bifid

to multifid. Fruits capsular, uni-, bi- or trilobate,

surface smooth and variously pubescent,

dehiscing septicidally into bivalved cocci, or

rarely indehiscent. Seeds ovoid, obloid, ellipsoid

or subglobose; testa crustaceous to woody;

albumen fleshy; caruncles entire, non-arilloid;

cotyledons broad, flat.

Over 800 species in the tropics and

subtropics. Thirty-one species in Australia, with

three naturalised and twenty-seven native

species with twenty-two being endemic. One

species is of dubious origin.

Webster (1967) chose to retypify Croton

with C. aromaticus L. as the lectotype species,

but this does not supersede the

lectotypification of Small (1913) with C. tiglium

L. and the status quo is maintained by Radcliffe-

Smith (2001) and here.

Key to the Australian species of Croton

1. Herbs.2

Lianes, shrubs or trees.4

2. Styles 1; fruits unilobate. 24. C. setigerus

Styles 3; fruits trilobate.3

3. Extrafloral nectaries 2 at top of petiole . 13. C. glandulosus

Extrafloral nectaries absent from leaf.7. C. capitatus

4. Lianes. 9. C.caudatus

Shrubs or trees.5

5. Leaf lamina palminerved.6

Leaf lamina penninerved.13

6. Leaf lamina silver-white below .7

Leaf lamina green below (not silver-white).8

7. Stipules linear-lanceolate, 3-6 mm long; leaf lamina below with dense

overlapping peltate scales. 23. C. schultzii

Stipules lanceolate, 1.2-1.8 mm long; leaf lamina below with dense stellate

trichomes. 28. C. tomentellus

8. Leaf lamina denticulate to crenate with 60-100 teeth. 4. C. arnhemicus

Leaf lamina denticulate to crenate with 30-52 teeth.9

9. Stipules lanceolate, 0.3-1 mm long. 3. C. armstrongii

Stipules linear to linear-lanceolate, 1.2-6 mm long.10

356 Austrobaileya 6 (3): 349-436

10. Leaf lamina with interlateral tertiary venation obscure below. 18. C. minimus

Leaf lamina with interlateral tertiary venation well developed below.11

11. Leaf lamina with 32-40 teeth; male flowers with pedicels 10-12 mm

long; stamens 32-38 . 31. C. waterhouseae

Leaf lamina with 10-28 teeth; male flowers with pedicels 1.5-7 mm long;

stamens 10-24.12

12. Male flower petals 4-4.5 mm long, 0.8-1 mm wide; fruit oblong-ovoid,

3-17 mm long; seed 8.5-10.5 mm long. 2. C. aridus

Male flower petals 1.5-3.2 mm long, 0.5-1.5 mm wide; fruit

depressed-globose, 5-8 mm long; seed 4-5 mm long. 19. C. multicaulis

13. Foliage silver-white to silver-green below due to presence of dense

adpressed silver scales and trichomes. 14

Foliage green to ferruginous below due to the absence of indumentum or

the presence of uncoloured or ferruginous indumentum.19

14. Leaf lamina with obscure lateral venation.15. C. insularis

Leaf lamina with discernible lateral venation.15

15. Branchlets with peltate scales only. 8. C. capitis-york

Branchlets with stellate trichomes, or with admixture of stellate trichomes

and peltate scales. 16

16. Branchlets with stellate trichomes only.17

Branchlets with either peltate trichomes, or an admixture of peltate scales

and stellate trichomes. 18

17. Leaf lamina with the lateral veins not prominently raised below; extrafloral

nectaries sessile, visible above only . 21. C. phebalioides

Leaf lamina with the lateral veins prominently raised below; extrafloral

nectaries stipitate to 1.8 mm long, visible above & below. 26. C. stigmatosus

18. Branchlets and leaf petioles with peltate trichomes only; extrafloral

nectaries absent or circular. 17. C. mamillatus

Branchlets and leaf petioles with admixture of peltate scales and stellate

trichomes; extrafloral nectaries ellipsoid.25. C. simulans

19. Leaf lamina below with indumentum of peltate scales or peltate trichomes .20

Leaf lamina below with indumentum of stellate trichomes .23

20. Leaf petioles 2-5 mm long; lamina narrow-ovate, oblanceolate or obovate,

margins entire to sinuate. 5. C. brachypus

Leaf petioles 4-36 mm long; lamina elliptic, lanceolate or ovate, margins

denticulate to weakly crenate with 4-34 teeth per side of midrib.21

21. Extrafloral nectaries stipitate.30. C. verreauxii

Extrafloral nectaries sessile.22

22. Stipules 0.5-1 mm long; extrafloral nectaries 0.8-1.2 mm long; leaf lamina

below with scattered peltate trichomes. 10. C. choristadenius

Stipules 1-4 mm long; extrafloral nectaries 0.4-0.6 mm long; leaf lamina

below with scattered peltate scales. 1. C. acronychioides

Forster, Croton in Australia 357

23. Leaf lamina with dense velutinous indumentum below.24

Leaf lamina with scattered to sparse indumentum below.26

24. Leaf indumentum yellow; extrafloral nectaries stipitate.11. C. densivestitus

Leaf indumentum orange-brown or ferruginous-silver; extrafloral nectaries

sessile.25

25. Leaf indumentum orange-brown; stipules lanceolate, 3-6 mm long;

inflorescence bracts linear-lanceolate.27. C. stockeri

Leaf indumentum ferruginous-silver; stipules subulate, 0.3-0.9 mm long;

inflorescence bracts lanceolate to oblanceolate.16. C. magneticus

26. Leaf lamina with 70-112 teeth.14. C. habrophyllus

Leaf lamina with 18-64 teeth.27

27. Leaf indumentum uncoloured to silver.28

Leaf indumentum ferruginous to yellow.29

28. Leaf lamina with 40-56 teeth, extrafloral nectaries stipitate; fruits globose,

c. 4 mm long and 4 mm diameter.12. C. dockrillii

Leaf lamina with 18-34 teeth, extrafloral nectaries sessile; fruits

depressed-globose, 4-5 mm long, 6-7 mm diameter.20. C. mutabilis

29. Stipules lanceolate, 4-4.5 mm long; male flower sepals 1.6-2 mm long.29. C. triacros

Stipules linear to linear-lanceolate, 0.7-3.9 mm long; male flower sepals

2-2.5 mm long.30

30. Male flower pedicels 1-2 mm long, sepals obovate; styles bifid; fruits

globose, 4-5 mm long, 4^1.5 mm diameter. 22. C. rarus

Male flower pedicels 2.5-7 mm long, sepals ovate; styles multifid, divided

twice; fruits depressed-globose, 4.5-5 mm long, 6-7 mm diameter. 6. C. byrnesii

1. Croton acronychioides F.Muell., Fragm. 4:142

(1864) (‘acronychoides’). Type:

Queensland. Port Curtis District: [label 1

in unidentified hand] “A handsome shrub

14 or 15 feet high growing in the scrub nr

Rockhampton” [label 2 in F.Mueller hand]

“Rockhampton Bowman ” (lecto [here

chosen]: MEL231235).

Croton affinis Maiden & R.T.Baker, Proc.

Linn. Soc. New South Wales, II, 9: 160, t.

12 (1894). Type: New South Wales, near

Tintenbar, August 1893, W. Baeuerlen s.n.

(lecto [herechosen]: NSW273894).

Illustrations : Floyd (1989: 141); James &

Harden (1990:419); Hauser (1992:180).

Shrub to 5 m high, monoecious, evergreen,

perennial. Indumentum ferruginous-yellow.

Branchlets rounded, with scattered to sparse

peltate trichomes and scales when young,

glabrescent. Stipules lanceolate, 1-4 mm long,

0.3-0.8 mm wide, entire and with dense peltate

scales. Leaves alternate, petiolate, discolorous;

petioles 4-13 mm long, 0.5-1 mm wide, with

dense peltate scales when young, glabrescent;

lamina elliptic to ovate, 20-140 mm long, 10-60

mm wide, penninerved with 8-14 lateral veins

each side of midrib, tertiary reticulate veins

obscure; upper surface glossy green, lateral

veins weakly visible, with scattered peltate

scales when young, glabrescent; lower surface

matt green, lateral veins prominent, with

scattered and ± persistent peltate scales, neither

scabrous or velutinous; margins denticulate to

weakly crenate with 12-26 teeth up to 0.3 mm

long, foliar glands prominent; tip acute,

obcordate or retuse; base cuneate; extrafloral

nectaries 2 at base, sessile, circular to elliptic,

0.4—0.6 mm long, 0.3-0.4 mm wide, visible above

and below. Inflorescence up to 40 mm long,

358

unbranched, often unisexual but occasionally

bisexual and androgynous, pedunculate up to

10 mm; axis with sparse to dense peltate scales;

bracts lanceolate, 0.5-1 mm long, 0.2-0.3 mm

wide, with sparse to dense peltate trichomes.

Male flowers 2.8-3 mm long, 3.5-5 mm diameter,

held singly on inflorescence, spaced up to 2

mm apart; pedicels 2-3.7 mm long, 0.3-0.5 mm

wide, with scattered peltate trichomes at base

or glabrous; sepals valvate, 5, lanceolate-ovate,

1.8-2.3 mm long, 1.2-1.6 mm wide, glabrous or

lanate in upper half; petals 5, lanceolate to

lanceolate-ovate, 2-2.8 mm long, 0.9-1.3 mm

wide, lanate in upper half; stamens 6, filaments

± flattened, 1.8-2 mm long, 0.4-0.6 mm wide,

with dense simple trichomes at base, anthers

oblong, 0.7-1.1 mm long, 0.7-0.9 mm wide.

Female flowers 3-3.5 mm long, 3-5 mm diameter,

held singly and spaced up to 5 mm apart;

pedicels 1.5^1 mm long, c. 1 mm diameter, with

sparse peltate trichomes; sepals valvate, 5,

lanceolate, 2-3 mm long, 0.5-1.5 mm wide,

glabrous or with scattered marginal cilia; petals

absent; styles 3, linear to obloid, up to 2 mm

long, bifid once for up to 1.5 mm long, connate

at base for c. 0.4 mm, glabrous; ovary 3-locular,

2.3-2.5 mm long, 2.3-2.5 mm diameter, with

dense, sessile peltate scales. Fruits trilobate,

globose, 7-9 mm long, 6-8 mm diameter, with

sparse, sessile peltate scales. Seeds ± obloid,

6-6.8 mm long, 3.5—4.5 mm wide, 2.7-3 mm thick,

pale brown to glossy dark brown, adaxial surface

bifacial, abaxial surface rounded, micropylar

ridge 3.5-5 mm long; caruncle crescent shaped,

1-1.5 mmlong, 1.5-1.8 mm wide, cream-yellow.

Fig. 2.

Selected additional specimens: Queensland. North

Kennedy District: Cromarty, ridge above Bruce Highway

in Bowling Green Bay N.P., 19°28’S, 147°53’E, Jan

1993, Forster PIF12748 & Bean (BRI). South Kennedy

District: S.F. 658 Carawatha, 20°47’S, 148°34’E, Apr

1991, Forster PIF8190 & McDonald (BRI, K, F, MEF,

QRS). Feichhardt District: Melaleuca Creek Scrub,

“Rookwood”, 23°12’S, 149°46’E, Apr 1991, Forster

PIF7953 & McDonald (BRI, K, F, MEF, QRS). Port

Curtis District: Yaparabah, 10 km SSE of Mardale,

24°39’S, 150°42’E, Dec 1982, Forster PIF1481 &

Marshall (BRI); S.F. 53, Dan Dan Scrub, Dec 1987,

Gibson 1006 (BRI, NSW); Mt Archer road, 23°21’S,

150°35’E, Nov 1986, Hoy 129 (BRI); Mt Etna,

23°10’S, 150°27’E, May 1990, Vavryn 101 (BRI).

Burnett District: Scientific Area 33, Coominglah S.F.

28, 24°54’S, 151°00’E, Apr 1990, Forster PIF6696

(BRI, MEF, QRS); Sanderson’s Scrub, Mt Blandy, 4 km

W of Mingo Crossing, 25°24’S, 151°44’E, Mar 1999,

Forster PIF24148 & Booth (BRI, QRS); S.F. 695

Austrobaileya 6 (3): 349-436

Kalpowar, Burnett Range road, 24°40’S, 151°2rE, Mar

2000, Forster PIF25415 & Booth (BRI, K, F, MEF,

QRS). Wide Bay District: 1 km SW of Booyal, 25°13’S,

152°02’E, Nov 1987, Forster PIF3287 (BRI); Oakview

S.F. 220 Malmaison, 12 km ESE of Kilkivan, 26°08’S,

152°20’E, Dec 2002, Forster PIF29213 (A, BRI, F,

MEF, NSW, WIS); Black Gin Creek, T.R. 580, 25°29’S,

151°55’E, Apr 1990, Forster PIF6601 (BRI, CBG,

MEF, QRS); Fairlies Knob N.P., 25°30’S, 152°17’E,

Dec 1992, Forster PIF12593 & Smyrell (BRI, MEF,

QRS). Darling Downs District: S.F. 197 Diamondy, Craig

Range, 32 km NE of Jandowae, 26°35’S, 151°20’E,

Mar 1999, Forster PIF24094 & Booth (BRI, QRS).

Moreton District: Pullen Creek, Moggill S.F., Feb 1980,

Bird [AQ331172] (BRI); Commissioner’s View,

Blackbutt Range, 26°52’S, 152°13’E, Nov 1987,

Forster PIF3251 & Bird (BRI, NSW, SAN, SAR). New

South Wales. Wiangaree S.F., Jan 1981, Bird

[AQ345019] (BRI); Oxley River (Middle Arm Creek),

just beyond end of Butler’s Road, NW of Tyalgum,

29°19’S, 153°09’E, Jul 1981, Guymer 1585 & Jessup

(BRI); 23 km NW of Kyogle, Toonumbar Forest Road,

Toonumbar S.F., 28°29’S, 152°48’E, Dec 1991, Halford

Q823 (BRI, MEF, NSW).

Distribution and habitat : Croton

acronychioides is widespread in south-eastern

Queensland and north-eastern New South Wales,

but is present in only a few populations in the

South Kennedy and North Kennedy districts

(Map 3). The species has been recorded from a

total of seventeen 1° squares. Plants grow in

microphyll to notophyll vineforests on a variety

of volcanic substrates and may be sympatric with

C. insularis and C. phebalioides.

Phenology: Flowering occurs from November

to May and commences after storm rain. Fmiting

occurs from November to August.

Notes: F.Mueller (1864) cited two elements in

the protologue of Croton acronychioides, one

collected at Fitzroy River by Thozet and the

other collected at Broad Sound by Bowman.

There are three sheets at MEL (MEL231221,

231235, 231233) that are possible syntypes.

MEL231221 is sterile and MEL231233 has two

conflicting dates (“155/62” and “166/62”)

indicating a mixed collection. MEL231235 is

fertile, but lacks a date and the locality of Broad

Sound or Fitzroy River. The labels do indicate

that MEL231235 was collected by Bowman near

Rockhampton. Mueller (1864) definitely

described a fertile plant, hence a lectotype is

chosen from MEL231235 as it fulfills more criteria

to qualify as a syntype of the name than do the

other contenders.

u ^

Forster, Croton in Australia

359

Fig. 2. Croton acronychioides. A. flowering branchlet. x 0.8. B. base of leaf lamina showing extrafloral nectaries,

x 16. C. node with stipule, x 4. D. inflorescence with male flowers, x 2. E. undersurface of leaf, x 1. F. male flower

(lacking one stamen), x 8. G. female flower, x 8. H & I. fruits, x 4. J. seed, x 6. A, B, D from Forster PIF3251 (BRI);

& G. from Forster PIF12616 (BRI); E, H-J from Forster PIF3287 (BRI); F from Forster PIF12593 (BRI). Del.

Smith.

360

There are three specimens that may be

considered as candidates for a type of Croton

affinis because they were all collected at

Tintenbar by Baeuerlen. There are two at NSW,

NSW273959 and NSW273894, only the latter

has a date - Feb 1894. There is also one

specimen at MEL dated Aug 1893. A lectotype

is selected from the Feb 1894 specimen at NSW

as this would appear to predate the publication

of the name.

Conservation status : Widespread and common,

but infrequent and usually disjunct in the

northern parts of its range. Present in at least 20

conservation reserves in south-eastern

Queensland alone (Forster etal. 1991).

Etymology : The specific epithet probably refers

to a resemblance of the foliage of this plant to

that of some species of Acronychia (Rutaceae).

2. Croton aridus P.I.Forst., sp. nov. affinis

C. arnhemico Muell. Arg. a qua foliis 20-

36-dentatis, fructibus oblongis

majoribusque (13-17 x 10-13 mm), et

seminibus oblongiovoideis majoribusque

(8.5-10.5 mm longis) differt. Typus:

Northern Territory, c. 130 km S of Tennant

Creek on Stuart Highway, 20 July 1968,

J.Z. Weber 1084A(holo: AD [1 sheet]; iso:

BRI, DNA, MEL).

Croton sp. (Barkly Downs S.L.Everist 3379)

(Forster & Henderson 1997:72; Forster &

Halford 2002:70)

Subshrub to 1.5 m high, multistemmed,

monoecious, evergreen, perennial. Indumentum

silver. Branchlets rounded, with dense stellate

trichomes when young, glabrescent. Stipules

linear-lanceolate, 3-5 mm long, 0.4-0.6 mm wide,

entire and with dense stellate trichomes. Leaves

alternate, discolorous, petiolate; petioles 5-15

mm long, c. 1 mm wide, with dense stellate

trichomes when young, rarely glabrescent;

lamina ovate to broadly ovate, 15-80 mm long,

10-80 mm wide, palminerved with 3-5 veins from

base and 6-8 lateral veins per side of midrib

further up lamina, tertiary reticulate veins

present; upper surface silver-green, venation

weakly visible, with dense stellate trichomes

when young, becoming sparse with age; lower

surface pale silver-green, lateral and reticulate

venation prominent, with dense and ± persistent

Austrobaileya 6 (3): 349-436

stellate trichomes, velutinous; margins crenate

with 10-18 teeth up to 3 mm long, foliar glands

prominent; tip obtuse to acute; base cordate to

truncate; extrafloral nectaries at lamina base

often absent or obscure, if present then 1 or 2

at lamina base, sessile, elliptic, c. 0.3 mm long

and 0.2 mm wide, visible below only.

Inflorescence up to 40 mm long, unbranched,

usually androgynous, often reduced to single

female flower, pedunculate up to 5 mm; axis with

dense stellate trichomes; bracts

linear-lanceolate to lanceolate, 1-2.5 mm long,

0.3-1 mm wide, with dense stellate trichomes.

Male flowers 3-4 mm long, 5-6 mm diameter,

densely clustered in upper portion of

inflorescence; pedicels 6-7 mm long, c. 0.5 mm

wide, with dense stellate trichomes; sepals

valvate, 5, lanceolate-ovate, 3-4 mm long, 1.5-

2.5 mm wide, with dense stellate hairs; petals 5,

oblanceolate, 4-4.5 mm long, 0.8-1 mm wide,

lanate in upper half; stamens 20, filaments +

filiform, 2.2-3 mm long, c. 0.1 mm wide, glabrous;

anthers oblong, 0.7-0.8 mm long, 0.6-0.7 mm

wide. Female flowers 5-6 mm long, 4-5 mm

diameter, held singly and spaced 6 mm apart;

pedicels 3-8 mm long, c. 1 mm diameter, with

dense stellate trichomes; sepals valvate, 5,

lanceolate, 3-3.5 mm long, 1.7-2 mm wide, with

dense stellate trichomes; petals absent; styles

3, linear-obloid, up to 3 mm long, bifid once for

up to 2.8 mm long, + free at base, glabrous;

ovary 3-locular, c. 3 mm long and 3 mm diameter,

with dense, + sessile stellate trichomes. Fruits

trilobate, oblong-ovoid, 13-17 mm long, 10-13

mm diameter, with dense, + sessile stellate

trichomes. Seeds ± obloid, 8.5-10.5 mm long,

7-9 mm wide, 4-6 mm thick, matt brown, ventral

surface bifacial or + rounded, dorsal surface

rounded, micropylar ridge 8-8.8 mm long;

caruncle ovate, 1.5-mm long, c. 1.8 mm wide,

cream. Fig. 3.

Additional specimens : Western Australia. Near Edgar

Range, SE of Broome, 18°28’S, 123°03’E, Aug 1976,

Kenneally 5733 (CANB); c. 40 km NE of Callawa

Station HSD, E of Shay Gap, 20°26’S, 120°47’E, Aug

1997, Mitchell PRP1823 (BRI); 79 km SE of Broome

on Dampier Downs road, NW of Edgar Ranges, 18°15’S,

122°45’E, Jul 1989, White 150 (PERTH); 9 miles [15

km] N of Bonney Well, Aug 1963, Winkworth 1579

(DNA). Northern Territory. Barkly Tablelands,

Barkly Stock route, 19°52’S, 137°07’E, Mar 1988,

Brock 375 (DNA); 16 km WSW Soudan, 20°05’S,

136°48’E, Jun 1960, Chippendale NT7284 (AD, BRI,

CANB, MEL); Barkly Tableland, 40 km WNW

Forster, Croton in Australia

361

Fig. 3. Croton aridus. A. branchlet. x 1. B. undersurface of leaf, x 1.5. C. node showing stipule, x 6. D. base of leaf

lamina showing extrafloral nectaries, x 12. E. node with inflorescence, x 3. F. male flower, x 6. G. female flower,

x 6. H. seed, x 3. A-D, H from Everist 4243 (BRI); E from Chippendale 7284 (BRI); G & F from Weber 1084

(BRI). Del. W. Smith.

362

Frewena, 19°18’S, 135°02’E, Jun 1960, Chippendale

NT7349 (BRI, CANB, DNA); Singleton, 240 miles

[400 km] N of Alice Springs, Jan 1950, Everist 4243

(BRI); 98 km N of Annitowa, 20°24’S, 136°50’E, Mar

1981, Henshall 3454 (DNA); 10 km S of Wauchope,

20°45’S, 134°15’E, Jul 1977, Latz 7511 (AD, DNA);

2 km W of Lake Surprise, 20°13’S, 131°46’E, May

1984, Latz 9908 (DNA, PERTH); 2 km W of Lake

Surprise, Tanami Desert, 20°12’S, 131°45’E, Jun 1985,

Latz 10073 (AD, DNA); c. 35 km W of Green Swamp

Well No. 4 on road to Lajamanu, 19°14’S, 132°19’E,

Sep 1985, Leach 815 (DNA); c. 55 km NE of Green

Swamp Well No. 4, 18°49’S, 132°54’E, Sep 1986,

Leach 868 (DNA); 102.5 km W of the Stuart Highway

on track to Lajamanu, 19°22’S, 133°20’E, Mar 1988,

Leach 1708 (DNA); 82 miles [131 km] S of Tennant

Creek Jul 1968, Must 282 (AD, DNA). Queensland.

Burke District: Red Sand hills NE of Barkly Downs

Homestead, Jul 2002, Bailey & Kelman 1 (BRI); Barkly

Downs, c. 50 miles [83.3 km] SW of Camooweal, Dec

1947, Everist 3379 (BRI, CANB).

Distribution and habitat: Croton aridus is

widespread in arid central Australia in Western

Australia, the Northern Territory and Burke

district in Queensland (Map 2). The Western

Australian populations are markedly disjunct

from those in central Australia. Although the

species has only been recorded from fourteen

1° squares, it is likely that many further

populations will be found. Plants grow on red

sand plains or ridges in Triodia hummock

grasslands, mulga shrublands or open

woodland dominated by Ventilago viminalis.

Phenology: Flowering occurs sporadically

throughout the year following storm rain. Fruits

mature two or three months after flowering.

Notes: Croton aridus is allied to C. arnhemicus

and may be a sister-taxon that has adapted to

the arid-zone. Croton aridus differs most

markedly from C. arnhemicus in the leaves with

20-36 marginal teeth, the larger (13-17 mm long,

10-13 mm diameter) oblong fruit with larger

(8.5-10.5 mm long) oblong-ovoid seed. Croton

arnhemicus has leaves with 60-100 marginal

teeth, globose fruit (6-11 mm long, 7-11 mm

diameter) and smaller, obloid to ovoid seeds

(4-7 mm long).

Conservation status: Croton aridus is

widespread and common in its known range.

Etymology: The specific epithet refers to the

distribution of this species in arid parts of

Australia.

Austrobaileya 6 (3): 349-436

3. Croton armstrongii S.Moore, J. Linn. Soc.,

Bot. 45: 219 (1920). Type: Northern

Territory. Port Essington, Armstrong s.n.

(holo: BM).

Shrub, height unknown, monoecious,

?evergreen or deciduous, perennial.

Indumentum uncoloured. Branchlets ± rounded,

with scattered stellate trichomes, glabrescent.

Stipules lanceolate, 0.3-1 mm long, 0.2-0.3 mm

wide, entire and with scattered stellate

trichomes. Leaves alternate, discolorous,

petiolate; petioles 1-5 mm long, 0.5-0.8 mm

wide, with sparse stellate trichomes; lamina

ovate, 15-55 mm long, 5-25 mm wide,

palminerved with 3-5 lateral veins from base

and 5 or 6 lateral veins per side of midrib further

up lamina, tertiary reticulate veins poorly

developed; upper surface dark green, lateral

veins not visible, with scattered stellate

trichomes; lower surface pale green, venation

weakly developed, with scattered to sparse

stellate hairs, neither scabrous nor velutinous;

margins crenate with 15-20 teeth 0.5-1.5 mm

long, foliar glands prominent; tip acute; base

truncate; extrafloral nectaries 1 or 2 at lamina

base, subulate, c. 0.6 mm long and 0.2 mm wide,

visible above only. Inflorescence up to 50 mm

long, unbranched, usually androgynous,

pedunculate for up to 2 mm; axis with sparse

stellate trichomes; bracts lanceolate, 0.9-1.2 mm

long, 0.4-0.5 mm wide, with scattered stellate

trichomes. Male flowers c. 2.5 mm long, 3.5-

4 mm diameter, densely clustered towards the

inflorescence tip; pedicels 1-1.5 mm long, c.

0.2 mm wide, with sparse stellate trichomes;

sepals valvate, 5, lanceolate-ovate, 1.8-2 mm

long, 1.2-1.3 mm wide, with sparse stellate

trichomes; petals 5, oblanceolate, c. 1.6 mm long

and 1 mm wide, lanate in upper half and with

scattered stellate hairs on backs; stamens 12,

filaments + flattened, 1.4-1.7 mm long and c. 0.1

mm wide, glabrous; anthers oblong, c. 0.8 mm

long and 0.6 mm wide. Female flowers c. 3.5 mm

long and 5 mm diameter, held singly 1-5 mm

apart; pedicels 1.5-2 mm long, 0.7-0.8 mm

diameter, with dense stellate trichomes; sepals

valvate, 5, lanceolate-obovate, c. 4 mm long and

2 mm wide, with sparse stellate trichomes; petals

absent; styles 3, linear, 1.8-2 mm long, multifid,

twice divided over 1.4-1.8 mm, connate at base

for c. 0.3 mm long, with scattered simple and

stellate trichomes; ovary 3-locular, c. 2 mm long

Forster, Croton in Australia

and 2.5 mm diameter, with dense stellate

trichomes. Fruits and seeds not seen.

Specimens examined : Known only from the

type.

Distribution and habitat : Apparently at Port

Essington, Northern Territory (Map 6).

Phenology : Unknown.

Notes: Croton armstrongii remains an enigma.

Airy Shaw (1980) reduced his own Croton

habrophyllus to synonymy with the earlier

name, but did not consult the type specimen of

C. armstrongii at BM. Wilmot-Dear (1987)

subsequently reinstated Croton habrophyllus

demonstrating that the type of C. armstrongii

is not conspecific with that of the later name.

The type collection of Croton

armstrongii was supposedly made at Port

Essington. No further collections that are

conspecific with the type have been made in

the Northern Territory, despite an intensive

survey of closed forests by staff of the Northern

Territory Conservation Commission in the late

1980s. This may indicate that the taxon is not

present in Australia and that the type specimen

is incorrectly labelled. There is circumstantial

evidence for the BM specimen being incorrectly

labelled. An Armstrong collection at NSW

(NSW270599) that is labelled as Croton

armstrongii , is conspecific with the type of

C. habrophyllus. Armstrong also collected in

Vanuatu and the Banks Islands (Lanjouw &

Stafleau 1954), hence it is possible that the type

of Croton armstrongii originates from one of

these regions. Nevertheless the BM type is of a

distinct taxon warranting recognition. Whether

the taxon is native to Australia remains to be

determined. Until this is resolved there seems

little choice but to include the taxon as

Australian.

Conservation status: Unknown.

Etymology: Named for Sir Alexander Armstrong

(1818-1899) who made plant collections in

Melanesia and at Port Essington.

4. Croton arnhemicus Muell.Arg., Linnaea 34:

112 (1865); Oxydectes arnhemicus

(Muell.Arg.) Kuntze, Rev. Gen. PI. 2: 611

(1891). Type: Queensland. Cape York,

MacGillivray 514 (lecto [here

designated]: K n.v., photo at BRI!);

363

Northern Territory, ‘in Arnhemsland

Novae-Hollandiae septentrionalis’,

F. Mueller (lectopara: G-DC n.v., fiche at

BRI!); Sea Range, towards the

Fitzmaurice, October 1855, F. Mueller

(lectopara: MEL231274 & MEL231258).

Croton arnhemicus var. urenifolius Baill.,

Adansonia 6: 300 (1866). Type:

Queensland. Port Denison, E. Fitzalan

(syn: MEL231243, MEL231249 &

MEL231251); Edgecombe Height, Aug

1863, Dallachy (syn: MEL231246).

Croton arnhemicus var. typicus Domin,

Biblioth. Bot. 89: 329 (1827), nom. inval.

Type: same as C. arnhemicus Baill.

Illustrations: Brock (1988: 129); Dunlop

(1995:214, fig. 71).

Shrub to 5 m high, monoecious, evergreen,

perennial. Indumentum ferruginous-silver.

Branchlets rounded, with dense overlapping,

stellate trichomes when young, glabrescent.

Stipules linear to linear-lanceolate, 1.7-11 mm

long, 0.2-0.8 mm wide, entire and with dense

stellate trichomes. Leaves alternate,

discolorous, petiolate; petioles 3-75 mm long,

0.9-2 mm wide, with dense stellate trichomes

when young, rarely glabrescent; lamina elliptic,

orbicular, ovate, obovate, 20-240 mm long, 10-

200 mm wide, palminerved with 3-5 veins from

base and 5-7 lateral veins per side of midrib

further up lamina, tertiary reticulate veins

present; upper surface grey-green, venation

weakly visible, with scattered to sparse stellate

trichomes; lower surface ferruginous-silver,

lateral and reticulate venation prominent, with

sparse to dense, overlapping stellate trichomes,

scabrid or velutinous; margins denticulate to

weakly crenate with 30-50 teeth up to 2 mm

long, foliar glands prominent; tip acute to

rounded; base cordate, cuneate, rounded or

truncate; extrafloral nectaries 2 at lamina base,

sessile or stipitate to 1 mm long, ellipsoid, 0.5-

2 mm long, 0.5-1.2 mm wide, visible above

and below. Inflorescence up to 200 mm

long, unbranched, usually androgynous,

pedunculate up to 24 mm; axis with dense

stellate trichomes; bracts linear to lanceolate,

0.5-2 mm long, 0.2-0.7 mm wide, with dense

stellate trichomes. Male flowers 3.5-6 mm long,

3-7 mm diameter, held singly or rarely in groups

364

of 2-5 in upper portions of inflorescence;

pedicels 1.5-10 mm long, 0.5-1 mm wide, with

dense stellate trichomes; sepals valvate, 5,

lanceolate-ovate to ovate, 2-4.5 mm long, 1-2

mm wide, with sparse to dense stellate hairs;

petals 5, oblanceolate to obovate, 2-4.2 mm

long, 0.6-2.5 mm wide, lanate in upper half;

stamens 20-44, filaments ± filiform, 1.5-4.5 mm

long, 0.1-0.2 mm wide, glabrous; anthers

oblong, 0.6-1.1 mm long, 0.4-1 mm wide. Female

flowers 3-6 mm long, 3.5-6 mm diameter, held

singly and spaced up to 20 mm apart; pedicels

1-7 mm long, 0.4-2 mm diameter, with dense

stellate trichomes; sepals valvate, 5,

lanceolate-ovate to ovate, 2.2-3.5 mm long, 1-

2.2 mm wide, with dense stellate trichomes;

petals absent; styles 3, linear, 1.8-5.5 mm long,

bifid for 1.7-5.3 mm, connate at base for c. 0.1

mm, with sparse stellate trichomes; ovary 3-

locular, 2-4 mm long, 2-4 mm diameter, with

dense, sessile and stalked stellate trichomes.

Fruits trilobate, globose, 6-11 mm long, 7-11

mm diameter, with dense, sessile and stalked

stellate trichomes. Seeds obloid to ovoid, 4-7

mm long, 3-5 mm wide, 2.2-4 mm thick,

brown-black, ventral surface bifacial, dorsal

surface rounded, micropylar ridge 3-5 mm long;

caruncle ellipsoid-ovate, 0.8-1.5 mm long, 1.3-

2.2 mm wide, cream. Fig. 4.

Selected additional specimens : Northern Territory.

Humpty Doo, 12°32’S, 130°50’E, Sep 1984, Brock 19

(DNA); c. 12 miles [20 km] S of Katherine, Jan 1973,

Byrnes 2875 (BRI, CANB, DNA); Stuart Highway, Edith

Falls turnoff, 14°15’S, 132°01’E, Dec 1990, Evans

3476 (BRI, CANB, DNA); Vicinity of El Sharana, Jan

1973, Martensz & Schodde AE400 (BRI, CANB);

Upper East Alligator River, 13°01’S, 133°25’E, Nov

1987, Russell-Smith 4151 & Lucas (DNA); 9 km S

Koolpinyah Homestead, 12°28’S, 131°10’E,

Russell-Smith 8126 & Lucas (BRI, CANB, DNA).

Queensland. Cook District: Mt Scatterbrain, Butchers

Hill Station near Lakeland, 15°52’S, 144°53’E, Jan

1992, Forster PIF9523 (BRI, K, L, MEL, QRS); 13 km

past Chillagoe on Mungana road, 17°06’S, 144°24’E,

Jan 1992, Forster PIF9578 (BRI, DNA, L, MET,, QRS);

Mt Eliza, 8 km NW of Mt Surprise, 18°06’S, 144°15’E,

Jan 1993, Forster PIF12799 (BRI, MEL, QRS); Agate

Creek, Robinhood Station, 18°50’S, 143°25’E, Apr

1996, Forster PIF19076 et al. (BRI); Badu Island,

Torres Strait, 10°07’S, 142°07’E, Dec 1979, Garnett

253 (BRI); Archer River Crossing on Peninsula

Development road, 13°26’S, 142°55’E, Nov 1986,

Jessup 768 (BRI). North Kennedy District: Harold Island,

20°14’S, 149°09’E, Nov 1985, Batianoff 3456 &

Dalliston (BRI); Turkey Scrub, Whitewater, 18°10’S,

144°38’E, Jan 1993, Fensham 343 (BRI); Mingela

Bluff, 19°53’S, 146°45’E, Jan 1992, Forster PIF9414

Austrobaileya 6 (3): 349-436

& Bean (A, B, BRI, DNA, K, L, MEL, MO, NY, QRS);

Swamp Bay, Conway Range N.P., 20°16’S, 148°46’E,

Jan 1993, Forster PIF12738 (BRI, MEL, QRS); Nellie

Bay, Dingo Beach, 20°05’S, 148°30’E, Dec 1999,

Forster PIF25263 & Booth (BRI, JE, MEL, QRS); 23

km NNE of Proserpine, 11 km from Bruce Highway

on road to Dingo Beach, Box Creek crossing, 20°14’S,

148°31’E, Nov 1991, Halford Q671 (BRI, K, MEL,

QRS); Mt Inkerman, 12 km S of Home Hill, 19° 46’S,

147°30’E, Mar 1992, Halford Q877 (BRI, MEL).

SOUTH KENNEDY DISTRICT: on hill above Lake

Elphinstone, 21°33’S, 148°14’E, Jan 1978, Anderson

271 (BRI); Havilah, 20°58’S, 147°52’E, Dec 1992,

Fensham 542 (BRI).

Distribution and habitat : Croton arnhemicus

is widespread in northern Australia in the “Top

end” of the Northern Territory and northern

Queensland (Map 2). This species has been

recorded from forty-six 1° grid squares and is

undoubtedly the most widespread of the

Australian Croton taxa. Plants grow in

deciduous vinethickets or in open eucalypt

woodland on a variety of soil types but

predominantly hard laterites or limestone.

Phenology: Flowering and fruiting occurs

throughout the year following storm rain. Most

flowering records are from the October to

February period.

Notes: J. Mueller (1865) listed several syntypes

in the protologue of Croton arnhemicus. All of

these have been located and a lectotype is

selected from the best available specimen.

Croton arnhemicus is an extremely variable

species in terms of its habit and vegetative

morphology. Some variation seen in specimens

may be explained by its deciduous habit,

resulting in the early flowers of the season often

being collected with young foliage. Later

flowers and fruit are always collected with older

and more mature foliage. Plants in deciduous

vinethickets grow into quite large shrubs or small

trees up to 6 m in height, whereas those that

occur in the eucalypt woodlands in the Northern

Territory are often multistemmed due to regular

burning back of the above ground parts. These

Northern Territory plants develop into small

trees if fire is excluded, whereas the allied

Croton multicaulis P.I.Forst. in Queensland

always retains the multistemmed subshrub

habit. Croton arnhemicus and C. multicaulis

are superficially similar, and apart from the

difference in habit, C. arnhemicus has leaves

Forster, Croton in Australia

365

Fig. 4. Croton arnhemicus. A. flowering branchlet. x 0.5. B. undersurface of leaf, x 1. C. base of leaf lamina

showing extrafloral nectaries, x 6. D. node showing stipules, x 2. E. inflorescence with female flowers in lower half

and male flower buds in upper half, x 1. F. female flower, x 6. G. male flower, x 6. H & I. fruits, x 3. J. seed, x 4.

A, C, D from Champion 319 (BRI); B, E, F, G from Forster PIF9414 (BRI); H-J from Forster PIF9523 (BRI). Del.

W. Smith.

366

with 60-100 marginal teeth and male flowers with

20-44 stamens, whereas C. multicaulis has

leaves with 32-56 marginal teeth and male

flowers with 11-24 stamens.

Croton arnhemicus was included in

C. section Cascarilla Griseb. by Webster

(1993a), but transgresses the character states

for this section in the palmate venation (versus

pinnate) and the stamen number.

Conservation status : Very common.

Etymology : The specific epithet alludes to one

of the original syntypes being collected in

Arnhem Land.

5. Croton brachypus Airy Shaw, Muelleria 4:

224 (1980). Type: Queensland. Cook

District: Tozer Range, 0.5 mile [0.8 km] east

of Mt Tozer, 6 July 1948, L.J. Brass 19462

(holo: Krc.v.;iso: BRI, CANB).

Shrub to 3 m high, monoecious, evergreen,

perennial. Indumentum ferruginous. Branchlets

rounded, with scattered to sparse peltate

scales when young, glabrescent. Stipules

linear-lanceolate, c. 1.7 mm long, c. 0.7 mm wide,

entire and with sparse peltate scales. Leaves

alternate, discolorous, petiolate; petioles 2-5

mm long, 1-1.4 mm wide, with sparse peltate

scales; lamina narrow-ovate, oblanceolate or

obovate, 18-180 mm long, 12-70 mm wide,

penninerved with 7-11 lateral veins per side of

midrib, tertiary reticulate veins weakly

developed; upper surface matt dark green,

venation not visible, glabrous; lower surface

pale green, lateral and tertiary reticulate veins

weakly developed, glabrous or with scattered

peltate scales when young, neither scabrid nor

velutinous; margins entire or weakly sinuate,

foliar glands prominent; tip acute to shortly

acuminate; base cordate to truncate; extrafloral

nectaries 2 at lamina base, shortly stipitate to

0.5 mm long, circular, 0.4-0.5 mm long, 0.4-0.5

mm wide, visible below only. Inflorescence up

to 65 mm long, unbranched, usually

androgynous, pedunculate up to 12 mm; axis

with scattered peltate scales; bracts lanceolate,

0.5-0.8 mm long, 0.2-0.3 mm wide, with scattered

to sparse peltate scales. Male flowers 2-2.5 mm

long, 3.5-4 mm diameter, sparsely to densely

clustered on inflorescence; pedicels 2-2.5 mm

long, c. 0.4 mm wide, with sparse peltate scales;

Austrobaileya 6 (3): 349-436

sepals valvate, 5, lanceolate, 1.5-2 mm long, c.

1 mm wide, lanate in upper half; petals 5,

lanceolate, 1.8-2.4 mm long, 0.8-1 mm wide,

lanate in upper half; stamens 10-11, filaments

filiform, 2.3-2.5 mm long, c. 0.1 mm wide,

glabrous or with scattered simple trichomes at

base, anthers oblong, c. 0.8 mm long and 0.4

mm wide. Female flowers 3-3.5 mm long, 5-6

mm diameter, held singly and spaced 2-6 mm

apart; pedicels 0.8-1 mm long, c. 0.5 mm diameter,

with sparse peltate scales; sepals valvate, 5,

lanceolate, 1.8-2 mm long, 0.5-0.8 mm wide, with

sparse peltate scales; petals absent; styles 3,

linear, 2.3-2.5 mm long, bifid for 1.8-2 mm,

connate at base for c. 0.6 mm, glabrous; ovary

3- locular, 1-1.3 mm long, 1-1.3 mm diameter, with

dense, + sessile peltate scales. Fruits trilobate,

globose, 5-6 mm long, 5-6 mm diameter, with

sparse, + sessile peltate scales. Seeds ovoid,

4- 5 mm long, 3-4 mm wide, c. 3.5 mm thick,

brown and white blotched, ventral surface

bifacial, dorsal surface rounded, micropylar

ridge 2.8-3.8 mm long; caruncle obloid, 1.2-1.5

mm long, c. 0.4 mm wide, cream. Fig. 5.

Additional specimens: Queensland. Cook District:

Tozer Range, north end, Jun 1948, Brass 19355

(CANB); Lower northern slopes of Mt Tozer, 12°45’S,

143°15’E, Jun 1972, Dockrill 441 (BRI, QRS); Iron

Range N.R, 1.3 km NE of Mt Tozer, 12°44’S,

143°13’E, May 1994, Fell DGF4083 et al. (BRI); at

base of Paps, Tozer Gap, 12°43’S, 14°3°12’E, Jul 1991,

Forster PIF9093 (BRI, K, L, MEL, QRS); Garraway

Creek area, 12°43’S, 143°08’E, Jul 1993, Forster

PIF13552 & Tucker (BRI, K, L, MEL, QRS); ditto, Jul

1994, Forster PIF15441 (BRI); Puffdelooney Ridge,

12°45’S, 143°13’E, Jul 1972, Irvine 249 (BRI, QRS);

8.3 km E of Garraway Creek on road to Portland

Roads, 12°45’S, 143°14’E, Jul 1991, Neldner 3538 &

Clarkson (BRI, DNA, MBA, QRS); Hill E of Mt Tozer,

Iron Range area, 12°45’S, 143°13’E, Nov 1977, Tracey

14211 (BRI); Mt Tozer near Iron Range, 12°45’S,

143°12’E, Nov 1977, Tracey 14849 (BRI).

Distribution and habitat : Croton brachypus

appears to be endemic to the Iron Range area of

northern Cape York Peninsula in Queensland

(Map 3) and is known from only a single degree

square. Plants grow on creek banks or on ridges

in notophyll to mesophyll semi-deciduous

vineforest on volcanic substrates.

Phenology: Flowering occurs sporadically

throughout the year with records in October,

November, December, June and July. Fruiting

probably occurs two or three months later.

Forster, Croton in Australia

367

Fig. 5. Croton brachypus. A. flowering branchlet. x 0.6. B. undersurface of leaf, x 0.6. C. base of leaf lamina

showing extrafloral nectaries, x 4. D. node showing stipules, x 8. E. inflorescence with female flowers towards base

and male flowers towards tip. x 2. F. female flower, x 8. G. male flower, x 8. H & I. Fruit, x 4. J. seed, x 6. A-G from

Forster PIF13552 (BRI); H-J from Sankowsky 1445 (BRI). Del. W. Smith.

368

Notes: Croton brachypus is distinctive

amongst Australian taxa of Croton most notably

in the leaves with short petioles.

Conservation status : This species is apparently

endemic to a small area of Cape York Peninsula

but is locally common. The species is quite

common in Iron Range National Park. No

conservation coding is thought necessary.

Etymology: The specific epithet is derived from

the Greek brachy (short) and -pus (footed) and

presumably alludes to the leaves of this species.

6. Croton byrnesii Airy Shaw, Muelleria 4:225

(1980). Type: Northern Territory. Cannon

Hill, 18 December 1972, N. Byrnes 2833

(holo: DNA; iso: BRI, CANB; K n.v.).

Shrub to 4 m high, monoecious, deciduous,

perennial. Indumentum ferruginous to yellow.

Branchlets ± rounded, with scattered stellate

trichomes when young, glabrescent. Stipules

linear, 0.7-2.5 mm long, c. 0.2 mm wide, entire

and with scattered stellate trichomes. Leaves

alternate, discolorous, petiolate; petioles 10-45

mm long, 0.5-1 mm wide, with sparse stellate

trichomes; lamina elliptic, ovate, lanceolate-

ovate, 40-170 mm long, 15-90 mm wide,

penninerved with 11-13 lateral veins per side of

midrib and indistinct tertiary reticulate veins;

upper surface dark matt-green, lateral veins

weakly visible, glabrous; lower surface pale

green, venation weakly developed, with

scattered stellate trichomes, neither scabrid nor

velutinous; margins crenate with 20-29 short

teeth up to 0.5 mm long, foliar glands prominent;

tip acute, acuminate; base cuneate, rounded;

extrafloral nectaries 2 on petiole 0.4-1 mm below

lamina base, sessile or stipitate up to 0.5 mm,

ellipsoid, 0.7-1 mm long, 0.5-0.8 mm wide, visible

above and below. Inflorescence up to 150 mm

long, unbranched, androgynous or with mixed

glomerules, pedunculate up to 30 mm; axis with

scattered stellate trichomes; bracts lanceolate,

0.8-1.2 mm long, 0.3-0.4 mm wide, with scattered

stellate trichomes. Male flowers 2.2-2.5 mm long,

3-4 mm diameter, in dense glomerules of many

flowers clustered towards top of inflorescence;

pedicels 2.2-4 mm long, 0.4-0.5 mm wide,

glabrous or with scattered stellate trichomes;

sepals valvate, 5, ovate, 2-2.5 mm long, 1.3-1.5

mm wide, with lanate tip; petals 5, oblanceolate,

Austrobaileya 6 (3): 349-436

2-3 mm long, 0.7-0.8 mm wide, with lanate tip;

stamens 9-11, filaments flattened, 1.5-2 mm

long, c. 0.2 mm wide, glabrous, anthers oblong,

0.8-1 mm long, c. 0.7 mm wide. Female flowers

3.8-4 mm long, 3.5-3.8 mm diameter, densely

clustered with males, sometimes single and up

to 15 mm apart; pedicels 2.5-7 mm long, 0.5-0.9

mm diameter, with scattered stellate trichomes;

sepals valvate, 5, lanceolate to lanceolate-ovate,

2-3 mm long, 1-2.5 mm wide, with lanate tip;

petals absent; styles 3, linear, 1.8-2.7 mm long,

multifid, twice divided for 1-1.5 mm, connate at

base for 0.2-0.3 mm, glabrous; ovary 3-locular,

1.5-2.3 mm long, 2-2.3 mm diameter, with dense,

sessile stellate trichomes. Fruits trilobate,

depressed-globose, 4.5-5 mm long, 6-7 mm

diameter, with scattered, sessile stellate

trichomes. Seeds ± obloid, 3.5-4.5 mm long, 3.2-

4 mm wide, 2.5-2.8 mm thick, tan-brown, ventral

surface rounded to weakly bifacial, dorsal

surface rounded, micropylar ridge 2.3-2.5 mm

long; caruncle crescent-shaped, 1-1.5 mm long,

1.7-2.5 mm wide, cream. Fig. 6.

Additional specimens : Northern Territory. Cannon

Hill, 12°22’S, 132°56’E, Nov 1976, Airy Shaw

(DNA1079); East Alligator River, 12°29’S, 133°03’E,

Feb 1973, Dunlop 3235 (DNA); East Alligator River,

12°50’S, 133°22’E, Dec 1989, Dunlop 7628 (AD, BRI,

CANB, DNA, MEL, NSW) 1 mile [1.7 km] SW Cannon

Hill, Feb 1973, Martensz & Schodde AE648 (BRI,

CANB, DNA); 2.5 miles [4.2 km] N Cannon Hill

airstrip, Feb 1973, Martensz AE812 (BRI, DNA);

10 km S Cannon Hill, 12°28’S, 132°55’E, Nov 1983,

Russell-Smith 845 (BRI, CANB, DNA); Upper East

Alligator River, 12°49’S, 133°22’E, Oct 1987, Russell-

Smith 3860 & Lucas (BRI, DNA); Upper East Alligator

River, Arnhem Land, 12°50’S, 133°20’E, Apr 1988,

Russell-Smith 5283 & Lucas (DNA); 12 km E of

Mudginberri Homestead, Kakadu N.R, 12°35’S,

132°59’E, Jan 1991, Russell-Smith 8402 & Brock

(BRI, DNA, MEL).

Distribution and habitat: Croton byrnesii is

restricted to the headwaters of the Alligator

River in Arnhem Land (Map 4) where it has been

collected from two 1 degree grid squares.

Plants grow in fragmented vinethickets,

often dominated by Allosyncarpia ternata

S.T.Blake and Lophostemon lactifluus

(F.Muell.) Peter G. Wilson & Waterhouse, along

streams in sandstone gorges.

Phenology: Flowering occurs from November

to April, following storm or wet season rain.

Fruiting occurs two or three months later.

Forster, Croton in Australia 369

Fig. 6. Croton byrnesii. A. fruiting branchlet. x 0.8. B. undersurface of leaf, x 1. C. node showing stipule, x 8.

D. base of leaf lamina showing extrafloral nectaries, x 8. E. male flower, x 12. F. female flower, x 8. G & H. fruit,

x 4. I. seed, x 8. A,B,I from Brock 8402 (BRI); C-F from Russell-Smith 845 (BRI); G & H from Dunlop 7628

(BRI). Del. W. Smith.

370

Notes : Croton bymesii is a distinctive species

that has been confused at times by collectors

with C. habrophyllus . As noted by Airy Shaw

(1981), Croton bymesii differs from that species

in the near glabrescence of the foliage (the

trichomes ferruginous to yellow) and the long

stipitate foliar glands that are on the petiole 0.4-

1 mm below the lamina base.

Conservation status'. Croton bymesii is

restricted in its distribution but appears relatively

common in its known range. The species is

present in Kakadu N.P. No conservation coding

is thought necessary.

Etymology: The epithet honours the late Norm

Byrnes (1922-1998), former botanist at DNA and

BRI, and first collector of the species. Norm made

many pioneering collections of plants in eastern

Arnhem Land while resident botanist at DNA,

including the type of this species.

7. Croton capitatus Michx., FI. Bor.-Amer. 2:214

(1803); Pilinophytum capitatum (Michx.)

Klotzsch in Wiegmann, Arch. Naturges.

7: 255 (1841). Type: United States of

America. Illinois, Michaux (holo: P-

Michaux n.v., fiche at BRI!; iso: P-JU n.v.,

ficheatBRI!).

Illustrations: Small (1913: 454, fig. 2714);

James & Harden (1990:420).

Erect herb to 80 cm high, monoecious, annual.

Indumentum silver. Stems rounded, with dense

sessile and stalked stellate trichomes. Stipules

linear, 2-3 mm long, c. 0.1 mm wide, entire and

with dense stellate trichomes. Leaves alternate,

discolorous, petiolate; petioles 5-48 mm long,

0.7-0.8 mm wide, with dense sessile and stalked

stellate trichomes; lamina lanceolate-ovate,

oblong to ovate, 18-55 mm long, 9-25 mm wide,

palminerved with 3 veins at base and 3-5 lateral

veins per side of midrib further up the lamina,

tertiary reticulate veins obscure; upper surface

dark silver-green, venation obscure, with sparse

to dense stellate trichomes; lower surface silver,

venation weakly visible, with dense stellate

trichomes; margins entire to weakly sinuate,

foliar glands inconspicuous; tip acute to

rounded; base cuneate to truncate; extrafloral

nectaries absent at leaf lamina base.

Inflorescence up to 20 mm long, androgynous,

Austrobaileya 6 (3): 349-436

+ sessile; axis with dense sessile and stalked

stellate trichomes; bracts linear, 3^1 mm long, c.

0.1 mm wide, with dense stalked stellate

trichomes. Male flowers 2-3 mm long, 2^1 mm

diameter, held singly or in glomerules of 2-3

flowers on inflorescence, spaced up to 2 mm

apart; pedicels 1.8-2.5 mm long, c. 0.2 mm wide,

with dense stalked stellate trichomes; sepals

valvate, 5, obovate, c. 1.6 mm long and 1.2 mm

wide, with dense stalked stellate trichomes;

petals 5, oblanceolate, 1.6-2.2 mm long, 0.4-0.5

mm wide, lanate; stamens 10-12, filaments

flattened, 1.8-2 mm long, c. 0.2 mm wide,

glabrous, anthers oblong, 0.7-0.8 mm long, 0.4-

0.5 mm wide. Female flowers c. 11 mm long and

12 mm diameter, held singly and spaced up to 3

mm apart, ± sessile; sepals valvate, 6-8,

oblanceolate to obovate, 4-4.8 mm long, 0.5-2

mm wide, with dense stalked stellate trichomes;

petals absent; styles 3, linear, 3-3.5 mm long,

multifid, thrice divided, connate at base for c.

0.2 mm, with sparse sessile stellate trichomes;

ovary 3-locular, c. 2.5 mm long and 4 mm

diameter, with dense stalked and sessile stellate

trichomes. Fruits trilobate, globose, 7-9 mm long,

7-9 mm diameter, with dense, sessile and

stalked stellate trichomes. Seeds orbicular, c. 5

mm long, 4-4.5 mm wide, 2.5 mm thick, orange-

brown, ventral surface bifacial, dorsal surface

rounded, micropylar ridge 4.5-5 mm long;

caruncle crescent shaped, c. 0.7 mm long and

0.7 mm wide, brown-red.

Additional specimens: United States of America.

OHIO: near Cincinnati, Sep 1880, Lloyd [AQ206109]

(BRI). MISSOURI: St Louis, Aug 1878, Martindale

[AQ206110] (BRI). Australia. New South Wales.

Dubbo to Collie road, 35 miles NW of Dubbo & 15

miles S of Collie, Feb 1955, Wheeler (NSW293822).

Distribution and habitat: Croton capitatus is

native to the United States of America where it

is known from New Jersey, Illinois, Kentucky,

Missouri, Arkansas, Louisiana, Tennessee,

Texas, Oklahoma, Kansas, Iowa, Alabama and

North Carolina (Ferguson 1901; Small 1913;

Johnston 1959). It is reported as being sparingly

naturalised in agricultural land in southern New

South Wales south of Collie (Map 9) (James &

Harden 1990). I have seen only the cited

specimen and it needs to be determined if this

plant is still naturalised in Australia.

Phenology: Unknown in Australia.

Forster, Croton in Australia

Notes: Michaux (1803) did not specify a

herbarium location for the single cited collection

from Illinois and no mention of type material for

Croton capitatus was made by Ferguson

(1901). J ohnston (1959)sawa photo at GH of a

specimen in P but was not specific as to the

location within that herbarium. There is a

Michaux collection of Croton capitatus from

Illinois that is present in P-Michaux and P-JU.

This collection is presumed to be the type with

the P-Michaux sheet considered as the

holotype.

Webster (1993a) placed C. capitatus in Croton

section Pilinophyton (Klotzsch) A.Gray.

8. Croton capitis-york Airy Shaw, Muelleria 4:

226 (1980). Type: Queensland. Cook

District: Silver Plains Holding between

Rocky River and Massey Creek, 13

September 1973, GC. Stocker 1077 (holo:

QRS; iso: BRI, CANB).

Croton capitis-york var. pilosus Airy Shaw,

Muelleria 4:226 (1980). Type: Queensland.

Cook District: 2 km south of Temple Bay

Outstation, 12°22’S, 143°05’E, Sep 1976,

B. Hyland 8995 (holo: QRS).

Illustrations: Airy Shaw (1981:619, fig. 2A);

Forster (1991:571).

Shrub to 5 m high, monoecious, evergreen,

perennial. Indumentum silver. Branchlets

rounded, with dense peltate scales. Stipules

minute, triangular, < 0.3 mm long and 0.3 mm

wide, entire. Leaves alternate, discolorous,

petiolate; petioles 3-27 mm long, c. 1 mm wide,

with dense stellate trichomes when young,

glabrescent; lamina elliptic-ovate, chartaceous,

30-160 mm long, 12-60 mm wide, penninerved

with 7-11 lateral veins per side of midrib, tertiary

reticulate veins absent; upper surface grey-

green, lateral veins indistinct, with scattered to

sparse peltate scales; lower surface pale grey-

green to silver, lateral veins strongly developed,

with sparse to dense peltate scales, neither

scabrid nor velutinous; margins entire or weakly

sinuate, foliar glands inconspicuous; tip acute,

acuminate; base cuneate, truncate; extrafloral

nectaries 2, just below leaf lamina base, stipitate

to 0.6 mm long, ellipsoid, c. 0.9 mm long and 0.5

mm wide, visible above and below. Inflorescence

up to 70 mm long, unbranched, mainly unisexual

371

but occasionally androgynous, pedunculate up

to 20 mm; axis with dense peltate scales; bracts

lanceolate, c. 0.8 mm long and 0.3 mm wide, with

dense peltate trichomes. Male flowers 1.8-2.3 mm

long, 3.5^1 mm diameter, densely clustered on

inflorescence in glomerules of 3-5 flowers, or

spaced to 5 mm apart; pedicels 1.4-1.5 mm long,

c. 0.5 mm wide, with dense peltate scales; sepals

valvate, 5, lanceolate-ovate, 1.9-2 mm long, 1.3-

1.4 mm wide, lanate in upper half; petals 5,

lanceolate-ovate, 1.4-1.5 mm long, c. 0.6 mm

wide, lanate in upper half; stamens 10-12,

filaments filiform, 1.5-2.2 mm long, c. 0.1 mm

wide, glabrous, anthers oblong, 0.8-0.9 mm

long, 0.4—0.5 mm wide. Female flowers not seen;

styles 3, obloid, up to 1.3 mm long and 0.2 mm

wide, bifid for up to 0.8 mm long, glabrous. Fmits

trilobate, subglobose, 6-7 mm long, 7-8 mm

diameter, with dense, stellate trichomes on

mamillate protuberances. Seeds ± obloid, 4.9-5

mm long, 3.2-3.3 mm wide, c. 3 mm thick, pale

glossy brown, ventral surface bifacial, dorsal

surface rounded, micropylar ridge c. 3.4 mm

long; caruncle weakly crescent shaped, c. 1.4

mm long and 1 mm wide, tan-yellow. Fig. 7.

Selected additional specimens: Queensland. Cook

District: 4 km SW of Cape Weymouth - Scrubby Creek,

12°38’S, 143°24’E, May 1990, Fell DGF2118 (BRI,

QRS); Scrubby Creek, between the Rocky and Chester

Rivers, Silver Plains, 13°46’S, 143°30’E, Dec 1990,

Fell DGF2286 (BRI); Kalpowar Pastoral Holding, 10

km ESE of the Normanby River mouth, 14°26’S,

144°13’E, Nov 1992, Fell DGF2750 & Stanton (BRI,

QRS); 4.5 km WSW of the Nesbit River mouth, 57 km

NE of Coen, 13°33’S, 143°22’E, Aug 1993, Fell

DGF3451 et al. (BRI); Carron Valley road, 44 km from

Moreton Telegraph Station, 12°29’S, 142°57’E, Jun

1988, Forster PIF4547 & Tucker (BRI); Maloneys

Springs, 40 km E by road of Moreton Telegraph Station,

12°28’S, 142°55’E, Jun 1989, Forster PIF5468 (BRI,

DNA, LAE); 2 km NW of Bolt Head, Temple Bay,

12°15’S, 143°04’E, Jul 1991, Forster PIF8957 (BRI,

DNA, K, MEL, QRS); 27 km along road to Leo Creek

mine, Mcllwraith Range, 13°42’S, 143°17’E, Forster

PIF10050 et al. (BRI, K, L, MEL, QRS); 31 km along

road to Leo Creek mine, Mcllwraith Range, 13°42’S,

143°18’E, Jun 1992, Forster PIF10262 et al. (BRI,

QRS); T.R. 9, Lankelly Creek, 13°53’S, 143°14’E, Jun

1992, Forster PIF10331 et al. (BRI, MEL, QRS); 3

km N of Massy Creek Crossing, Silver Plains Station,

13°53’S, 143°31’E, Jun 1992, Forster PIF10579 et al.

(BRI, QRS); 3 km SSW of Rocky River Crossing, Silver

Plains, 13°49’S, 143°27’E, Jul 1993, Forster PIF13668

et al. (BRI); Captain Billy Landing turnoff, on Coen

to Bamaga road, 11°41’S, 142°41’E, Jun 1994, Forster

PIF15360 (BRI, MEL, NSW, QRS); West Claudie River,

12°45’S, 143°15’E, Jun 1972, Hyland 6170 (BRI, QRS);

Olive River, 12°10’S, 143°05’E, Sep 1974, Hyland

372

7449 (BRI, QRS); 0.5 km from main Peninsula road,

on Captain Billy road, 11°41’S, 142°42’E, Feb 1992,

Johnson 4945 (BRI, DNA); Nesbit River, 13°26’S,

143°10’E, Sep 1974, Tracey 14103 (BRI); North bank

of Olive River near mouth, 12°07’S, 143°05’E, Sep

1974, Tracey 14488 (BRI); Bamaga Mission, 11.2 km

SW of Cape York, Jan 1965, Smith 12393 (BRI); Mt

Tozer, Iron Range area, 12°45’S, 143°15’E, Oct 1968,

Webb & Tracey 8702 (BRI).

Distribution and habitat: Croton capitis-york

is restricted to northern Cape York Peninsula in

Queensland (Map 5) where it has been collected

from five 1° grid squares. Plants grow in semi-

deciduous or evergreen microphyll to notophyll

vinethickets and vineforest on sandy soils

derived from sandstone.

Phenology: Flowering records are few and have

been made in April and June. Fruiting probably

occurs two or three months later. This species

generally has buds present for most of the year

and probably flowers after storm or wet season

rain.

Notes: The recognition of varieties in Croton

capitis-york has been previously refuted

(Forster 1991). A putative hybrid individual

between this species and Croton dockrillii was

found at the Rocky River, Silver Plains ( Forster

13669 et al. : BRI, QRS).

At Bolt Head there appears to exist a mixed

population of typical C. capitis-york and

atypical plants that are more densely silver

pubescent below. These atypical plants

[Forster PIF19391 (BRI, MEL) & PIF19406 (A,

BRI, K, MEL, QRS)] also have leaf lamina glands

that are visible only from below due to their

position at the base of the leaf lamina. In typical

Croton capitis-york these glands are at the

junction of the petiole and lamina and are visible

from both above and below the leaf. These

atypical plants have not been collected in a

fertile state so their status cannot be adequately

resolved at this stage, although it is likely that

they may represent an undescribed taxon.

Croton capitis-york shares the unusual

character of fruit with stellate trichomes on

mamillate protuberances, as found also in

C. mamillatus and C. stigmatosus from southern

Queensland.

Conservation status: Croton capitis-york is

not rare or threatened. It is present in

conservation reserves at Iron Range and

Heathlands National Parks.

Austrobaileya 6 (3): 349-436

Etymology: The specific epithet alludes to the

occurrence of this plant on Cape York Peninsula.

9. Croton caudatus Geisel., Croton. Monograph.

73 (1807). Type: ex India orient., Dr. Rottler

(holo: C n.v., fiche at BRI!).

Woody scrambler or liane, monoecious,

deciduous, perennial. Indumentum uncoloured.

Branchlets somewhat angular, with dense

stellate trichomes when young, glabrescent.

Stipules linear-subulate, 2.6-8 mm long, 0.3-0.58

mm wide, entire or divided and with dense

stellate trichomes. Leaves alternate, discolorous,

petiolate; petioles 2-3 mm long, c. 1 mm diameter,

with dense stellate trichomes; lamina elliptic to

ovate, 12-140 mm long, 5-100 mm wide,

palminerved with 5 veins at base and 3-6 lateral

veins per side of midrib further up the lamina,

tertiary reticulate veins obscure; upper surface

dark matt-green, lateral veins weakly visible, with

scattered to sparse, stellate trichomes; lower

surface pale silver-green, venation weakly

visible, with dense stellate trichomes,

glabrescent, neither scabrid nor velutinous;

margins crenate with 25-30 short teeth up to 2

mm long, foliar glands prominent; tip acute,

acuminate; base cuneate, cordate, lobate;

extrafloral nectaries 2 at lamina base, sessile or

stipitate up to 1 mm, circular, c. 0.6 mm diameter,

visible above and below. Inflorescence up to

250 mm long, unbranched, usually unisexual but

occasionally androgynous, pedunculate up to

50 mm; axis with dense stellate trichomes; bracts

linear-lanceolate, 1.2-6 mm long, 0.2-0.3 mm

wide, with sparse to dense stellate trichomes.

Male flowers 5-7 mm long, 6-8 mm diameter,

held singly or in glomerules of 1-6 flowers on

inflorescence, spaced up to 2 mm apart; pedicels

3-9 mm long, c. 0.5 mm wide, with dense stellate

trichomes; sepals valvate, 5, lanceolate-ovate,

2-3 mm long, 1.2-1.8 mm wide, with dense

stellate trichomes; petals 5, obovate, 2.5-3.5 mm

long, 1.2-1.7 mm wide, lanate; stamens 23-36,

filaments filif orm, 3-5.5 mm long, c. 0.1 mm wide,

glabrous, anthers oblong, 0.8-0.9 mm long, 0.6-

0.7 mm wide. Lemale flowers 3.5^1 mm long, 5-

7 mm diameter, held singly and spaced up to 10

mm apart; pedicels 1-3.5 mm long, 1-1.2 mm

diameter, with sparse stellate trichomes; sepals

valvate, 5, lanceolate to lanceolate-ovate, 2.8-6

mm long, 1.5-2.5 mm wide, with stellate

trichomes; petals absent; styles 3, linear, 3-5.5

mm long, bifid for 2.5-5 mm, connate at base for

Forster, Croton in Australia

373

Fig. 7. Croton capitis-york. A. habit, x 0.75. B. leaf base showing extrafloral nectaries, x 0.75. C. detail of leaf

surface showing stellate trichomes. x 12. D. stalked stellate trichome from leaf, x 25. E. male flower, x 6. F & G.

fruit, x 3. H. dehisced coccus, x 3. J. seed, x 3. A-E. from Clarkson 3641 (BRI); F-J from Morton 905 (BRI). Del.

M. Saul. Plate reproduced with permission from Forster (1991: 571).

374

0.2-0.6 mm, with scattered stellate trichomes in

lower half; ovary 3-locular, 2.8-3.5 mm long, 2.8-

3.5 mm diameter, with dense, sessile and stalked

stellate trichomes. Fruits ± globose, 14-22 mm

long, 14-24 mm diameter, with dense, sessile

and stalked stellate trichomes. Seeds ± ovoid,

c. 9 mm long, 7 mm wide, 5 mm thick, brown,

ventral surface rounded to weakly bifacial,

dorsal surface rounded, micropylar ridge c. 7

mm long; caruncle flattened-ovate, 1-1.5 mm

long, c. 2.5 mm wide, pale-brown. Fig. 8.

Selected additional specimens'. Malaysia. S. Pahang,

Fort Iskandar, Mar 1959, Woods 1716 (L); Sumatra,

Palembang, forest N of G. Roepit, Forbes 2572 (L);

Sarawak, G. Berloban, 10 km from Tebakang, Tebedu

road, Jul 1982, Yii & Othman S46180 (L); Sabah,

Limbuah darat Banggi Island, Aug 1964, Ampuria

SAN40384 (L); Hap Seng Plantation road to Sg.

Tangkulap, Karamuak, Jun 1982, Sundaling SAN90427

(L). Singapore. Bukit Timah Nature Reserve, Nov

1982, Maxwell 82-286 (L). Indonesia, Kalimantan,

East Kutai Resrve, vicinity of Sengata & Mentoko

rivers, 0°30’N, 117°20’E, Dec 1978, Leighton 354

(L); Java, Prov. Besuki, 1895, Koorders 20572(L);

Sulawesi, northern central part, near Palu, 0°53’S,

119°53’E, Apr 1975, Meijer 9200 (L). Philippines.

Palawan, May 1913, Merrill 1243 (BRI, L). Australia.

Christmas Island. Bordering settlement, Poon San

road, 10°26’S, 105°42’E, Sep 1984, Mitchell 167

(CANB). Queensland. Cook District: Chili Creek,

12°39’S, 143°23’E, Jul 1993, Forster PIF13570 et al.

(BRI, MEL); ditto, Jan 1982, Hyland 11568 (QRS);

ditto, Mar 1982, Hyland 11742 (QRS); ditto, Dec 1982,

Hyland 12432 (QRS).

Distribution and habitat : Croton caudatus is

widespread in western parts of Malesia and Asia

(Bangledesh, Bhutan, China, India, Myanmar,

Nepal, Pakistan, Sri Lanka, Thailand)

(Chakrabarty & Balakrishnan 1997), but is

known in mainland Australia only from one

population at Chili Creek on Cape York Peninsula

(Map 7). It is also recorded from Christmas

Island in the Indian Ocean, an Australian

territory (Du Puy & Telford 1993). Plants grow

as canopy lianes in semi-deciduous notophyll

vineforest.

Phenology : In Australia flowering occurs from

December to January and fruiting from March

to April.

Notes : Geiseler (1807) based his taxa on

specimens in the Vahl herbarium. This

herbarium is now deposited at C and sheet 15

appears to represent type material of Croton

caudatus. A long list of synonyms is given for

Austrobaileya 6 (3): 349-436

this species by Chakrabarty & Balakrishnan

(1997) and is not repeated here.

Croton caudatus is the only climbing Croton

in Australia and this lifeform is very rare in the

genus throughout its range (Chakrabarty &

Balakrishnan 1997; Secco & Rosa 1992).

Conservation status: Croton caudatus is very

restricted in its Australian occurrence and has

perhaps arrived in Australia by accidental

human intervention at some time. The only

known population is within the Iron Range

National Park. No conservation coding is

thought necessary at this stage, however, the

population could be construed as being

endangered due to its proximity to a road.

Etymology: The specific epithet is derived from

Latin and probably refers to the shape of the

leaf base in this species.

10. Croton choristadenius K.Schum., Nachtr.

295 (1905). Type: Papua New Guinea.

Augusta-Station, September 1887,

M. Hollrung 705 (syn: K n.v., photo at

BRI!; L n.v.); Ramufluss, 15 July 1898,

Tappenbeck 116 (syn: n.v.).

Crotonphilombros Croizat, J. Arnold Arb. 23:

371 (1942). Type: Papua New Guinea.

Western Province: Penzara, between the

Morehead & the Wassi Kussa Rivers,

December 1936, L.J. Brass 8455A(holo: A

n.v.; iso: BRI; L n.v).

Croton pusilliflorus Croizat, J. Arnold Arb.

23: 374 (1942). Type: Papua New Guinea.

Western Province: Palmer River, below

the junction with the Black River, July

1936, L.J. Brass 7226 (holo: An.v.; iso: BRI;

Ln.v).

Croton semunculus Croizat, J. Arnold Arb.

23:374 (1942). Type: Papua New Guinea.

Central Province: Nakeo district, Baroka,

10 April 1933, L.J. Brass 3770 (holo: An.v.;

iso: BRI).

Shrub or tree to 10 m high, monoecious,

evergreen, perennial. Bark nondescript,

somewhat tessellated; blaze reddish-pink; wood

straw. Indumentum silver or silver-ferruginous.

Branchlets ± rounded, with admixture of dense

stellate trichomes, peltate trichomes and peltate

Forster, Croton in Australia

375

Fig. 8. Croton caudatus. A. fruiting branchlet. x 0.5. B. base of leaf lamina showing extrafloral nectaries, x 8. C.

node showing stipule, x 4. D. inflorescence with male flower buds, x 2. E. male flower, x 6. F. female flower, x 8.

G & H. fruits, x 1.5. I. seed, x 3. A, B, G, H from Hyland 11568 (BRI); C from Forster PIF13570 (BRI); D-F from

Hyland 12432 (QRS); I from Hyland 11742 (QRS). Del. W. Smith.

376

scales, glabrescent. Stipules linear, 0.5-1 mm

long, 0.1-0.2 mm wide, entire and with sparse to

dense peltate scales. Leaves alternate,

discolorous, petiolate; petioles 10-40 mm long,

0.7-1 mm wide, with scattered peltate trichomes

and peltate scales; lamina elliptic to ovate, 30-

110 mm long, 15-40 mm wide, penninerved with

10-12 lateral veins per side of midrib, tertiary

reticulate veins indistinct; upper surface dark

green, venation weakly visible, glabrous; lower

surface pale green, lateral veins weakly

prominent, with widely scattered, peltate

trichomes, neither scabrid nor velutinous;

margins crenate to denticulate with 20-36 teeth

up to 0.2 mm long, foliar glands prominent; tip

acute, short to long acuminate; base rounded;

extrafloral nectaries 2 at base of lamina, sessile,

ellipsoid, 0.8-1.2 mm long, 0.7-0.8 mm wide,

visible below only. Inflorescence up to 120 mm

long, androgynous, pedunculate up to 20 mm;

axis with sparse peltate trichomes; bracts linear-

lanceolate, 0.8-1.3 mm long, 0.2-0.5 mm wide,

with sparse, peltate trichomes. Male flowers 2.5-

3 mm long, 2.5-3 mm diameter, evenly

distributed in upper 3/4 of inflorescence;

pedicels 1.5-2 mm long, c. 0.4 mm wide, with

sparse peltate trichomes; sepals valvate, 5,

lanceolate-triangular, 1.3-1.5 mm long, 0.7-0.8

mm wide, with scattered peltate trichomes,

lanate at tip; petals 5, oblanceolate, 1.8-2 mm

long, c. 0.5 mm wide, lanate near tip and near

base; stamens 11 or 12, filaments filiform, 1.2-

1.5 mm long, c. 0.1 mm wide, glabrous, anthers

oblong, 0.5-0.6 mm long, 0.5-0.8 mm wide.

Female flowers 2.5-3 mm long, 2.5-3 mm

diameter, held singly and spaced up to 7 mm

apart; pedicels 1-1.5 mm long, c. 1 mm diameter,

with sparse to dense, peltate trichomes; sepals

valvate, 5, lanceolate, 1.5-1.8 mm long, 1.2-1.3

mm wide, with scattered peltate trichomes and

stellate trichomes; petals absent; styles 3, linear-

flabellate, 1-1.2 mm long, bifid for 0.7-0.9 mm,

with scattered peltate trichomes near base;

ovary 3-locular, 1.8-2 mm long, 1.4-1.5 mm

diameter, with dense, peltate trichomes and

occasional stellate trichomes. Fruits trilobate,

depressed-globose, 4-5 mm long, 5-6 mm

diameter, with sparse, sessile stellate trichomes.

Seeds ± obloid, 3-3.5 mm long, 2.5-2.8 mm wide,

2-2.2 mm thick, glossy brown, ventral surface

bifacial, dorsal surface rounded, micropylar

ridge 1.8-2 mm long; caruncle ellipsoidal, c. 1.8

mm long and 1 mm wide, yellowish. Fig. 9.

Austrobaileya 6 (3): 349-436

Additional specimens : Papua New Guinea. Morobe

Province: Lower Inokanda L.A., Bulolo, 7°10’S,

146°40’E, Jun 1962, Havel NGF9168 (BRI); Crooked

Creek L.A., Bulolo, Jul 1964, Havel NGF25550 (BRI);

Bulolo Valley, 7°10’S, 146°40’E, Oct 1955, McVeigh

& Ridgwell NGF7368 (BRI); Busu River, 7°25’S,

147°10’E, Aug 1970, Streimann NGF45101 (BRI).

Central Province: c. 2 miles W of Kanosia Plantation,

Jul 1962, Darby shire 602 (BRI); Cape Rodney, TP 107,

near P.I.T. Sawmill, Jun 1968, Henty NGF38618 (BRI).

Northern Province: near Davatutu Village, Jul 1953,

Hoogland 3396 (BRI); Oive Ridge, Jul 1964, Millar

NGF23521 (BRI). Australia, Queensland. Cook

District: Macrossan Range, Turrel Hill, Silver Plains,

13°30’S, 143°30’E, Jul 1997, Forster PIF21326 et al.

(A, AD, BRI, DNA, L, MEL, MO, NSW, QRS); ditto,

Forster PIF21327 et al. (A, AD, BRI, DNA, MEL,

NSW, QRS).

Distribution and habitat: Croton

choristadenius occurs in Papua New Guinea in

the Morobe, Central and Northern Provinces

and is here newly recorded from a single locality

on Cape York Peninsula in Queensland (Map 3)

in the most speciose grid square in Australia

for Croton taxa. At Turrel Hill Croton

choristadenius forms a canopy tree in low semi-

deciduous complex notophyll vineforest on

metamorphic derived substrate.

Phenology: Flowers were recorded in July in

Queensland. Specimens from Papua New

Guinea have had flowers in the months of June-

August and fruit in the months of July and

October.

Notes: Croton choristadenius was renamed

several times by Croizat (1942) based on a series

of collections by Brass in southern New Guinea.

These names were subsequently reduced to the

synonymy of C. choristadenius by Airy Shaw

(1980a).

This species was included in Croton

section Tiglium (Klotzch) Baill. by Webster

(1993a), but transgresses the characters given

for that section in the admixture of peltate scales

and stellate trichomes (versus stellate trichomes

only).

Croton choristadenius is one of four

Australian species of Croton that form small trees,

the others being C. insularis, C. phebalioides

and C. stigmatosus. Consequently I have

included brief descriptors on bark, blaze and wood

in the descriptions for these taxa.

Forster, Croton in Australia

377

Fig. 9. Croton choristadenius. A. flowering branchlet. x 0.6. B. undersurface of leaf, x 1. C. base of leaf lamina

showing extrafloral nectaries, x 6. D. inflorescence with female flowers in lower half and male flower buds in upper

half, x 2. E. male flower, x 10. F. female flower, x 10. A-D, F from Forster PIF21326 (BRI); E from Forster

PIF21327 (BRI). Del. W. Smith.

378

Conservation status : In Queensland Croton

choristadenius is a very rare tree and has only

been seen on one ridge of Turrel Hill where it

was locally common over about a hectare.

Further investigation of other parts of Turrel

Hill and the adjacent Xena Hill in 1998 did not

reveal further individuals of this species.

Further unexplored hills in the Macrossan Range

require examination for additional populations

of this species. Croton choristadenius has not

been found in Iron Range and surrounds or near

Bamaga, so it may be reasonable to assume that

it is truly restricted in its Queensland

occurrence. Turrel Hill is now included in

Aboriginal controlled land and the future of land

management in favour of rare plants in this area

remains uncertain.

Etymology : From the latin chorisis (to split into

parts) and adenius, and probably referring to

the leaf lamina glands.

11. Croton densivestitus C.T.White &

W.D.Francis, Proc. Roy. Soc. Queensland

35: 80-83, fig. 9 (1923) (‘densivestitum’).

Type: Queensland. Cook District:

Harvey’s Creek, 1889, F.M. Bailey (holo:

BRI; iso: MEL).

Crotonpubens Domin, Biblioth. Bot. 89: 882,

t. 31, fig. 11-19 (1928). Type: Queensland.

Cook District: Harvey’s Creek and near

estuary of Russell River, 1909-10, Domin

(holo: ?PRn.v.).

Shrub to 3 m high, monoecious, evergreen,

perennial. Indumentum yellow. Branchlets

rounded, with dense stellate trichomes. Stipules

linear-lanceolate, 3.5-4 mm long, 0.3-1 mm wide,

entire and with dense stellate trichomes. Leaves

alternate, discolorous, petiolate; petioles 3-30

mm long, 1-1.5 mm wide, with dense stellate

trichomes; lamina elliptic to ovate, 40-180 mm

long, 20-80 mm wide, penninerved with 12-14

lateral veins per side of midrib, tertiary reticulate

veins obscure; upper surface dark green,

venation not visible, glabrous or with scattered

stellate trichomes; lower surface pale green,

lateral veins weakly prominent, with dense,

stellate trichomes, markedly velutinous; margins

denticulate with 20-23 teeth up to 0.2 mm long,

foliar glands prominent; tip acute, short to long

acuminate; base rounded, weakly cordate;

extrafloral nectaries 2 at base of lamina, stipitate

Austrobaileya 6 (3): 349-436

to 1.5 mm, circular, 0.7-0.9 mm long, 0.7-0.9 mm

wide, visible below only. Inflorescence up to 70

mm long, androgynous, pedunculate up to 20

mm; axis with dense stellate trichomes; bracts

lanceolate, 0.8-1.3 mm long, 0.2-0.3 mm wide,

with sparse to dense stellate trichomes. Male

flowers c. 2.5 mm long and 3.5 mm diameter,

densely clustered towards top of inflorescence;

pedicels 1.8-2 mm long, 0.1-0.3 mm wide, with

dense stellate trichomes; sepals valvate, 5,

lanceolate to lanceolate-ovate, 1.3-1.6 mm long,

c. 0.8 mm wide, with dense stellate trichomes,

lanate at tip; petals 5, oblanceolate, c. 1.5 mm

long and 0.4 mm wide, lanate in upper half;

stamens 11 or 12, filaments filif orm, 1.5-1.8 mm

long, c. 0.1 mm wide, glabrous, anthers oblong,

0.4-0.5 mm long, c. 0.3 mm wide. Female flowers

2.5- 3.5 mm long, 3—4.5 mm diameter, held singly

and spaced up to 10 mm apart; pedicels 1.2-2

mm long, 0.5-0.6 mm diameter, with dense

stellate trichomes; sepals valvate, 5, lanceolate,

1.3-1.8 mm long, 0.4-0.8 mm wide, with dense

stellate trichomes; petals absent; styles 3, linear,

2.5- 3 mm long, bifid for 1.5-2.2 mm, with sparse

stellate trichomes in lower half; ovary 3-locular,

1-2 mm long, 1.2-2 mm diameter, with dense, +

sessile stellate trichomes. Fruits trilobate,

depressed-globose, 4.5-6 mm long, 6-8 mm

diameter, with scattered, + sessile stellate

trichomes. Seeds ± ovoid, 4.5^4.8 mm long, 3.8-

4.5 mm wide, 2.5-3 mm thick, brown and white

blotched, ventral surface ± rounded, dorsal

surface rounded, micropylar ridge 3-3.5 mm

long; caruncle truncate-ovate, c. 0.5 mm long

and 1.2 mm wide, cream. Fig. 10.

Additional specimens examined : Queensland. Cook

District: Harvey’s Creek, s.dat., Bailey [AQ202099]

(BRI); Mt Bellenden Ker, 17°16’S, 145°54’E, Sep 1992,

Christensen 792 (AD, BRI); Carrington road,

Carrington, S of Atherton, 17°18’S, 145°27’E, Jun

1999, Ford 2236 (BRI, QRS); Base of Bellenden Ker,

17°15’S, 145°53’E, Jul 1993, Forster PIF13740 &

Lyons (BRI, MEL, QRS); Harvey’s Creek headwaters,

17°15’S, 145°53’E, Jul 1994, Forster PIF15499 et al.

(BRI); Bellenden Ker Cable Car Station, 17°16’S,

145°54’E, Jan 2002, Forster PIF28215 et al. (A, BRI,

K, L, MEL, WIS); Cooroo Lands, W of Innisfail,

17°31’S, 145°53’E, Jul 1971, Hyland 5259 (BRI, QRS);

S.F.R. 607, Parish of Cairns, Shoteel L.A., 16°56’S,

145°36’E, Oct 1991, Hyland 14258 (QRS); N.P.R.

226, Bellenden Ker, 17°16’S, 145°53’E, Aug 1992,

Hyland 14517 (QRS); Y.C.L. Parish of Glady, 17°31’S,

145°56’E, Sep 1992, Hyland 14583 (QRS); Harvey

Creek, c. 1.3 km upstream from Bruce Highway,

17°15’S, 145°54’E, Jan 1994, Jago 3061 (BRI);

Warramami Hill, c. 11 km by road W of Bruce Highway

Forster, Croton in Australia

379

Fig. 10. Croton densivestitus. A. flowering branchlet. x 0.4. B. undersurface of leaf, x 0.6. C. base of leaf lamina

showing extrafloral nectaries, x 4. D. node showing stipules, x 4. E. inflorescence with female flowers towards base

and male flower buds towards apex, x 2. F. male flower, x 12. G. female flower, x 8. H & I. fruit, x 4. J. seed, x 6.

A & F from Jago 3068 (BRI); B-E, G-J from Forster PIF13740 (BRI). Del. W. Smith.

380

bridge over North Johnstone River, 17°32’S, 145°55’E,

Nov 1982, Jessup 509 (BRI, QRS); Russell River, 1892,

Johnson s.n. (MEL); North Johnstone River, Jun 1985,

Sankowsky 401 & Sankowsky (BRI); Harvey’s Creek,

Russell River, 1887, Sayer s.n. (MEL); Bellenden Ker,

Mar 1922, White 1291 (BRI); Russell River, 1886, Sayer

s.n. (MEL); Warramami Hill, Jun 1992, Tucker

[AQ625430] (BRI, DNA, L, MEL, QRS).

Distribution and habitat : Croton densivestitus

is restricted to the ‘Wet Tropics’ of north¬

eastern Queensland (Map 5) where it has been

found in two 1° grid squares. Plants generally

grow along creeks in lowland notophyll to

mesophyll vineforest on alluvium, although

there is one disjunct occurrence at 800 m in

vineforest on a seasonal watercourse.

Phenology: Flowers have been collected from

June to January and fruits in March and July,

but it is probable that flowering and fruiting

can occur sporadically throughout the year.

Notes: Croton pubens was included in the

synonymy of C. densivestitus by Airy Shaw

(1981). No material under this name was received

in a loan to BRI from PR, but this does not

preclude the existence of type material at that

institution. From Domin’s original description,

there seems little doubt as to the placement of

his name here.

Croton densivestitus is distinctive

amongst Australian taxa of Croton in the dense

yellow indumentum of the foliage.

Conservation status: This is a poorly collected

plant, although it may be reasonably common

in the extant localities. The species is present in

Wooroonooran National Park.

Etymology: The specific epithet is derived from

the Latin densus (dense) and vestitus (clothed)

and refers to the dense indumentum on the

foliage of this species.

12. Croton dockrillii Airy Shaw, Muelleria 4:

227 (1980). Type: Queensland. Cook

District: Alligator Creek, 12°35’S, 143°24’E,

14 October 1972, A. Dockrill 589 (holo:

QRS; iso: BRI).

Illustration: Airy Shaw (1981:619, fig. 2D).

Shrub to 3 m high, monoecious, evergreen,

perennial. Indumentum uncoloured. Branchlets

Austrobaileya 6 (3): 349-436

rounded, with dense stellate trichomes when

young, glabrescent. Stipules linear-lanceolate,

1.5-5 mm long, 0.2-0.8 mm wide, entire and with

dense stellate trichomes. Leaves alternate,

discolorous, petiolate; petioles 3-26 mm long,

c. 1.5 mm wide, with sparse stellate trichomes;

lamina elliptic, 30-100 mm long, 20-50 mm wide,

penninerved with 10 or 11 lateral veins per side

of midrib and reticulate tertiary veins; upper

surface dark glossy green, venation not visible,

glabrous; lower surface pale green, venation

weakly prominent, with scattered stellate

trichomes, neither scabrid nor velutinous;

margins sinuate to denticulate with 20-28 teeth

up to 0.2 mm long, foliar glands prominent; tip

acute, short to long acuminate; base cuneate;

extrafloral nectaries 2 at base of lamina, stipitate

to 1.5 mm, circular, 0.4—0.5 mm long, 0.4—0.5 mm

wide, visible above and below. Inflorescence

up to 50 mm long, androgynous, pedunculate

up to 8 mm; axis with sparse stellate trichomes;

bracts lanceolate, 0.7-1.5 mm long, 0.3-0.4 mm

wide, with sparse to dense stellate trichomes.

Male flowers 3-3.5 mm long, 3.5-4.5 mm

diameter, sparsely clustered or held singly

towards top of inflorescence; pedicels 2-3 mm

long, 0.3-0.4 mm wide, with sparse stellate

trichomes; sepals valvate, 5, lanceolate-ovate,

2-2.3 mm long, c. 1.6 mm wide, glabrous, weakly

lanate at tip; petals 5, oblanceolate, 2.5-2.8 mm

long, 0.7-0.8 mm wide, lanate in upper half;

stamens 9-11, filaments filiform, 2.2-2.5 mm

long, c. 0.2 mm wide, glabrous, anthers oblong,

0.5-0.6 mm long, 0.4-0.5 mm wide. Female

flowers c. 3 mm long, 3-5 mm diameter, held

singly and spaced up to 7 mm apart; pedicels

0.5-2 mm long, c. 1 mm diameter, with dense

stellate trichomes; sepals valvate, 5, lanceolate

to lanceolate-ovate, 2.6-3 mm long, 1-1.5 mm

wide, with sparse to dense stellate trichomes;

petals absent; styles 3, linear, 2.5-3 mm long,

bifid for 1.8-2.6 mm, with scattered stellate

trichomes in lower half; ovary 3-locular, 1.7-

2 mm long, 1.7-2 mm diameter, with dense, +

sessile stellate trichomes. Fruits trilobate,

globose, c. 5 mm long and 5 mm diameter, with

sparse, ± sessile stellate trichomes. Seeds ovoid,

c. 4 mm long, 4 mm wide, 3.5 mm thick, glossy

tan-brown, ventral surface bifacial, dorsal

surface rounded, micropylar ridge c. 4 mm long;

caruncle truncate-ovate, c. 1-1.2 mm long, 2.3-

2.5 mm wide, cream. Fig. 11.

Forster, Croton in Australia

381

Fig. 11. Croton dockrillii. A. flowering branchlet. x 0.6. B. undersurface of leaf, x 1. C. node with arrangement of

stipules, x 3. D & E. base of leaf lamina showing extrafloral nectaries, x 4. F. inflorescence with male flowers, x 2.

G. male flower, x 8. H. female flower, x 8. I. fruit on inflorescence, x 4. J. face view of fruit, x 4. K. seed, x 6. A,

I-J from Sankowsky 1444 (BRI); B & D from Sankowsky 1050 (BRI); C, E-H from Forster PIF13588 (BRI); K

from Fell DGF2138 (BRI). Del. W. Smith.

382

Additional specimens: Queensland. Cook District:

King Park, Claudie River, 12°36’S, 143°17’E, Jul 1990,

Fell DGF2136 (QRS); ditto, Fell DGF2137 (QRS);

ditto, Fell DGF2138 (BRI, QRS); 9 km SW of King

Park Ranger Station, Claudie River, 12°46’S, 143°17’E,

Apr 1992, Fell DGF2495 (BRI, MEE, QRS); Portland

Roads, Apr 1944, Flecker N.Q.N.C. 8507 (QRS); Rocky

River Scrub, eastern fall Mcllwraith Range, 13°49’S,

143°27’E, Jun 1992, Forster PIF10627 et al. (BRI,

DNA, K, L, MEL, NSW, QRS); Ham Hill, 12°43’S,

143°18’E, Jul 1993, Forster PIF13588 et al. (BRI,

MEL, QRS); 3 km SSW of Rocky River Crossing, Silver

Plains, 13°49’S, 143°27’E, Jul 1993, Forster PIF13665

et al. (BRI, CANB, MEL, QRS); Rocky River, 13°50’S,

143°25’E, Sep 1973, Hyland 6814 (BRI, CANB, QRS);

N.P.R. 8, Parish of Weymouth, 12°40’S, 143°21’E,

Jan 1982, Hyland 11588 (QRS); cult. Forestry &

Timber Bureau, Atherton (ex Claudie River), Jan 1975,

Irvine 1115 (BRI, QRS); cult. Tolga (ex Ham Hill,

12°43’S, 143°19’E), Dec 1989, Sankowsky 1050 (BRI,

CANB); Iron Range, Sep 1962, Volck 2418 (BRI);

Galloways Creek, Bamaga, May 1962, Webb & Tracey

6083 (BRI).

Distribution and habitat : Croton dockrillii is

endemic to Cape York Peninsula, Queensland

and has been found in the rainforest areas at

Cape York, Iron Range and the Mcllwraith Range

(Map 4) over a total of three 1° grid squares.

Plants grow in evergreen notophyll to mesophyll

vineforest on volcanic substrates or alluvium.

Croton dockrillii occurs in association with

C. capitis-york at some sites and occasional

hybrids have been recorded.

Phenology : Flowers have been collected

sporadically throughout the year. Fruiting

probably occurs two or three months later.

Notes: Croton dockrillii is distinctive in the

narrow leaves with markedly stipitate glands.

Conservation status: Croton dockrillii is

uncommon in its known range, but not

endangered or threatened at this stage.

Etymology: Named for Alick Dockrill of

Atherton, who collected the type specimen.

13. Croton glandulosus L., Syst. Nat. ed. 10,

1275 (1759). Decarinium glandulosum (L.)

Raf., Neogenyton 1 (1825).Type: [Jamaica]

P. Browne, jam. 346. n. 1 (lecto: LINN

1140.7 n.v. ; BRI fiche n.v. ;fide Johnston

(1959:182).

Illustrations: Ferguson (1901: plate 16); Small

(1913:454, fig. 2713).

Austrobaileya 6 (3): 349-436

Erect herb to 40 cm high, monoecious, annual.

Indumentum uncoloured. Stems rounded, with

sparse, sessile and stalked stellate trichomes.

Stipules linear-lanceolate, 0.8-1 mm long, c.

0.3 mm wide, entire and with sparse stellate

trichomes. Leaves alternate, discolorous,

petiolate; petioles 3-25 mm long, 0.7-1 mm wide,

with sparse, sessile and stalked stellate

trichomes; lamina elliptic to oblanceolate, 5-

22 mm long, 4-18 mm wide, palminerved with 5

veins at base and 2 or 3 lateral veins per side of

midrib further up the lamina, tertiary reticulate

veins obscure; upper surface dark green,

venation + obscure, with scattered stellate

trichomes; lower surface pale green, venation

weakly visible, with sparse, sessile and stalked

stellate trichomes; margins crenate, with 4-7

teeth up to 3.5 mm long, foliar glands

inconspicuous; tip acute to obtuse; base

cordate to truncate; extrafloral nectaries 2 at top

of petiole, sessile, ellipsoid, 0.5-0.8 mm long,

0.4-0.5 mm wide, visible above and below.

Inflorescence up to 10 mm long, androgynous,

+ sessile; axis with dense, sessile and stalked

stellate trichomes; bracts linear, 0.8-1 mm long,

0.1-0.2 mm wide, with dense stellate trichomes.

Male flowers c. 1.5 mm long and 1.5 mm diameter,

held singly, spaced < 0.5 mm apart; pedicels c.

0.3 mm long and 0.1 mm wide, with dense, sessile

and stalked stellate trichomes; sepals valvate,

5, lanceolate-ovate, 0.9-1 mm long and c. 0.5

mm wide, lanate at tip and with sparse, stalked

stellate trichomes; petals 5, obovate, c. 0.8 mm

long and 0.4 mm wide, lanate; stamens 9-12,

filaments flattened, c. 1 mm long and 0.1 mm

wide, glabrous, anthers oblong, 0.3-0.4 mm

long, 0.3-0.4 mm wide. Female flowers c. 4 mm

long and 3 mm diameter, held singly and densely

crowded at base of inflorescence, + sessile;

sepals valvate, 5, lanceolate-ovate to obovate,

1.5-4 mm long, 1-1.5 mm wide, with sparse to

dense, sessile and stalked stellate trichomes;

petals absent; styles 3, linear, 1-1.5 mm long,

bifid for c. 1 mm, with sparse sessile stellate

trichomes; ovary 3-locular, c. 1 mm long and 1

mm diameter, with dense stellate trichomes.

Fruits trilobate, globose, 4-5 mm long, 4-5 mm

diameter, with sparse, sessile and stalked stellate

trichomes. Seeds oblong, c. 3.5 mm long, 2.8mm

wide, 2 mm thick, grey, ventral surface bifacial,

dorsal surface rounded, micropylar ridge 2.5-

2.8 mm long; caruncle crescent shaped, c. 0.5

mm long and 1.2 mm wide, yellowish. Fig. 12.

Forster, Croton in Australia

383

Fig. 12. Croton glandulosus. A. flowering and fruiting branchlets. x 0.6. B. undersurface of leaf, x 2. C. base of leaf

lamina showing extrafloral nectaries, x 12. D. node showing stipule, x 8. E. node with inflorescence, x 4. F. male

flower, x 12. G. female flower, x 12. H & I. fruit enclosed by calyx, x 4. J. seed, x 8. All from Forster PIF28046.

Del. W. Smith.

384

Additional specimens examined : U.S.A. New Jersey:

Camden, Aug 1878, Martindale AQ206128 (BRI).

GEORGIA: College Park, Fulton Co., Apr 1964,

Schallert 842 (BRI). Australia, Queensland.

Moreton District: Jacobs Well road, Woongoolba, Dec

1994, Blatch 1813 (BRI); Jacobs Well, A.Brumm

property, 27°47’S, 153 0 21’E, Dec 2001, Forster

PIF28046 & Leiper (A, AD, BRI, DNA, K, L, MEL,

MO, NSW, WIS); Jacobs Well, Apr 2000, Leiper

[AQ667881] (BRI).

Distribution and habitat : Croton glandulosus

is native to the U.S.A. where it occurs in the

south-eastern and eastern states, through

Central America to Brasil (Ferguson 1901;

Johnston 1959, 1962). It has been recorded

several times from south-eastern Queensland

near Jacobs Well (Map 9) as a weed in sugar¬

cane paddocks. Between April 2000 and

December 2001, this species had experienced a

population explosion at the Jacobs Well locality,

with thousands of plants growing in dense

swards below a sugar-cane crop.

Phenology: The Australian collection had both

flowers and fruits in December.

Notes: Linnaeus (1759) lists “Croton

glandulofum Br. jam. 1.” in the protologue for

C. glandulosus. P. Browne donated his

Jamaican collections to Linnaeus (Stafleu &

Cowan 1981) and the Linnean herbarium (as

on the fiche at BRI) has the specimen “Brown,

jam. 346. n. 1” which I am equating with being

the lectotype of this name as informally

designated by Johnston (1959). Several

varieties have been recognised for this species

based mainly on the density of foliage

indumentum (Ferguson 1901; Johnston 1959,

1962).

Webster (1993a) included C. glandulosus

in Croton section Geiseleria (Klotzsch) Baill.

Etymology: The specific epithet probably refers

to the conspicuous glands on the leaf lamina.

14. Croton habrophyllus Airy Shaw, Kew Bull.

31: 386 (1976). Type: Northern Territory.

Port Darwin, June 1870, Schultz 680 (holo:

K n.v., photo at B RI!).

Illustrations: Dunlop etal. (1995: 214, Fig.

71); Wheeler (1992: 598, Fig. 182B);

Kenneally etal (1996:100-101).

Small tree or shrub to 4 m high, monoecious,

deciduous, perennial. Indumentum silver.

Austrobaileya 6 (3): 349-436

Branchlets ± rounded, with dense overlapping

stellate trichomes when young, sparse with age.

Stipules linear-lanceolate, 0.4-2.2 mm long, 0.1-

0.2 mm wide, entire and with scattered stellate

trichomes. Leaves alternate, discolorous,

petiolate; petioles 3-120 mm long, 0.6-2 mm

wide, with scattered to dense stellate trichomes;

lamina elliptic, elliptic-ovate or obovate, 15-200

mm long, 7-135 mm wide, with + penninerved or

somewhat palminerved, 9-11 lateral veins per

side of midrib and reticulate tertiary veins; upper

surface glossy green, lateral venation just

visible, with dense stellate trichomes becoming

scattered with age; lower surface pale green,

venation weakly prominent, with dense stellate

trichomes becoming scattered with age, neither

scabrid nor velutinous; margins sinuate to

denticulate with 35-56 teeth up to 0.5 mm long,

foliar glands conspicuous; tip acuminate, acute

or rounded; base cordate, rounded or truncate;

extrafloral nectaries 2 at lamina base, stipitate to

1 mm, ellipsoid, 0.4—1 mm long, 0.3-0.7 mm wide,

visible only below. Inflorescence up to 170 mm

long, androgynous or sometimes with glomerules

of mixed male and female flowers, pedunculate

up to 25 mm; axis with sparse to dense stellate

trichomes; bracts linear-lanceolate to lanceolate,

0.7-1.2 mm long, 0.2-0.3 mm wide, with scattered

stellate trichomes. Male flowers 2.5^4 mm long,

3-5 mm diameter, held singly or sparsely

clustered towards top of inflorescence; pedicels

1.5-6 mm long, 0.3-0.5 mm wide, glabrous or

with scattered stellate trichomes; sepals valvate,

5, lanceolate-ovate, 1.8-2.5 mm long, 0.8-1.8 mm

wide, glabrous, weakly lanate at tip; petals 5,

oblanceolate, 2-3 mm long, 0.5-1 mm wide, lanate

at top; stamens 8-12, filaments filiform, 2-2.5

mm long, 0.1-0.2 mm wide, glabrous, anthers

oblong, 0.6-0.8 mm long, 0.5-0.7 mm wide.

Female flowers 2-4 mm long, 2.2-3 mm diameter,

held singly and spaced up to 13 mm apart, or

mixed with male flowers; pedicels 1-4 mm long,

0.5-1 mm diameter, glabrous or with sparse

stellate trichomes; sepals valvate, 5, lanceolate-

ovate, 1.8-2.3 mm long, 1-1.3 mm wide, with

scattered stellate trichomes and lanate tip; petals

absent; styles 3, obloid-flabellate to linear, 1-

2.8 mm long, bifid for 0.5-1.8 mm, connate at

base for c. 0.2 mm, glabrous; ovary 3-locular,

1.3-2 mm long, 1.3-2 mm diameter, with dense,

sessile stellate trichomes. Fruits trilobate,

depressed-globose, 5-6 mm long, 6-7 mm

diameter, with sparse, sessile stellate trichomes.

Forster, Croton in Australia

385

Fig. 13. Croton habrophyllus. A. flowering branchlet. x 0.6. B. undersurface of leaf, x 0.8. C. base of leaf lamina

showing extrafloral nectaries, x 8. D. node showing stipule, x 8. E. male flower, x 8. F. female flower, x 8. G & H

fruit, x 4. I. seed, x 8. A, D, F from O’Keefe 5 (BRI); B, C from Lucas 4635 (BRI); E from Innis 205 (BRI); G-I from

O’Keefe AQ454826 (BRI). Del. W. Smith.

386

Seeds ± obloid, 3.5-4 mm long, 2.6-3.5 mm wide,

2.2-2.5 mm thick, mottled cream and tan-brown,

ventral surface bifacial, dorsal surface rounded,

micropylar ridge 2-2.8 mm long; caruncle

reniform, 0.5-1 mm long, 1-2 mm wide, cream-

yellow. Fig. 13.

Selected additional specimens: Western Australia.

One Arm Point, N. Dampierland, 16°26’S, 123°05’E,

Nov 1987, Carter 139 (DNA); Cape Leveque,

Gnamagun Well, 16°27’S, 122°55’E, Apr 1988, Dunlop

7811 (BRI, DNA, MEL); Galen, Dampierland, W of

Skeleton Point, 16°32’S, 122°58’E, Jan 1988, Martin

207 (BRI, CANB, PERTH). Northern Territory.

1 km past Adelaide River on Daly River road, 13°30’S,

131°05’E, Nov 1990, Cowie 1405 & Dunlop (DNA);

Melville Island, Snake Bay, Sand Spear Jungle, Nov

1983, Dunlop 6510 & Wightman (BRI, DNA, MEL);

East Alligator River, 12°50’S, 133°22’E, Dec 1989,

Dunlop 7628 (BRI, CANB, DNA, MEL); Middle Arm,

near Palmerston, 12°34’S, 130°34’E, Nov 1989,

Forster PIF5942 et al. (BRI, DNA, MEL); 3 km W of

Nhulunbuy, 12°11’S, 136°45’E, Nov 1989, Forster

PIF6048 (BRI, DNA); Wessell Islands, lTll’S,

136°44’E, Sep 1972, Latz 3289 (BRI, DNA); North

Point, Kapalga, 12°23’S, 132°21’E, Nov 1983, Russell-

Smith 885 (BRI, DNA); Bennett Bay, Eastern Arnhem

Land, 13°42’S, 135°42’E, Nov 1987, Russell-Smith

4222 & Lucas (BRI, DNA); Mt Ranken, Walker River,

13°35’S, 135°32’E, Oct 1987, Russell-Smith 4322 &

Lucas (BRI, DNA); Cape Wirawawo, Gove, 12°10’S,

136°47’E, Feb 1988, Russell-Smith 4635 & Lucas (BRI,

DNA); Nhulunbuy, East Woody Island, 12°10’S,

136°45’E, Feb 1988, Wightman 4128 (BRI, DNA,

MEL); Glasswater Creek, Litchfield, 13°18’S,

130°30’E, Oct 1988, Russell-Smith 5977 & Lucas

(BRI, DNA); High Black Range, Moroak, 14°39’S,

133°37’E, Jan 1989, Russell-Smith 6614 & Lucas (BRI,

DNA); Redbank Mine, Wollogorang Station, 17°10’S,

137°46’E, Nov 1984, Thomson 763 (BRI, DNA); SW

corner Centre Island, Sir Edward Pellew Group, 15°06’S,

136°44’E, Jan 1989, Thomson 2946 (BRI, DNA).

Queensland. Burke District: Louie Creek, 18°48’S,

138°32’E, Dec 1989, Innis 205 (BRI); Lawn Hill N.P.,

18°45’S, 138°30’E, Dec 1989, O’Keefe 10 (BRI).

Distribution and habitat : Croton

habrophyllus is widespread (twenty-four 1° grid

squares) in northern Australia, particularly in

the “Top End” of the Northern Territory. There

are disjunct populations in the Kimberley region

of Western Australia and at Lawn Hill National

Park in Burke district in Queensland (Map 5).

Plants grow in deciduous vinethickets on a

variety of substrates, including granite, laterite,

sandstone and limestone.

Phenology: Flowering occurs from August to

January, especially after storm rains. The plant

becomes deciduous for a short period and

Austrobaileya 6 (3): 349-436

flowers as the new leaves are expanding.

Fruiting occurs from October to April, by which

time the leaves are fully expanded.

Notes: This species was placed in the synonymy

of Croton armstrongii by Airy Shaw (1980c,

1981) and later reinstated by Wilmot-Dear

(1987). It is easily distinguished from related

species such as Croton byrnesii , C. mutabilis

and C. rarus in the silver indumentum and

leaves with more marginal teeth (70-112).

Conservation status : Croton habrophyllus is

common and widespread.

Etymology: The specific epithet is derived from

the Greek habros (shaggy) and phyllus (leaf)

and refers to the dense indumentum on the

young foliage.

15. Croton insularis Baill., Adansonia 2: 217

(1862), (‘insulare’); Oxydectes insularis

(Baill.) Kuntze, Rev. Gen. PL 2:612 (1891).

Type: New Caledonia, Pancher 360 (lecto:

P ,fide McPherson & Tirel 1987).

Illustrations: Willi ams (1979:78); McPherson

& Tirel (1987, pi. 15: 1-6); Floyd (1989:

142); James & Harden (1990:420); Hauser

(1992: 88); Logan River Branch S.G.A.P.

(Qld Region) Inc. (2002:221).

Tree or shrub to 15 m high, monoecious,

evergreen, perennial; bark lenticellate, cream;

blaze pale pink-red; wood straw. Indumentum

silver. Branchlets rounded, with dense

overlapping peltate scales. Stipules apparently

obsolete. Leaves alternate, discolorous,

petiolate; petioles 8-18 mm long, c. 1 mm wide,

with sparse peltate scales; lamina elliptic,

lanceolate-ovate to ovate, 12-100 mm long, 8-

45 mm wide, venation obscure; upper surface

dark matt green, with sparse peltate scales;

lower surface silver, with dense, overlapping

peltate scales, neither scabrid nor velutinous;

margins entire or somewhat sinuate, foliar

glands inconspicuous; tip acute to acuminate;

base cuneate to obtuse; extrafloral nectaries 2

at top of petiole, sessile, circular, 0.2-0.5 mm

long, 0.2-0.5 mm wide, visible above only.

Inflorescence up to 120 mm long, unisexual or

androgynous, pedunculate up to 20 mm; axis

with dense, overlapping peltate scales; bracts

lanceolate-triangular, 0.6-1 mm long, 0.3-1 mm

Forster, Croton in Australia

wide, with dense peltate scales. Male flowers

4- 5 mm long, 3-4 mm diameter, held singly or

densely clustered towards top inflorescence,

spaced up to 3 mm apart; pedicels 2.3-5 mm

long, 0.5-1 mm wide, with dense peltate scales;

sepals valvate, 4 or 5, lanceolate-triangular, 1.5-

2.5 mm long, 1.2-1.6 mm wide, with sparse

peltate scales; petals 5, oblanceolate, 1.7-2.6

mm long, 0.6-1 mm wide, lanate; stamens 14-

18, filaments filif orm, 1.8-4 mm long, c. 0.1 mm

wide, glabrous, anthers oblong, 0.8-1.1 mm

long, 0.6-1 mm wide. Female flowers 2.5-3 mm

long, 3.5-5 mm diameter, held singly and spaced

up to 5 mm apart; pedicels 3-8 mm long, 0.6-1

mm diameter, with dense peltate scales; sepals

valvate, 5, lanceolate, 1.8-2.6 mm long, 1-1.6

mm wide, with dense peltate scales; petals

absent; styles 3, obloid, 0.8-2 mm long, bifid

for 0.5-1.2 mm, connate at base for c. 0.2 mm,

glabrous; ovary 3-locular, 1.5-2 mm long, 1.5-

2.2 mm diameter, with dense, sessile peltate

scales. Fruits trilobate, globose, 7-9 mm long,

5- 8 mm diameter, with dense, sessile peltate

scales. Seeds ovoid, 3.6-6.5 mm long, 2-3.2

mm wide, 2-2.5 mm thick, brown, ventral surface

bifacial, dorsal surface rounded, micropylar

ridge 2.3-5.7 mm long; caruncle crescent

shaped, 0.5-0.8 mm long, 0.7-1.5 mm wide,

cream-yellow. Fig. 14.

Selected additional specimens’. Queensland. Cook

District: S.F. 185 Danbulla, 7 km SW of Hoop Pine

Triangle, 17°09’S, 145°35’E, Jan 1993, Forster

PIF13082 & Bean (BRI, L, MEL, QRS); Possum Scrub,

Weipa to Stones Crossing road, 12°27’S, 142°09’E, Jul

1993, Forster PIF13512 et al. (BRI, MEL); Mt

Windsor Tableland, S.F 144, 9 km past Spencer Creek

Crossing, 16°18’S, 145°05’E, Jul 1993, Forster

PIF13703 et al. (BRI, MEL, QRS). North Kennedy

District: Fern Creek Spring, catchment of Burdekin

River, St Pauls Scrub, Mt Cooper Station, 42 km S of

Ravenswood, Aug 1989, Fell DGF1964 (BRI). South

Kennedy District: 26.5 km W of St Anns Homestead,

21°13’S, 146°39’E, Jun 1992, Thompson BUC582 &

Sharpe (BRI). Leichhardt District: Coxens Peak,

22°12’S, 148°27’E, Aug 1990, Forster PIF7310 (BRI,

MEL, QRS); 17 km from Cracow on Nathan Gorge

road, 25°26’S, 150°19’E, Sep 1992, Forster PIF 11207

& Sharpe (BRI, L, MEL, QRS); Palmgrove N.P., Bigge

Range, 25°01’S, 149°16’E, Nov 1998, Forster

PIF23653 & Booth (BRI, MEL, QRS). Maranoa

District: Chesterton Range, Mt Moffat, NW of Marlong

Plain & SW of Mt Sugarloaf, Nov 1990, Henderson

3504 & Robins (BRI, NSW). Port Curtis District: c.

17 km ESE of Duaringa, 23°46’S, 149°50’E, Sep 1988,

Anderson 4520 (BRI); Barren Island, SE of Great

Keppel Is, 23°10’S, 151°05’E, Batianoff 9691 &

387

Dillewaard (BRI, NSW). Burnett District: Mt

Wooroolin, 26°32’S, 151°48’E, Apr 1990, Forster

PIF6662 (BRI, L, MEL, QRS); Coominglah Range,

S.F.28 Coominglah, 24°5LS, 150°56’E, Nov 1994,

Forster PIF 15908 (BRI, MEL, QRS). WideBay District:

5 km SW of Mt Walsh, Coongara Rock road, 25°36’S,

152°00’E, Oct 1990, Forster PIF7549 (BRI, K, L,

MEL, QRS); Fairlies Knob, 10 km NNE of Brooweena,

25°30’S, 152°17’E, Dec 1990, Forster PIF7672 (BRI,

MEL, QRS). Darling Downs District: Chinchilla, May

1912, Beasley 4 (BRI). Moreton District: end of

Steinharts road, Lark Hill, 4 km N of Marburg, 27°32’S,

152°36’E, May 1983, Forster PIF1586 (BRI); Splityard

Creek, Wivenhoe Dam, 27°23’S, 152°38’E, Nov 1990,

Forster PIF7612 & Sharpe (BRI, DNA, K, L, MEL,

MO, QRS); Welk Remnant, Mt Berryman, 27°43’S,

152°18’E, Sep 1999, Forster PIF24931 (BRI, QRS).

New South Wales. 36 miles [60 km] W of Wauchope

on Oxley Highway, Aug 1967, Telford 58 (CANB).

Distribution and habitat: Croton insularis is

found in Australia in eastern Queensland from

near Weipa in the north, to north-eastern New

South Wales in the south (Map 6) over a total

of forty-four 1° grid squares. As a result it is the

third most widespread Croton, after

C. amhemicus and C. phebalioides, in mainland

Australia. It also occurs in New Caledonia and

Vanuatu (McPherson & Tirel 1987). Plants grow

in vinethickets or vineforests on a variety of

volcanic substrates.

Phenology: Flowering and fruiting occurs

througout the year following rain. The peak

flowering period is from October to December.

Notes: Croton insularis is easily distinguished

from all other taxa of Australian Croton by the

foliage silver below with totally obscure lateral

and interlateral venation. Usually this species

is encountered as a shrub up to 5 m tall but at

some localities (e.g. Cathu, Windsor Tableland

and Bridle Creek State Forests) it may grow up

to 15 m tall as a canopy tree.

There is a variant population at Eight Mile

Mountain, Emu Creek Station, 9 km NNE of

Petford C Ford 3668 (BRI); Forster PIF28192 et

al. (BRI, MEL, NSW, WIS)) that appears to be

intermediate in vegetative morphology between

C. insularis and C. phebalioides.

Conservation status: Croton insularis is very

common. It is present in 23 conservation

reserves in south-eastern Queensland alone

(Forster et al. 1991) and four in New South Wales

(Floyd 1989).

388

Austrobaileya 6 (3): 349-436

Fig. 14. Croton insularis. A. flowering branchlet. x 0.8. B & C undersurface of leaf, x 0.5. D. base of leaf lamina

showing extrafloral nectaries, x 8. E. inflorescence with female flowers in lower half and male flowers in upper

half, x 2. F. male flower, x 8. G. female flower, x 8. H & I. fruit, x 4. J. seed, x 8. A, B & G from Forster PIF25193

(BRI); C from Forster PIF6662 (BRI); D-F from Forster PIF7549 (BRI); H-J from Ryan 1416 (BRI). Del.

W. Smith.

Forster, Croton in Australia

Etymology : The specific epithet refers to the

island origin of the type collection.

16. Croton magneticus Airy Shaw, Muelleria 4:

227 (1980). Type: Queensland. North

Kennedy District: Magnetic Island, 24

July 1938, DA. Goy 329 (holo: BRI).

Small tree or shrub to 5 m high, monoecious,

deciduous, perennial. Indumentum ginger to

silver. Branchlets ± rounded, with dense stellate

trichomes when young, glabrescent. Stipules

subulate, 0.3-0.9 mm long, c. 0.2 mm wide, entire

and with sparse to dense stellate trichomes.

Leaves alternate, petiolate, discolorous; petioles

5-25 mm long, c. 1 mm wide, with dense stellate

trichomes; lamina cuneate-obovate, elliptic,

elliptic-ovate, 20-115 mm long, 12-60 mm wide,

penninerved with 7-9 lateral veins per side of

midrib, tertiary reticulate veins obscure; upper

surface matt green, venation obscure, glabrous

or with scattered stellate trichomes; lower

surface silver, lateral veins weakly developed,

with sparse to dense, stellate trichomes, scabrid

to weakly velutinous; margins denticulate to

weakly crenate with 8-24 short teeth up to 2

mm long, foliar glands prominent; tip obtuse to

rounded; base weakly cordate, cuneate or

truncate; extrafloral nectaries 2 at lamina base,

sessile, ellipsoid, 0.6-0.7 mm long, 0.3-0.4 mm

wide, visible only below. Inflorescence up to 80

mm long but often reduced to a single flower,

often unisexual but occasionally bisexual and

androgynous, pedunculate up to 10 mm; axis

with dense stellate trichomes; bracts lanceolate

to oblanceolate, 1.8-4 mm long, 0.4-1.5 mm

wide, with dense stellate trichomes. Male

flowers 3-5 mm long, 4.5-5 mm diameter, held

singly on inflorescence, spaced up to 2 mm

apart; pedicels 2.5-7 mm long, 0.2-1 mm wide,

with sparse to dense stellate trichomes; sepals

valvate, 5, lanceolate-ovate to obovate, 3-3.5

mm long, 1.8-2.2 mm wide, with dense stellate

trichomes; petals 5, obovate, 3-4 mm long, c.

1.5 mm wide, lanate; stamens 15, filaments

filiform, 2.2-3 mm long, 0.2-0.5 mm wide,

glabrous; anthers oblong, 0.8-1.2 mm long, 1-

1.3 mm wide. Female flowers 4-4.5 mm long,

3.5-5 mm diameter, held singly and spaced up

to 2 mm apart; pedicels 3-8 mm long, 1-1.2 mm

diameter, with dense stellate trichomes; sepals

valvate, 5, lanceolate to lanceolate-ovate, 2.3-3

mm long, 1.2-1.8 mm wide, with dense stellate

389

trichomes; petals absent; styles 3, obloid, 1.8-

3 mm long, bifid for 1.6-2.8 mm long, connate at

base for 0.2 mm, glabrous; ovary 3-locular, 3-4

mm long, 3-4 mm diameter, with dense, stalked

stellate trichomes. Fruits trilobate, globose, c. 8

mm long and 8 mm diameter, with dense, stalked

stellate trichomes. Seeds obloid-ovoid, 5-5.5

mm long, 4.2^1.5 mm wide, c. 3 mm thick, pale

brown, ventral surface bifacial, dorsal surface

rounded, micropylar line 4-4.5 mm long; caruncle

crescent shaped, 1.5-1.7 mm long, 0.7-1 mm

wide, cream. Fig. 15.

Additional specimens : Queensland. North Kf.nnf.dy

District: Georges Point, 20°04’S, 148°35’E, Sep 1992,

Batianoff 9209286 & Carter (BRI, MEL); Montes

Resort, Cape Gloucester, 20°04’S, 148°27’E, Mar 1994,

Batianoff 9403250 & Dillewaard (BRI); Gloucester

Island, E. side, 19°59’S, 148°27’E, Apr 1994, Batianoff

94037 & Figg (BRI); Gloucester Island, S. end, 20°02’S,

148°26’E, Apr 1994, Batianoff 940415G & Figg (BRI);

Gloucester Island, E. side, 20°01’S, 148°28’E, Apr 1994,

Batianoff 9 A0 A A3 & Figg (BRI); Mt Abbot, 50 km W

of Bowen, 20°06’S, 147°43’E, Jul 1992, Bean 4734

(BRI); Mt Blackjack, ‘Wietalaba’, 21°01’S, 147°56’E,

Jan 1993, Fensham 490 (BRI); ‘Havilah’, 20°58’S,

147°52’E, Dec 1992, Fensham 601 (BRI); ‘Fanning

River’, 19°44’S, 146°27’E, Jan 1993, Fensham 731

(BRI); Turtle Creek, SW of Greenvale, 19°18’S,

144°50’E, Fensham 914 (BRI); Leichhardt Range,

20°03’S, 147°03’E, Jul 1993, Fensham 997 (BRI); West

Point, Magnetic Island, 19°07’S, 146°46’E, Jan 1993,

Forster PIF12761 & Bean (BRI, MEL, QRS);

‘Wietalaba’, 21°01’S, 147°56’E, Jul 1993, Forster

PIF13408 & Tucker (BRI, QRS); ditto, Feb 1994,

Forster PIF14863 & Bean (A, BISH, BRI, CANB, DNA,

K, L, MEL, NSW, QRS); Mt Blackjack, Wietalaba

Station, 21°00’S, 147°55’E, Jun 1996, Forster

PIF19197 & Tucker (BRI, QRS); Magnetic Island, Jun

1922, Helms 1125 (BRI); Balding Bay, Magnetic Island,

Aug 1982, Sandercoe 751 (BRI); Magnetic Island, Aug

1982, Sandercoe 860 (BRI); Magnetic Island, Jun 1983,

Tracey 14101 (BRI).

Distribution and habitat : Croton magneticus

is restricted to an area between Greenvale in

the north to near Collinsville in the south in

north-eastern Queensland (Map 5) over seven

1° grid squares. Plants grow in deciduous

vinethicket on soils derived from sandstone,

granite or acid agglomerate, often in association

with Croton arnhemicus and C. phebalioides.

Phenology: Flowering occurs from December

to February following storm or seasonal rains,

fruiting occurs from January to March. Dormant

buds are held on the plants for much of the

year.

390

Austrobaileya 6 (3): 349-436

Fig. 15. Croton magneticus. A. flowering branchlet. x 1. B & C. undersurface of leaves, x 0.5. D. base of leaf lamina

showing extrafloral nectaries, x 6. E. node showing stipules, x 4. F. inflorescence with female flowers towards base

and male flowers towards apex, x 2. G. female flower, x 6. H. male flower, x 6. I & J. dehiscing fruit, x 3. K. seed,

x 4. A, C, F-H from Forster PIF14863 (BRI); B from Forster PIF12761 (BRI); I-K from Fensham 490 (BRI). Del.

W. Smith.

Forster, Croton in Australia

Notes: Croton magneticus is often sympatric

with C. amhemicus and sterile collections could

possibly be confused with small-leaved forms

of that species. Croton magneticus is easily

distinguished by its penninerved leaves, as

opposed to the palminerved leaves of

C. amhemicus.

Conservation status : Croton magneticus is

now known to be much more widespread than

was previously thought (Airy Shaw 1981). It is

relatively common at the listed localities, but

these are disjunct and several may be subject

to land clearing. The species is present in the

National Park at Magnetic Island and is

currently listed as Vulnerable under Queensland

Government legislation.

Etymology: The specific epithet refers to the

plant’s occurrence on Magnetic Island, where

it was once thought to be endemic.

17. Croton mamillatus P.I.Forst., sp. nov. affinis

C. insulari autem venatione foliorum ex

12-14 venis lateralibus constanti (vice

venatione obscura), pedicellis trichomatis

peltatis (vice squamis peltatis) vestitis,

floribus masculinis staminibus 9 vel 10

(vice 14-18), fructibus processis

mamillatis praeditis trichomatis stellatis

vestitis (vice processis emamillatis

squamis sessilibus peltatis vestitis) differt.

Typus: Queensland. Moreton District:

Bahr’s Scrub, 5 km SSW of Beenleigh,

27°45’S, 153 °10’E, 19 December 2001,

P.l. Forster PIF28049 & GLeiper (holo: BRI

[2 sheets + spirit]; iso: A, L, MEL, NE,

NSW).

Shrub to 4 m high, monoecious, evergreen,

perennial. Indumentum uncoloured to silver.

Branchlets + rounded, with dense peltate

trichomes, glabrescent. Stipules shortly

lanceolate, c. 0.5 mm long and 0.3 mm wide, entire

and with dense peltate trichomes. Leaves

alternate, petiolate, discolorous; petioles 3-10

mm long, 1.2-1.5 mm wide, with dense peltate

trichomes; lamina elliptic to oblanceolate, 30-

70 mm long, 10-32 mm wide, venation

penninerved with 12-14 lateral veins per side

of midrib, very indistinct, tertiary reticulate veins

obscure; upper surface matt dark-green, more

glossy when fresh, venation weakly visible,

glabrous; lower surface silver-white, lateral

veins indistinct, with dense peltate trichomes

391

and peltate scales, neither scabrid nor

velutinous; margins ± entire, or very weakly

denticulate with barely discernible foliar glands;

tip acute to acuminate; base rounded to retuse;

extrafloral nectaries absent or if present, then 2,

circular, ± sessile, c. 0.3 mm long and 0.2 mm

wide, visible above and below. Inflorescence

up to 35 mm long, unbranched, androgynous

(rarely with male and female flowers mixed in

same glomerule), pedunculate up to 12 mm; axis

with dense peltate trichomes; bracts shortly

lanceolate, 0.5-0.7 mm long, c. 0.3 mm wide, with

dense peltate trichomes. Male flowers c. 2 mm

long and 3 mm diameter, held singly, spaced up

to 1 mm apart, usually towards top of

inflorescence; pedicels 2.5-3 mm long, 0.4-0.5

mm wide, with dense peltate trichomes; sepals

valvate, 5, lanceolate-ovate to ovate, 1.8-2.2 mm

long, 1.4-1.5 mm wide, with dense peltate

trichomes; petals 5, obovate, 1.5-2 mm long,

0.6-0.7 mm wide, lanate in upper half; stamens

9 or 10, filaments filiform, 2-2.2 mm long, c. 0.1

mm wide, with dense simple trichomes at base,

anthers oblong, 0.6-0.8 mm long, 0.3-0.4 mm

wide. Female flowers 3-3.2 mm long, 2.8-3.5 mm

diameter, usually held singly and spaced up to

3 mm apart; pedicels 3-4 mm long, 0.8-1 mm

diameter, with dense peltate trichomes; sepals

valvate, 5, ovate to obovate, 2.5-3 mm long,

1.8-2 mm wide, with dense peltate trichomes,

lanate; petals absent; styles 3, linear, 1.2-1.5

mm long, multifid, twice divided for 1-1.2 mm

long, connate at base for c. 0.2 mm, glabrous;

ovary 3-locular, 1.8-2 mm long, 2.5-2.7 mm

diameter, with dense stalked stellate trichomes.

Fruits trilobate, weakly depressed-globose, 9-

10 mm long, 10-10.5 mm diameter, with dense

stellate trichomes on fleshy mamillate

protuberances to 1 mm long and 0.5 mm diameter

that are topped by a stellate trichome. Seeds

oblong, 6-7 mm long, c. 3.5 mm wide, 2.5-3 mm

thick, grey-brown, ventral surface bifacial, dorsal

surface rounded, micropylar ridge 4.5-5 mm

long; caruncle oblong-rectangular, 1.4-1.5 mm

long, 0.8-1 mm wide, cream. Fig. 16.

Additional specimens : Queensland. Moreton District:

Bahr’s Scrub, 5 km SW of Beenleigh, 27°45’S, 153°09’E,

Feb 2001, Bean 17373 (BRI, MEL); slopes of Mt

French, SW of Boonah, 28°00’S, 152°37’E, Jan 2002,

Bean 18336 (BRI); Wolfdene area, 6 km SW of

Beenleigh, 27°46’S, 153°10’E, Sep 2002, Forster

PIF28882 et al. (A, BRI, L, MEL, NE, NSW, NY);

Bahr’s Scrub, Beenleigh, 27°45’S 153°10’E, Jan 2000,

Leiper [AQ667034] (BRI, QRS); French’s Creek road,

near Boonah, Jun 1984, Williams 84050 (BRI).

392

Distribution and habitat: This new species is

known from only four localities, two near

Beenleigh and two near Boonah, all southwest

of Brisbane in south-eastern Queensland (Map

5) on two 1° grid squares. All populations occur

in the understorey of dry rainforest (araucarian

microphyll vineforest or notophyll vineforest)

amongst rocks, on red soil derived from chert.

Notes: Recognition of this taxon as a new and

critically endangered species, is very much due

to the keen eye of Glen Leiper who first brought

the Bahr’s Scrub population to my attention.

Croton mamillatus appears to have affinities

with both C. insularis and C. stigmatosus, but

is immediately distinctive in its spindly habit

and the highly mamillate fruit, with the fleshy

processes topped by a stellate hair. This feature

of mamillate fruit is also present in Croton

capitis-york and C. stigmatosus but is less well

developed in those species.

Conservation status: Croton mamillatus is

known from only four localities, all on private

land. The populations at the four localities

comprise less than 100 individuals in total. It is

likely that the species was once more

widespread in the region but that other

populations have been destroyed in land

clearing over the last 150 years. Plants of Croton

mamillatus are also rather insignificant, and

superficially similar to species such as

C. insularis or C. stigmatosus, so may well have

been overlooked by collectors. On present

evidence the species should be regarded as

critically endangered using the IUCN (2001)

categories of A. 1(c), B. 2(a, b (ii, iii), C. 2a(i).

Etymology: The specific epithet mamillatus is

from the Latin word mamillatus, meaning

mamillate or having small nipple-like projections,

and refers directly to the distinctive fruit of this

species.

18. Croton minimus P.I.Forst., sp. nov. affinis

C. arnhemico Muell.Arg. a qua habitu

suffrutice usque 30 cm alto, foliis dentibus

paucioribus (22-28) et venatione

secundaria obscura, et staminibus florum

marium paucioribus (16-18) differt. Typus:

Queensland. Cook District: 11 km from

Petford towards Dimbulah, 2.5 km E of

Eight Mile Mountain, 17°15’S, 144°58’E,

30 January 1994, P.I. Forster PIF14708

Austrobaileya 6 (3): 349-436

(holo: BRI [1 sheet + spirit]; iso: MEL,

NSW).

Croton sp. (Mt Mulligan H.Flecker

NQNC6457) (Forster & Henderson 1997:

72; Forster & Halford 2002:70).

Multistemmed subshrub to 30 cm high,

monoecious, evergreen, perennial. Indumentum

ferruginous-silver. Branchlets rounded, with

dense stellate trichomes when young,

glabrescent. Stipules linear-lanceolate, 2-5 mm

long, 0.4-0.5 mm wide, entire and with dense

stellate trichomes. Leaves alternate, discolorous,

petiolate; petioles 3-8 mm long, 1-1.2 mm wide,

with dense stellate trichomes; lamina broadly

ovate to lanceolate-ovate, 10- 40 mm long, 8-

25 mm wide, palminerved with 3-5 veins from

the base and 3 or 4 lateral veins per side of

midrib further up lamina, tertiary reticulate veins

obscure; upper surface green-grey, venation

obscure, with sparse stellate trichomes; lower

surface ferruginous-silver, lateral veins weakly

prominent, with dense, stellate trichomes,

velutinous; margins irregularly crenate with 11-

14 teeth up to 1.5 mm long, foliar glands

prominent; tip acute; base cordate to rounded;

extrafloral nectaries absent at base of leaf lamina.

Inflorescence up to 20 mm long, androgynous,

pedunculate up to 6 mm; axis with dense stellate

trichomes; bracts linear-lanceolate, 0.8-2 mm

long, 0.2-0.3 mm wide, with sparse stellate

trichomes. Male flowers 2.5-6 mm long, 2.5-6

mm diameter, densely clustered on

inflorescences in glomerules of 1-3 flowers;

pedicels 1.4-2.5 mm long, c. 0.5 mm wide, with

dense stellate trichomes; sepals valvate, 5,

lanceolate-ovate to ovate, 1.5-2.5 mm long, 0.9-

2 mm wide, with dense stellate trichomes; petals

5, oblanceolate, 2-2.2 mm long, 0.5-1 mm wide,

lanate in upper half; stamens 16-18, filaments

filiform, 1-3.5 mm long, c. 0.1 mm wide, glabrous;

anthers oblong, 0.7-0.8 mm long, 0.4-0.6 mm

wide. Female flowers 2.5-3 mm long, 2-2.5 mm

diameter, held singly and spaced up to 3 mm

apart; pedicels 0.5-0.8 mm long, 0.5-0.6 mm

diameter, with dense stellate trichomes; sepals

valvate, 5, lanceolate-ovate, 1.8-3 mm long, 1-

1.8 mm wide, with dense stellate trichomes;

petals absent; styles 3, linear 1.8-2 mm long,

bifid for 0.8-1 mm, with scattered stellate

trichomes in lower half; ovary 3-locular, 1.8-2.6

mm long, 1.5-2.6 mm diameter, with dense,

Forster, Croton in Australia

393

Fig. 16. Croton mamillatus. A. flowering branchlet. x 0.6. B. undersurface of leaf, x 2. C. base of leaf lamina

showing extrafloral nectaries, x 12. D. node showing stipule, x 8. E. node with inflorescence with female flower

near base and male flowers near apex, x 1.5. F. male flower, x 8. G. female flower, x 8. H & I fruit, x 3. J. mamillate

process on fruit with stellate hairs, x 20. K. seed, x 6. All from Forster PIF28049 (BRI). Del. W. Smith.

394

sessile and stalked stellate trichomes. Fruits

trilobate, globose, 6-6.5 mm long, 5-6.5 mm

diameter, with dense, sessile and stalked stellate

trichomes. Seeds ± ovoid to obloid, 4-4.5 mm

long, c. 4 mm wide, 3-3.2 mm thick; dorsal surface

rounded, ventral surface bifacial; micropylar line

3.8-4 mm long; caruncle + reniform, 0.8-1 mm

long, 1.2-1.5 mm wide, cream. Fig. 17.

Additional specimens : Queensland. Cook District:

bank of Hodgkinson River, Mineham’s, Flecker

N.Q.N.C.1165 (BRI); between Mt Mulligan &

Thornborough, Dec 1937, Flecker N.Q.N.C.6459

(BRI); Chillagoe - Mungana road, c. 200 m SE of Red

Dome turnoff, 17°07’S, 144°26’E, Nov 2000, Ford

AF2484 & Tucker (BRI, CANB, K, L, MEL, NE, NSW);

N.RR. 98, near Belgravia Creek, off Burke

Development road, Mungana, 17°06’S, 144°24’E, Apr

2002, Ford AF3330 et al. (BRI, DNA, MEL, NSW);

between Eureka Creek & Mt Pinnacle, NNE of Summit,

SW of Dimbulah, 17°13’S, 145°03’E, Apr 2002, Ford

AF3338 & Sankowsky (BRI); Mt Pinnacle, SSW of

Dimbulah, 17°14’S, 145°03’E, Jan 1993, Forster

PIF12950 & Bean (BRI, MEL); Near Spring Creek

Crossing, Mt Carbine to Lakeland Downs road, 16°27’S,

144°50’E, Forster PIF18128 & Spokes (BRI, MEL);

Mungana, near Red Dome Mine turnoff, 17°06’S,

144°25’E, Jan 2002, Forster PIF28145 et al (BRI);

Eight Mile Mountain, 10.5 km NE of Petford, 17°15’S,

144°59’E, Jan 2002, Forster PIF28146 et al. (BRI,

MEL, NSW, WIS); Eight Mile Mountain, Emu Creek

Station, 9.5 km NNE of Petford, 17°15’S, 144°57’E,

Jan 2002, Forster PIF28196 et al. (A, BRI, DNA, L,

MEL, NSW, WIS); Richards Creek, Mt Mulligan,

16°52’S, 144°53’E, Nov 1999, Holmes 131 (BRI); 3 km

from Dimbulah on the Mareeba road, 17°10’S,

145°10’E, Oct 1975, Hyland 8472 (BRI, QRS).

Austrobaileya 6 (3): 349-436

Distribution and habitat : Croton minimus is

known only from west of Dimbulah, around

Chillagoe and north of Mt Carbine in north-east

Queensland, over three 1° squares (Map 4).

Plants grow in open forest with Corymbia

clarksoniana (D.J.Carr & S.GM.Carr) K.D.Hill

& L.A.S.Johnson on skeletal soils derived from

granite on steep ridges.

Phenology: Flowering occurs from December

to April with fruiting from December to May.

Notes: Croton minimus appears to be a

miniature derivative of C. arnhemicus or

C. multicaulis that has evolved in response

to the xeric environment where it has been

found. A comparison of these three species is

given in Table 1. Both Croton minimus and

C. multicaulis subsp. velutinus come into close

proximity to one another in the Spring Creek

area north of Mt Carbine (vouchers: Forster

PIF12928 & Bean', Forster PIF18128 & Spokes )

and further field work is required to see if the

two taxa intergrade.

Conservation status : This species has been

rarely collected, but is an inconspicuous plant

and vast areas of suitable habitat exist in the

known range. Hence it is probable that many

more populations can be found once this area

is properly explored. No conservation coding

is considered necessary.

Etymology: The specific epithet is derived from

the Latin minimus and refers to the small stature

of this plant.

Table 1. Comparison of morphological characters for Croton arnhemicus , C. minimus and

C. multicaulis

Character

C. arnhemicus

C. minimus

C. multicaulis

Habit

shrub or tree to 5 m high

subshrub 30cm high

subshrub 1.5m high

no. of marginal

teeth on leaf

60-100

22-28

32-56

leaf lamina

lateral veins

prominent

weak

prominent

leaf lamina

prominent

obscure

prominent

interlateral veins

no. of stamens/

flower

20-44

16-18

11-24

Forster, Croton in Australia

395

Fig. 17. Croton minimus. A. flowering branchlet. x 1. B. undersurface of leaf, x 1. C. leaf lamina base demonstrating

absence of extrafloral nectaries, x 6. D. inflorescence, x 2. E. male flower, x 8. F. female flower, x 8. G & H. fruit,

x 4. I. seed, x 6. All from Forster PIF 14708 (BRI). Del. W. Smith.

396

19. Croton multicaulis P.I.Forst., sp. nov. affinis

C. arnhemico Muell.Arg. a qua habitu

suffrutice multicauli semper, foliis

dentibus minus quam 60, et staminibus

11-24 (plerumque minus quam 20) differt.

Typus: Queensland. Cook District: 20.5

km along Weipa road, off Peninsula

Development road, 13°03’S, 142°36’E, 7

December 1993, P.I. Forster PIF14367

(holo: BRI [2 sheets + spirit]; iso: DNA, L,

MEL).

Croton sp. (Myall Creek P.I.Forster PIF14368)

(Forster & Henderson 1997:72; Forster &

Halford 2002:71).

Multistemmed shrub to 1.5 m high, monoecious,

deciduous, perennial. Indumentum ferruginous-

silver. Branchlets rounded, with dense stellate

trichomes. Stipules linear to linear-lanceolate,

1.2-6 mm long, 0.2-0.6 mm wide, entire and with

dense stellate trichomes. Leaves alternate,

discolorous, petiolate; petioles 2-40 mm long,

1-1.7 mm wide, with dense stellate trichomes;

lamina broadly ovate elliptic, lanceolate-ovate,

or ovate, 15-140 mm long, 11-115 mm wide,

palminerved with 3-5 veins at base, 4-6 lateral

veins per side of midrib further up lamina and

tertiary reticulate veins; upper surface matt

green, lateral veins weakly visible, with

scattered to sparse stellate trichomes; lower

surface pale matt green, lateral and tertiary veins

prominent, with sparse to dense stellate

trichomes and sometimes with stalked yellow

glandular trichomes, scabrid to velutinous;

margins crenate with 16-28 teeth up to 4 mm

long, foliar glands prominent; tip acute to

rounded; base cordate, rounded or truncate;

extrafloral nectaries absent or 2 at lamina base,

sessile or stipitate to 0.8 mm long, circular to

ellipsoid, 0.3-2 mm long, 0.2-1 mm wide, visible

above and below. Inflorescence up to 150 mm

long, usually androgynous, pedunculate up to

45 mm; axis with sparse to dense stellate

trichomes; bracts linear to linear-lanceolate, 0.5-

4 mm long, 0.2-0.5 mm wide, with dense stellate

trichomes. Male flowers 1.8-4 mm long, 2.5-5

mm diameter, held singly on inflorescence or in

Austrobaileya 6 (3): 349-436

dense clusters of 1-5 flowers, spaced up to 4

mm apart; pedicels 1.5-7 mm long, 0.3-1 mm

wide, with dense stellate trichomes; sepals

valvate, 5, lanceolate-ovate to ovate, 1.8-3 mm

long, 1-2.2 mm wide, with sparse to dense

stellate trichomes; petals 5, oblanceolate to

obovate, 1.5-3.2 mm long, 0.5-1.5 mm wide,

lanate; stamens 11-24, filaments filiform, 1-3.1

mm long, 0.1-0.2 mm wide, with dense simple

trichomes at base; anthers oblong, 0.5-1 mm

long, 0.4-0.6 mm wide. Female flowers 3^4.5 mm

long, 2.5-5 mm diameter, held singly and spaced

up to 15 mm apart; pedicels 0.8-6 mm long, 0.8-

1.2 mm diameter, with dense stellate trichomes;

sepals valvate, 5, lanceolate-ovate, 1.7-3 mm

long, 0.9-1.8 mm wide, with dense stellate

trichomes; petals absent; styles 3, linear, 1.2-

3.2 mm long, bifid for 1.1-3 mm long, connate at

base for c. 0.2 mm, with scattered stellate

trichomes in lower parts; ovary 3-locular, 2-3.5

mm long, 2-3.5 mm diameter, with dense, stalked,

stellate trichomes. Fruits trilobate, depressed-

globose, 5-8 mm long, 7-8 mm diameter, with

dense, stalked, stellate trichomes. Seeds obloid

to ovoid, 4-5 mm long, 2.5^1.5 mm wide, 2-3.5

mm thick, black-brown to grey, ventral surface

bifacial, dorsal surface ± rounded, micropylar

ridge 3-4 mm long; caruncle crescent shaped,

0.6-1.5 mm long, 0.8-3.5 mm wide, cream.

Distribution : Croton multicaulis is endemic to

north Queensland from the islands of Torres

Strait in the north, south to Porcupine Gorge

near Hughenden, covering a total of thirteen

1° grid squares.

Notes: Croton multicaulis is superficially very

similar to C. arnhemicus but differs in several

ways. Croton multicaulis is always a

multistemmed subshrub with less than 60 teeth

on the margins of the leaf lamina and with 11-

24 (mostly less than 20) stamens per flower.

Croton arnhemicus may persist as a

multistemmed subshrub for an indefinite period,

but ultimately grows into a small tree, has 60 or

more teeth on the margins of the leaf lamina and

has 20-44 (mostly more than 28) stamens per

flower.

Key to subspecies of Croton multicaulis

Foliage scabrid below.

Foliage soft velutinous below

subsp. multicaulis

. subsp. velutinus

Forster, Croton in Australia

19a. Croton multicaulis subsp. multicaulis

Foliage scabrid below with sparse stellate

trichomes. Fig. 18.

Selected additional specimens : Queensland. Cook

District: Thursday Island, Jun 1897, Bailey [AQ202076]

(BRI); c. 18 km NW of Silver Plain s Station, 13°52’S,

143°24’E, Nov 1980, Clarkson 3614 (BRI); 0.5 km S

of Watson River Crossing on the Aurukun to Merluna

road, c. 40 km NE of Aurukun, 13°08’S, 142°01’E,

Dec 1981, Clarkson 4043 (BRI, QRS); Chester River

Scrub, eastern fall of Mcllwraith Range, 13°40’S,

143°29’E, Jun 1992, Forster PIF10400 et al. (BRI);

Granny Scrub, Weipa to Stones Crossing road, 42 km

from Weipa, Jul 1993, Forster PIF13500 et al. (BRI);

Scrubby Creek Scrub, Silver Plains, 13°44’S, 143°28’E,

Jul 1993, Forster PIF13626 et al (BRI, MEL, NSW);

Northern end of Bamboo Range, 14°36’S, 143°27’E,

Dec 1993, Forster PIF14364 (BRI); Myall Creek

Crossing, Weipa road, 12°39’S, 142°16’E, Dec 1993,

Forster PIF14368 (BRI); Cowal Creek floodplain, Cape

York, 10°55’S, 142°18’E, Jun 1994, Forster PIF15336

(BRI, QRS); Tragia Scrub, 3.5 km ESE of Mutee Head,

Cape York, 10°55’S, 142°16’E, Jun 1994, Forster

PIF15337 (BRI, QRS); 6.5 km from Captain Billy

Landing, 11°37’S, 142°48’E, Jun 1994, Forster

PIF15361 (BRI, MEL); 27 km SE of Heathlands,

11°52’S, 142°38’E, Feb 1992, Johnson 4993 (BRI,

DNA, MEL, NSW); Weipa, 3 km E of Lorim Point,

12°41’S, 141°53’E, Jan 1981, Morton 1050 (BRI);

Herring Oil Slot, Weipa, 12°39’S, 141°50’E, Dec 1989,

O’Reilly 500 (BRI); Bamaga Mission, 11.2 km SW of

Cape York, Oct 1965, Smith 12364 (BRI); 5 km ENE

of Weipa Mission, 12°38’S, 141°56’E, Jul 1974, Specht

347 & Salt (BRI); ditto, Dec 1974, Specht W43 & Salt

(BRI); 23.5 km ENE of Weipa Mission, Dec 1974,

Specht W190 & Salt (BRI); Laradeenya Creek, Cape

York, Jun 1963, Stephens [AQ202078] (BRI); New

Mapoon, Northern Peninsula area, 10°52’S, 142°23’E,

Jan 1998, Waterhouse BMW4785 (BRI).

Distribution and habitat: Croton multicaulis

subsp. multicaulis is endemic to Torres Strait

and the northern parts of Cape York Peninsula

north of Musgrave (Map 8). Plants often grow

on the margins of vineforest, but are more

common in adjacent Eucalyptus tetrodonta

F.Muell. woodland on white sandy or red lateritic

soils.

Phenology : Flowering occurs from November

to July with fruiting two to three months later.

Conservation status: Croton multicaulis

subsp. multicaulis is common throughout its

known range; however, no populations appear

to be present in conservation reserves at

present. No conservation coding is required.

Etymology : The specific epithet is derived from

397

the Latin multi (many) and caulis (stemmed)

and refers to the habit of this plant.

19b. Croton multicaulis subsp. velutinus

P.I.Forst., subsp. nov. affinis C. multicauli

P.I.Forst. a qua foliis velutinis

trichomatibus stellatis densis abaxialiter

differt Typus: Queensland. Cook District:

19 km from Laura on road to New Laura

homestead, 15°25’S, 144°25’E, 22 January

1993, P.I. Forster PIF12829 &A.R. Bean

(holo: BRI [1 sheet + spirit]; iso: DNA,

MEL).

Foliage velutinous below with dense stellate

trichomes. Fig. 19.

Selected additional specimens: Queensland. Cook

District: Musgrave Telegraph Station, s.dat., Barclay-

Millar [AQ202070](BRI); Little Laura River, SSW of

Laura, 15°42’S, 144°17’E, Jul 1990, Bean 1905 (BRI);

Blue Hills, 49 km from Mt Surprise township, 17°58’S,

144°02’E, Mar 1988, Champion 340 (BRI); 0.9 km E

of the West Normanby River on the Lakeland Downs

to Cooktown road, 15°46’S, 144°59’E, May 1987,

Clarkson 6754 & McDonald (BRI, MBA, QRS); Cape

Melville N.R, Altanmoui Range section, 14°30’S,

144°35’E, May 1993, Fell DGF3132 & Stanton (BRI);

2 km E of Mt Gibson, 16 km SSE of Lakeland Downs,

West Normanby River catchment, May 1993, Fell

DGF3281 & Daunt (BRI); Birthday Mt., Rokeby N.P.,

23 km N of Coen aerodrome, Aug 1993, Fell DGF3522

& Jensen (BRI); 31 km from Laura on road to New

Laura Homestead, 4 Mile Swamp area, Lakefield N.R,

15°18’S, 144°25’E, Jan 1993, Forster PIF12822 &

Bean (BRI, MEL); New Laura Homestead area,

Lakefield National Park, 15°11’S, 144°20’E, Jan 1993,

Forster PIF12895 & Bean (BRI, DNA, MEL); Spring

Creek, Mt Carbine to Laura road, 16°22’S, 144°43’E,

Jan 1993, Forster PIF12928 & Bean (BRI, MEL);

Lake Emma, Lakefield N.P, 15°17’S, 144°38’E, Jan

1993, Forster PIF 12939 & Bean (BRI, DNA, MEL);

Giant Horse Gallery (Laura), 15°40’S, 144°30’E, Mar

1975, Hyland 8111 (BRI, QRS); 19.2 km N of Laura,

15°25’S, 144°25’E, Oct 1974, Robinson [AQ196265];

Lakefield N.P., 20 km SW of Lakefield Homestead,

Aug 1983, Stanton [AQ349838] (BRI). BURKE

DISTRICT: Porcupine Gorge, 53 km NNE of

Hughenden, 20°25’S, 144°26’E, May 1990, Halford

Q228 (BRI); 55 km NE of Hughenden at Porcupine

Gorge N.P. lookout, 20°24’S, 144°26’E, Nov 1992,

Thompson HUG72 & Turpin (BRI, DNA).

Distribution and habitat'. Croton multicaulis

subsp. velutinus has been found in the southern

part of Cape York Peninsula from near Musgrave

south to Lakefield National Park near Laura and

has been found in six 1° grid squares (Map 3).

There are also several apparently disjunct

populations at Porcupine Gorge and Mt Surprise

398

Austrobaileya 6 (3): 349-436

Fig. 18. Croton multicaulis subsp. multicaulis. A. habit of flowering branchlet. x 0.6. B. undersurface of leaf,

x 0.8. C. base of leaf lamina showing extrafloral nectaries, x 8. D. node showing stipules, x 8. E. inflorescence with

female flowers towards base and male flowers towards apex, x 2. F. male flower, x 6. G. female flower, x 8. H. seed,

x 6. A, D-E from Forster PIF14368 (BRI); B from Forster PIF15361 (BRI); C & H from Waterhouse 4785 (BRI);

F & G from Forster PIF13626 (BRI). Del. W. Smith.

Forster, Croton in Australia

that have been tentatively placed with this

taxon. Plants grow in open eucalypt woodland

with Eucalyptus leptophleba F.Muell. and

Melaleuca viridiflora Sol. ex Gaertn. on reddish

sandy soils or in open eucalypt forest with

Corymbia clarksoniana and E. tetrodonta on

red lateritic soils.

Phenology : Flowering occurs from November

to July with fruiting two to three months later.

Conservation status: Croton multicaulis

subsp. velutinus is common throughout its

known range. Some populations are present in

conservation reserves at Lakefield, Porcupine

Gorge and Rokeby National Parks.

Etymology: The subspecific epithet is derived

from the Latin velutinus (velvety) and refers to

the dense indumentum on the lower leaf

surfaces of this plant.

Conservation status : This subspecies is

widespread, and present in at least three

National Parks. No conservation coding is

required.

20. Croton mutabilis P.I.Forst., sp. nov. affinis

C. byrnesii Airy Shaw a qua indumento

ecolorato usque argenteo, foliis sinuatis

usque denticulatis dentibus 18-24, venis

lateralibus paucioribus (8-10), et petiolis

juvenibus trichomatibus dispersis stellate

et peltatis differt. Typus: Queensland.

Cook District: Possum Scrub, road to

Stone’s Crossing from Weipa, 12°27’S,

142°09’E, 8 December 1993, P.I. Forster

PIF14376 (holo: BRI [2 sheets + spirit];

iso: DNA, L, MEL, MO, NSW).

Croton sp. (Possum Scrub P.I.Forster

PIF14376) (Forster & Henderson 1997:72;

Forster & Halford 2002:71).

Shrub to 4 m high, monoecious, deciduous,

perennial. Indumentum uncoloured or silver.

Branchlets + rounded, glabrous or with

scattered stellate trichomes, glabrescent.

Stipules linear, 1.8-2 mm long, c. 0.2 mm wide,

entire and glabrous. Feaves alternate, petiolate,

discolorous; petioles 7-70 mm long, 0.5-1.5 mm

wide, with scattered stellate to peltate trichomes

when young, glabrescent; lamina elliptic,

obovate or ± orbicular, 13-170 mm long, 7-90

399

mm wide, penninerved with 8-10 lateral veins

per side of midrib and tertiary reticulate veins;

upper surface dark green, lateral veins weakly

developed, glabrous; lower surface pale green,

lateral and tertiary veins weakly developed,

glabrous or with stellate trichomes, neither

scabrid nor velutinous; margins sinuate or

denticulate with 9-17 teeth up to 1 mm long,

foliar glands prominent; tip acute to rounded;

base cuneate, rounded or truncate; extrafloral

nectaries 2 at lamina base, sessile, ellipsoid,

0.3-1.3 mm long, 0.2-0.7 mm wide, visible below

only. Inflorescence up to 70 mm long,

androgynous, pedunculate up to 25 mm; axis

glabrous; bracts lanceolate, 0.4-2.5 mm long,

0.2-0.7 mm wide, with scattered simple

trichomes and scattered stellate trichomes.

Male flowers 2.5-4 mm long, 4-6 mm diameter,

held singly or in pairs on inflorescence, spaced

up to 5 mm apart; pedicels 3-4 mm long, 0.4-0.7

mm wide, glabrous; sepals valvate, 5, lanceolate-

ovate, 2-2.5 mm long, 1.2-1.5 mm wide, lanate

on tips; petals 5, oblanceolate, 2-3 mm long,

0.5-1 mm wide, lanate on tips; stamens 10-12,

filaments ± flattened, 1.5-2.8 mm long, 0.2-0.3

mm wide, with dense simple trichomes at base;

anthers oblong, 0.8-1 mm long, 0.7-0.8 mm wide.

Female flowers 3-3.5 mm long, 3.5-4 mm

diameter, held singly and spaced up to 10 mm

apart; pedicels 2.5-4 mm long, 0.5-0.7 mm

diameter, glabrous; sepals valvate, 5, lanceolate-

ovate, 2-2.5 mm long, 1-1.3 mm wide, lanate on

tips; petals absent; styles 3, linear, 2-2.5 mm

long, bifid for 1.1-2.5 mm long, connate at base

fore. 0.5 mm, glabrous; ovary 3-locular, c. 2 mm

long and 2 mm diameter, with dense, sessile

stellate trichomes. Fruits trilobate, depressed-

globose, 4-5 mm long, 6-7 mm diameter, with

sparse, sessile stellate trichomes. Seeds ±

obloid, 3.5-3.8 mm long, 2.8-3 mm wide, 2-2.2

mm thick, glossy brown, ventral surface bifacial,

dorsal surface rounded, micropylar ridge 2.5-

2.8 mm long; caruncle crescent shaped, c. 1 mm

long and 1 mm wide, pale brown. Fig. 20.

Additional specimens: Queensland. Cook District:

Chester River Scrub, eastern fall of Mcllwraith Range,

Silver Plains Sation, 13°40’S, 143°29’E, Jun 1992,

Forster PIF10423 & Tucker (BRI); Nesbit River,

13°32’S, 143°31’E, Jun 1992, Forster PIF10507 et al.

(BRI); Massy Creek Scrub, Silver Plains Station,

13°55’S, 143°30’E, Jun 1992, Forster PIF10598 et al.

(BRI, L, MEL, NSW); Stones Crossing, 73 km from

Weipa, Jul 1993, Forster PIF13506 et al. (BRI); Possum

Scrub, Weipa to Stones Crossing road, 12°27’S,

400

Austrobaileya 6 (3): 349-436

Fig. 19. Croton multicaulis subsp. velutinus. A. flowering branchlet. x 0.6. B. undersurface of leaf, x 0.8. C. base

of leaf lamina showing extrafloral nectaries, x 6. D. inflorescences, x 1. E. male flower, x 6. F. female flower, x 6.

G & H. fruit, x 3. I. seed, x 6. A,B from Forster PIF12822 (BRI); C-I from Forster PIF12829 (BRI). Del. W.

Smith.

Forster, Croton in Australia

142°09’E, Jul 1993, Forster PIF13515 et al. (BRI);

Pascoe River crossing, road to Iron Range, 12°53’S,

143°00’E, Jul 1993, Forster PIF13532 et al (BRI,

QRS); Near Ham Hill (Weymouth Holding), 12°45’S,

143°20’E, Oct 1973, Hyland 6990 (BRI, QRS); Claudie

River, Oct 1974, Hyland 7817 (BRI, QRS); T.R. 14

Massy, 13°52’S, 143°25’E, Nov 1980, Hyland 10858

(BRI, QRS); Claudie River, Jan 1982, Hyland 11504

(BRI, QRS); ditto, Dec 1982, Hyland 12401 (BRI,

QRS); cult. Tolga (ex Possum Scrub, Weipa), Nov 1991,

Sankowsky 1316 & Sankowsky (BRI); Nesbit River,

Oct 1986, Tucker 61 (BRI); Claudie River, Oct 1968,

Webb & Tracey 8542 (BRI, CANB); between Iron Range

airstrip and Portland Roads - Coen road, 12°40’S,

143°23’E, Oct 1968, Webb & Tracey 8673 (BRI,

CANB).

Distribution and habitat: Croton mutabilis is

endemic to northern Cape York Peninsula,

Queensland and occurs in four 1° grid squares

(Map 9). Plants grow on the margins of

deciduous vine thickets and semi-deciduous

notophyll vineforests on alluvium, red laterite

or heavy black clay soils.

Notes: Sterile collections of Croton mutabilis

were misidentified by Airy Shaw (1981) as

C. storckii Seem, ex A.C.Sm. Smith (1981)

401

reduced Croton storckii to the synonymy of

C. microtiglium Burkill, a species endemic to

Fiji.

Croton mutabilis appears to be allied to

C. byrnesii and C. habrophyllus, all three being

deciduous and allopatric. Croton mutabilis

differs from these two species in a number of

characters (Table 2). While it appears similar to

C. microtiglium this latter species differs in a

number of characters (Table 3) and is not

necessarily closely related.

Considerable variation occurs in leaf

morphology of Croton mutabilis. Most

collections of this plant have been made in

winter when the plants are sterile and the leaves

are well developed and turning orange prior to

abscission. Flowering occurs following storm

rains, when the plant produces flushes of new,

and much smaller, foliage.

Conservation status: Croton mutabilis is

relatively common throughout its known range

and is present in Iron Range National Park.

Table 2. Comparison of morphological characters for Croton byrnesii , C. habrophyllus and

C. mutabilis

Character

C. byrnesii

C. habrophyllus

C. mutabilis

indumentum colour

ferruginous

to yellow

silver

uncoloured

silver

leaf lamina margins

crenate

sinuate to

denticulate

sinuate to

denticulate

no.of marginal teeth

per leaf lamina

40-58

70-112

18-24

no. of lateral veins each

side of midrib

11-13

9-11

8-10

trichomes on petioles

when young

sparse stellate

scattered to dense

stellate

scattered stellate

& peltate

trichomes on shoot tips

or when young

scattered

stellate

dense to sparse

stellate

glabrous or

scattered stellate

stamen filaments

flattened

filiform

flattened

402

Austrobaileya 6 (3): 349-436

Fig. 20. Croton mutabilis. A. flowering branchlet. x 0.6. B. base of leaf lamina showing extrafloral nectaries, x 6.

C. node with stipule, x 6. D & E. undersurface of leaf, x 0.8. F. inflorescence of mainly female flowers, x 1. G. shoot

tip with inflorescence of male flowers, x 1. H. male flower, x 8. I. female flower, x 8. J & K. parts of dehisced fruit

with seed inside, x 6. L. seed, x 8. A, B, E & F from Sankowsky 1316 (BRI); C & D from Tucker 61 (BRI); G-I from

Forster PIF14376 (BRI); J-L from Hyland 11504 (BRI). Del. W. Smith.

Forster, Croton in Australia 403

Table 3. Comparison of morphological characters for Croton microtiglium and C. mutabilis

Character

C. microtiglium

C. mutabilis

branchlet indumentum

peltate trichomes

stellate & peltate trichomes

petiole indumentum

peltate trichomes

stellate & peltate trichomes

lateral vein pairs

in leaf lamina

9-11

8-10

extrafloral nectaries

poorly formed

well developed,

embedded

sessile

pedicel indumentum

dense peltate

trichomes

glabrous

Etymology: The specific epithet is derived from

the Latin mutabilis (changeable) and refers to

the developmental change in leaf lamina

morphology in this species that occurs between

flowering and leaf abscission.

21. Croton phebalioides F.Muell. ex Muell.Arg.,

Flora 47: 485 (1864); Oxydectes

phebalioides (‘phebaliodes’) (F.Muell. ex

Muell.Arg.) Kuntze, Rev. Gen. PL 2: 612

(1891). Type: Queensland. North Kennedy

District: Burdekin River, F. Mueller (holo:

K n.v., photo atBRI!).

Croton maidenii R.T.Baker, J. & Proc. Roy.

Soc. New South Wales 48:444,1.12 (1915).

Type: New South Wales. Guthrie’s

Mountain (Read’s Mine), 1904,

A.Paddison (holo: NSW n.v.; iso: BRI).

Croton phebalioides var. acuminatus Domin,

Biblioth. Bot. 89: 326 (1927). type:

Queensland. Moreton District: Prope

Brisbane River, 1863-1865, A. Dietrich

2326 (syn: PR528548); Queensland. North

Kennedy District: Edgecombe Bay,

Dallachy (syn: K n.v., photo at BRI!;

MEL231441).

Croton phebalioides var. typicus Domin,

Biblioth. Bot. 89: 326 (1927), nom. inval.

type: same as for C. phebalioides

F.Muell. ex Muell.Arg.

Illustrations : James & Harden (1990: 420);

Hauser (1992:100).

Shrub or small tree to 8 m high, monoecious,

evergreen, perennial. Bark lenticellate, grey;

blaze thick, flaky, cream-tan; wood cream-tan.

Indumentum uncoloured to silver. Branchlets ±

rounded, with dense stellate trichomes. Stipules

lanceolate to lanceolate-ovate, 0.3-8 mm long,

0.2-0.5 mm wide, entire and with dense stellate

trichomes. Leaves alternate, petiolate,

discolorous; petioles 3-16 mm long, 0.8-1 mm

wide, with dense stellate trichomes; lamina

elliptic, lanceolate or ovate, 8-118 mm long, 3-

38 mm wide, penninerved with 7-14 lateral veins

per side of midrib, tertiary reticulate veins

obscure; upper surface matt dark-green,

venation obscure, glabrous or with sparse

stellate trichomes; lower surface silver, lateral

veins weakly visible, with dense overlapping

peltate trichomes, neither scabrid nor

velutinous; margins entire, sinuate or

denticulate with 11-18 weakly defined teeth less

than 0.2 mm long, foliar glands prominent; tip

acuminate, acute, obtuse or mucronate; base

cuneate; extrafloral nectaries 2 at lamina base,

sessile, ellipsoid, 0.1-0.3 mm long, 0.1-0.2 mm

wide, visible above only. Inflorescence up to 95

mm long, unbranched, androgynous,

pedunculate up to 12 mm; axis with dense

stellate to peltate trichomes; bracts lanceolate-

ovate to ovate, 0.4-0.8 mm long, 0.2-0.7 mm

wide, with dense peltate trichomes. Male flowers

2-3 mm long, 2.5-4.5 mm diameter, held singly

or 2 to 3 per glomerule, spaced up to 5 mm apart

but usually densely clustered towards top of

inflorescence; pedicels 1.5-3.5 mm long, 0.3-

0.5 mm wide, with dense peltate trichomes;

sepals valvate, 5, lanceolate-ovate to obovate,

404

1.2-2.7 mm long, 1.3-1.8 mm wide, with dense

peltate trichomes; petals 5, obovate, 1.3-3 mm

long, 0.5-0.7 mm wide, lanate; stamens 10-12,

filaments flattened, 1.6-2.5 mm long, 0.2-0.4 mm

wide, glabrous, anthers oblong, 0.8-1.2 mm

long, 0.6-1.2 mm wide. Female flowers 2.5^1 mm

long, 3-5.5 mm diameter, held singly and spaced

up to 15 mm apart; pedicels 2-6 mm long, 0.6-1

mm diameter, with dense peltate trichomes;

sepals valvate, 5, lanceolate-ovate to obovate,

2.5-3.5 mm long, 1.5-2.2 mm wide, with dense

peltate trichomes, lanate; petals absent; styles

3, flattened-flabellate, 1.2-2.5 mm long, multifid,

twice divided for 1-2 mm long, + free at base,

glabrous or sparsely papillose near base; ovary

3-locular, 2-3 mm long, 2.6-4 mm diameter, with

dense, sessile and stalked stellate trichomes.

Fruits trilobate, depressed-globose, 6-8 mm

long, 6.5-9 mm diameter, with dense, sessile and

stalked stellate trichomes. Seeds ovoid, 4-5 mm

long, 3.2-4 mm wide, 2.3-2.5 mm thick, pale to

dark brown, ventral surface bifacial, dorsal

surface rounded, micropylar ridge 2.8-3.8 mm

long; caruncle broadly ovate, 1-1.2 mm long,

1.4-1.5 mm wide, cream to cream-yellow. Fig.

21 .

Selected additional specimens: Queensland. Cook

district: Mungana, near Red Dome Mine turnoff,

17°06’S, 144°25’E, Jan 2002, Forster PIF28144 et al.

(A, BRI, MEL, WIS); Mt Elephant, Curramore Holding,

16°27’S, 144°56’E, Apr 1987, Wolfe 2 (QRS). Burke

District: Prairie Creek Gorge, 45 km NNE of

Hughenden, Jun 1986, Murray 62 & Morgan (BRI).

North Kennedy District: Mingela Bluff, 19°53’S,

146°45’E, Jan 1992, Forster PIF9436 & Bean (BRI,

K, MEL, QRS); Mt Inkerman, 19°44’S, 147°29’E, Mar

1999, Forster PIF24215 (A, AD, BRI, K, L, MEL,

QRS). South Kennedy District: Carlisle Is, c. 1 km W of

Turtle Bay & c. 35 km N of Mackay, 20°47’S,

149°17’E, Sep 1986, Sharpe 4450 & Batianoff (BRI).

Leichhardt District: Palmgrove N.P., NW of Taroom,

25°01’S, 149°15’E, Nov 1998, Forster PIF23808 (BRI,

MEL, QRS); Expedition N.P., Amphitheatre section,

Cannondale Scrub, 25°12’S, 148°59’E, Nov 1998,

Forster PIF23869 (BRI, MEL, QRS). Port Curtis

District: Mt Etna, 23°10’S, 150°27’E, Nov 1987, Vavryn

21 (BRI). Mitchell District: Gowan Range, c. 20 km

NNW of Idalia HS, 24°43’S, 144°41’E, Apr 1984,

Purdie 2071 (BRI). Burnett District: Kalliwa Creek,

S.F. 169, St Agnes, 25°18’S, 151°51’E, Dec 1990,

Forster PIF7717 (BRI, K, L, MEL, MO, QRS); S.F.

695 Kalpowar, Burnett Range road, 24°42’S, 151°20’E,

Mar 2000, Forster PIF25409 & Booth (BRI, MEL,

QRS); Coalstoun Lakes N.P, 16 km SW of Biggenden,

25°35’S, 151°54’E, Dec 2002, Forster PIF29182 (A,

BRI, L, MEL, NE, NSW, WIS). Wide Bay District: Mt

Austrobaileya 6 (3): 349-436

Biggenden, 25°32’S, 151°59’E, Jan 1991, Forster

PIF7738 (BRI, MEL, QRS); Lime Mine road, between

Didcot & Coalstoun Lakes, 25°33’S, 151°53’E, Dec

2001, Forster PIF28061 (A, BRI, K, L, MEL, WIS).

Warrego District: 13 km W of Morven, 26°09’S,

146°59’E, Jun 1978, Purdie 766d (BRI). Maranoa

District: “Stafford Park”, Ulandilla, Jan 1936, Hewitt

[AQ202176] (BRI). Darling Downs District: “Kilbumie”

area, 26°47’S, 150°27’E, Oct 1985, Hoy 92(BRI).

Moreton District: Ivorys Knob, west slopes, 10 km NE

of Boonah at end of Hansens road, Nov 1986, Bird

[AQ431621] (BRI, NSW). New South Wales. Duri

Mt, 20 km WSW of Tamworth, 31°12’S, 150°43’E,

Nov 2000, Copeland 2760 (BRI, NE).

Distribution and habitat : Croton phebalioides

is widespread in central and southern

Queensland with an apparent northern limit at

Mt Elephant. It also occurs in north-eastern New

South Wales (Map 8). It is the most widespread

Croton in mainland Australia by its presence in

fifty-five 1° grid squares. Plants grow in semi¬

evergreen vinethickets throughout much of the

range, although some of the northern

populations are present in deciduous

vinethicket.

Phenology: Flowering and fruiting occurs

throughout the year following storm rains, but

is concentrated from September to December.

Notes: Croton phebalioides is quite variable in

terms of leaf lamina size. Much of this variation

appears to be related to aridity, as the

populations in drier, more inland localities tend

to have smaller leaves than those from nearer

the coast.

Although Croton phebalioides is usually

a shrub, at least one locality it forms a small tree

to 8 m high (Forster PIF29182).

This species was included in Croton

section Croton by Webster (1993a), but does

not conform with the characters given for that

section, e.g. the penninerved foliage (versus

palminerved) and the 10-12 stamens (versus

15-35).

Conservation status: This is a very common

plant and is present in twelve conservation

reserves in south-eastern Queensland alone

(Forster etal. 1991).

Etymology: The specific epithet refers to a

resemblance between the foliage of this plant

and some species of Phebalium (Rutaceae).

Forster, Croton in Australia

405

Fig. 21. Croton phebalioides. A. flowering branchlet. x 1. B & C. undersurface of leaves showing variation in size

and primary venation, x 0.8. D. base of leaf lamina showing extrafloral nectaries, x 6. E. inflorescence with male

flowers, x 1.5. F. male flower, x 8. G. female flower, x 6. H & I. fruits, x 4. J. seed, x 6. A & F from Bird AQ431621

(BRI); B & D from Forster PIF7717 (BRI); Cl from Forster PIF6571 (BRI); C2 from Forster PIF13410 (BRI); G

from Forster PIF2176 (BRI); H-J from Gordon AQ202185 (BRI). Del. W. Smith.

406

22. Croton rarus P.I.Forst., sp. nov. affinis

C. dockrillii Airy Shaw a qua lamina

crenata et venis utroque costae 12-14,

indumento ferruginei-argenteo, glandibus

foliaribus sessilibus, et pedicello florum

marium breviore (1-2 mm longo) differt.

Typus: Queensland. Cook District: 4.5 km

from the Watson River Crossing on the

Aurukun - Merluna road, c. 40 km NE of

Aurukun, 13°07’S, 141°59’E, 3 December

1981, J.R. Clarkson 4062A (holo: BRI [1

sheet]; iso: QRS; DNA, K, L, MO n.v.)

Croton sp. (Watson River J.R. Clarkson

406IB) (Forster & Henderson 1997: 72;

Forster & Halford 2002:71).

Shrub to 5 m high, monoecious, evergreen,

perennial. Indumentum ferruginous-silver.

Branchlets ± rounded, with dense stellate

trichomes when young, glabrescent. Stipules

linear, 2-3.9 mm long, c. 0.2 mm wide, entire and

with sparse to dense stellate trichomes. Leaves

alternate, discolorous, petiolate; petioles 6-30

mm long, 0.7-0.8 mm wide, with dense stellate

trichomes; lamina narrowly elliptic to

oblanceolate, 20-110 mm long, 10-35 mm wide,

penninerved with 12-14 lateral veins per side

of midrib and poorly developed tertiary

reticulate veins; upper surface dark green,

venation not visible, glabrous; lower surface

pale green, venation weakly developed, with

scattered to sparse, stellate trichomes, neither

scabrid nor velutinous; margins crenate with

16-32 teeth up to 0.5 mm long, foliar glands

prominent; tip acute to rounded; base cuneate

to rounded; extrafloral nectaries 2 at base of

lamina, sessile, ellipsoid, 0.7-1 mm long, 0.5-

0.6 mm wide, visible mainly below. Inflorescence

up to 120 mm long, unbranched, androgynous,

pedunculate up to 5 mm; axis with dense stellate

trichomes; bracts linear-lanceolate to lanceolate,

1-3.5 mm long, 0.2-0.3 mm wide, with scattered

simple and stellate trichomes. Male flowers 3-

3.5 mm long, 3-4.5 mm diameter, densely

clustered in glomerules of 1-5 flowers towards

the top of the inflorescence; pedicels 2.2-2.6

mm long, 0.3-0.4 mm wide, with scattered stellate

trichomes; sepals valvate, 5, obovate, 2-2.3 mm

long, 1-1.2 mm wide, glabrous; petals 5,

Austrobaileya 6 (3): 349-436

oblanceolate, 2.3-2.5 mm long, 0.7-0.8 mm wide,

lanate in upper half; stamens 10-12, filaments

filiform-flattened, 2.2-3 mm long, c. 0.1 mm wide,

glabrous, anthers oblong, 0.7-0.8 mm long, 0.6-

0.7 mm wide. Female flowers 4-4.5 mm long, 3.5-

4 mm diameter, held singly or in groups of 2-4

and spaced up to 11 mm apart; pedicels 1-2 mm

long, 0.7-1 mm diameter, with sparse stellate

trichomes; sepals valvate, 5, lanceolate-ovate,

2-3.5 mm long, 1-2 mm wide, with scattered

stellate trichomes; petals absent; styles 3, linear,

2-3.2 mm long, bifid for 1.5-2.5 mm, glabrous,

connate at base for c. 0.2 mm; ovary 3-locular,

1.5-2 mm long, 1.8-2.5 mm diameter, with dense,

sessile stellate trichomes. Fruits trilobate,

globose, 4-5 mm long, 4-4.5 mm diameter, with

scattered, sessile stellate trichomes. Seeds

ovoid, 3.3-4 mm long, c. 3 mm wide and 2 mm

thick, dark-brown, micropylar line 2.2-2.5 mm

long; caruncle not seen. Fig. 22.

Additional specimens : Queensland. Cook District:

Kowanyama, Mitchell River, Mar 1978, Alpha & Black

202B (BRI); 4.5 km from the Watson River Crossing

on the Aurukun - Merluna road, c. 40 km NE of Aurukun,

13°07’S, 141°59’E, Dec 1981, Clarkson 4061B (BRI,

QRS); Rokeby N.R, old Archer River crossing,

Wenlock, 13°26’S, 142°42’E, Apr 1991, Fell DGF2291

(BRI); 12 km along road to Weipa, off Peninsula

Development road, 13°04’S, 142°40’E, Jul 1993,

Forster PIF13478 et al. (BRI, MEL, QRS); Stones

Crossing, c. 73 km from Weipa, 12°23’S, 142°10’E,

Jul 1993, Forster PIF13505 et al. (BRI); Walkers Creek,

Karumba - Normanton road, 17°28’S, 141°10’E, Mar

2001, Holmes [AQ498382] (BRI); cult. Tolga (ex

Stones Crossing, Wenlock River), Dec 1991,

Sankowsky 1370 & Sankowsky (BRI).

Distribution and habitat: Croton rarus is

endemic to western Cape York Peninsula,

Queensland where it has been collected from

near Weipa in the north to Walkers Creek in the

south (Map 4) over five 1° grid squares. Plants

grow in semi-deciduous notophyll vineforest

on alluvium along seasonally flooded

watercourses, or in one instance in deciduous

vinethicket on heavy black clay. Croton rarus

is sympatric with C. mutabilis at some localities.

Notes: Croton rarus is closely allied to both

C. byrnesii from Arnhem Land and C. dockrillii

from the east coast of Cape York Peninsula. A

macromorphological comparison of these three

taxa is made in Table 4.

Forster, Croton in Australia

407

Fig. 22. Croton rarus. A. flowering branchlet. x 0.8. B. undersurface of leaf, x 1. C. base of leaf lamina showing

extrafloral nectaries, x 8. D. node showing stipules, x 8. E. inflorescence with female flowers towards base, male

buds in upper two-thirds, x 1.5. F. male flower, x 8. G. female flower, x 8. H. coccus of dehisced fruit, x 8. I. seed,

x 8. A,B,G from Sankowsky 1370 (BRI); C & F from Clarkson 4061B (BRI); D from Sankowsky 1446 (BRI); E

from Clarkson 4062A (BRI); H & I from Alpha & Black 202B (BRI). Del. W. Smith.

408

Austrobaileya 6 (3): 349-436

Table 4. Morphological comparison of Croton byrnesii , C. dockrillii and C. rarus

Character

C. byrnesii

C. dockrillii

C. rarus

No. lateral veins

in leaf lamina

11-13

10-11

12-14

leaf lamina margin

crenate

denticulate

to sinuate

crenate

indumentum colour

ferruginous-

yellow

clear

ferruginous- silver

foliar glands

sessile to stipitate

stipitate

sessile

male pedicel length

2.5-7 mm

2-3 mm

1-2 mm

styles divided

twice

once

once

fruit shape

depressed-

globose

globose

globose

Conservation status: Croton rarus is an

uncommonly collected plant but is likely to be

more widespread than the available collections

would indicate. There are no immediate threats

to this species and no conservation coding is

thought necessary. The species has been

recorded from Rokeby National Park.

Etymology: The specific epithet is derived from

the Latin rarus (scattered or rare) and alludes

to the distribution and apparent paucity of this

species.

23. Croton schultzii Benth., FI. Austral. 6: 124

(1873). Type: Northern Territory. Port

Darwin, June 1870, Schultz 609 (holo: K

n.v., photo atBRI!).

[Croton argyratus auct., non Blume; Brock

(1988:129); Dunlop etal. (1995); Wheeler

(1992:597)].

Illustrations : Brock (1988:129); Dunlop et al.

(1995:214, Fig. 71); Wheeler (1992:598,

Fig. 182A).

Shrub to 4 m high, monoecious, seasonally

deciduous, perennial. Indumentum silver.

Branchlets ± rounded, with dense overlapping

peltate scales when young, glabrescent.

Stipules linear-lanceolate, 3-6 mm long, 0.3-mm

wide, entire and with dense peltate scales.

Leaves alternate, discolorous, petiolate; petioles

20-60 mm long, 0.8-1 mm wide, with dense

peltate scales; lamina elliptic to broadly ovate,

20-120 mm long, 15-100 mm wide, palminerved

with 1 or 2 veins per side from base and with 6-

10 lateral veins per side of midrib, tertiary

reticulate veins obscure; upper surface dark

green, lateral veins not visible, ± glabrous or

with scattered peltate scales, glabrescent; lower

surface irridescent silver, lateral veins weakly

visible, with dense, overlapping, peltate scales,

neither scabrid nor velutinous; margins entire

or weakly sinuate, foliar glands inconspicuous;

tip acute to short acuminate; base cordate;

extrafloral nectaries absent, or if present then 2

at base of lamina, sessile, ellipsoid, 0.4-0.6 mm

long, 0.2-0.5 mm wide, visible below only.

Inflorescence up to 70 mm long, unbranched,

usually androgynous, pedunculate up to 10 mm;

axis with dense peltate scales; bracts lanceolate,

1-2.8 mm long, 0.2-0.8 mm wide, with dense

peltate scales. Male flowers 3-4 mm long, 3-4

mm diameter, densely clustered towards the

inflorescence tip; pedicels 2-4 mm long, 0.4-0.5

mm wide, with dense peltate scales; sepals

valvate, 5, lanceolate to lanceolate-ovate, 2.5-

2.8 mm long, 1-1.6 mm wide, with dense peltate

scales; petals 5, lanceolate, 2.3-4 mm long, 0.8-

1.5 mm wide, lanate in upper half and with sparse

simple hairs on abaxial surface; stamens 10,

filaments + flattened, 2-3 mm long, c. 0.2 mm

Forster, Croton in Australia

409

Fig. 23. Croton schultzii. A. fruiting branchlet. x 0.4. B. undersurface of leaf, x 1. C. base of leaf lamina howing

extrafloral nectaries, x 8. D. node showing stipule, x 6. E. inflorescence with male flowers, x 2. F. male flower, x 6.

G. female flower, x 6. H & I. Fruit, x 4. J. seed, x 4. A,B,J from Brock 209 (BRI); C, H & I from Cowie 8781 (BRI);

D & G from Leach 2695 (BRI); E & F from Russell-Smith 8123 (BRI). Del. W. Smith.

410

wide, glabrous; anthers oblong, 0.8-1 mm long,

0.5-0.7 mm wide. Female flowers 3^1.5 mm long,

3-4 mm diameter, held singly 5-15 mm apart;

pedicels 2-8 mm long, 0.8-1 mm diameter, with

dense peltate scales; sepals valvate, 5,

lanceolate-obovate, 3-5.5 mm long, 1.5-2.5 mm

wide, with dense peltate scales; petals absent;

styles 3, linear to obloid, 2.6-4 mm long, bifid

for 1.5-2 mm long, connate at base for c. 0.3

mm, glabrous apart from sparse peltate scales

at base; ovary 3-locular, 2-3 mm long, 2-3 mm

diameter, with dense peltate scales. Fruits

trilobate, globose, 6-8 mm long, 7-8 mm

diameter, with sparse peltate scales. Seeds +

obloid, c. 5.5 mm long, 4-4.8 mm wide, 2-2.8 mm

thick, irregularly blotched brown and cream,

ventral surface bifacial, dorsal surface rounded,

micropylar ridge 4-4.8 mm long; caruncle

crescent shaped, c. 1 mm long, 2.3-2.5 mm wide,

cream. Fig. 23.

Selected additional specimens: Western Australia.

Near Cape Bernier, 14°06’S, 127°32’E, Jun 1988,

Hyland 13538 (QRS). Northern Territory. East Point,

Feb 1987, Brock 209 (BRI, DNA); East Point, 12°25’S,

130°49’E, Nov 1967, Byrnes 284 (DNA); Gunn Point,

Nov 1990, Cowie 1444 & Dunlop (BRI, DNA, MEL);

Gunn Point, 12°10’S, 131°02’E, May 1984, Dunlop

6706 & Wightman (DNA); East Point, Dec 1990,

Dunlop 8781 & Cowie (BRI, DNA); Gunn Point, Nov

1989, Forster PIF5918 & Russell-Smith (BRI, DNA,

MEL); Mt Briggs, Fish River Station, Mar 1989, Leach

2512 & Dunlop (BRI, DNA); Gunn Point, Feb 1990,

Leach 2695 & Dunlop (BRI, CANB, DNA); Lee Point,

Darwin, 12°26’S, 131°50’E, Aug 1984, Russell-Smith

1147 (DNA); Murganella, Wunya Beach, 11°42’S,

133°09’E, Mar 1987, Russell-Smith 1957 & Lucas

(DNA); Melville Island, Condor Point, 11°44’S,

131°17’E, May 1987, Russell-Smith 2390 & Lucas

(DNA); 3 km W Mt Muriel, Tipperary, 13°54’S,

131°10’E, Mar 1989, Russell-Smith 7981 & Lucas

(BRI, DNA); Gunn Point, Nov 1989, Russell-Smith

8123 & Lucas (BRI, DNA).

Distribution and habitat : Croton schultzii is

endemic to Australia occuring in a few localities

near Darwin in the Northern Territory and the

Kimberley region in Western Australia (Map 3).

Its distribution covers nine 1° grid squares.

Plants grow in deciduous vinethicket on red

laterite or limestone.

Phenology: Flowering occurs from November

to January, following storm rain. There is one

record from June, but this is considered

abnormal. Fruiting occurs from December to

March.

Austrobaileya 6 (3): 349-436

Notes: Croton schultzii has been referred to in

recent times as C. argyratus (Wheeler 1992;

Dunlop etal. 1995; Chakrabarty & Balakrishnan

1997). It is not conspecific with that species

and differs in the peltate scales on the foliage

(versus peltate trichomes), the male flowers with

shorter pedicels (2-4 mm versus 5-6 mm), the

smaller fruit (6-8 mm diameter versus 12-16 mm)

and smaller seeds (c. 5.5 mm long x 4-4.8 mm

wide, versus 10-11 mm long x 7-8 mm wide).

C. argyratus is restricted to peninsular

Thailand, the Malay Peninsula, Sumatra, Java,

Borneo and the Lesser Sunda Islands (H.-J.Esser

pers. comm. 2000), although it has also been

recorded for India in the Andaman and Nicobar

Islands (Chakrabarty & Balakrishnan 1997).

Conservation status : The species is common

at the known localities.

Etymology: The name honours M. Schultz (date

of birth & death unknown), who collected

specimens at Port Darwin as cited by Bentham

in the ‘Flora Australiensisk

24. Croton setigerus Hook., FI. Bor.-Amer. 2:

141 (1838); Eremocarpus setigerus

(Hook.) Benth., Bot. Voy. Sulph. 53, t. 26

(1844). Type: U.S.A., California, Douglas

(holo: K n.v., photo at BRI!).

Illustrations: James & Harden (1990: 420);

Webster (1993b: 575); Jeanes (1999: 67,

Fig. 10b); Radcliffe-Smith (2001: 325, Fig.

40).

Prostrate to semi-erect herb to 20 cm high,

monoecious, annual. Indumentum ferruginous-

silver. Branchlets rounded, with dense stellate

trichomes. Stipules linear-subulate, 5-7 mm

long, 0.2-0.5 mm wide, multifid and with dense

stellate trichomes. Leaves alternate, petiolate,

discolorous; petioles 3-45 mm long, c. 1 mm

wide, with dense stellate trichomes; lamina

ovate to suborbicular, 8-65 mm long, 8-40 mm

wide, palminerved with 3-5 lateral veins from

base, tertiary reticulate veins obscure; upper

surface silver-green, venation obscure, with

dense stellate trichomes; lower surface silver,

venation obscure, with dense stellate trichomes;

margins entire, foliar glands absent; tip acute

to rounded; base cuneate to rounded; extrafloral

nectaries absent. Inflorescence up to 15 mm

long, unbranched, androgynous, ± sessile; axis

Forster, Croton in Australia

411

Fig. 24. Croton setigerus. A. habit, x 1. B. stellate trichome. x 80. C. male flower, x 20. D. stamens, x 20. E. fruit,

x 13.5. F. fruit, dehisced, x 13.5. G. seed, x 13.5. All from Rose 41457 (K). Del. C. Speight. Plate reproduced with

permission from Radcliffe-Smith (2001: 325).

412

with dense stellate trichomes; bracts linear, 4-5

mm long, c. 0.1 mm wide, with dense stellate

trichomes. Male flowers 3-4 mm long, 2.8-3.2

mm diameter, held singly, usually densely

clustered towards top of inflorescence; pedicels

1.8^1 mm long, c. 0.1 mm wide, with sparse sessile

trichomes; sepals valvate, 5, obovate, 2-2.5 mm

long, 0.9-1 mm wide, with dense stellate

trichomes; petals absent; stamens 5-7, filaments

f ilif orm, 2-3.8 mm long, c. 0.1 mm wide, glabrous,

anthers oblong, 0.8-1 mm long, 0.3-0.4 mm wide.

Female flowers 6-7 mm long, c. 2 mm diameter,

held singly and spaced up to 5 mm apart, sessile;

sepals absent; petals absent; styles 1, linear-

subulate, 3.5-5 mm long, entire, with sparse

stellate trichomes; ovary 1-locular, c. 1.5 mm

long and 0.2 mm diameter, with sparse stellate

trichomes. Fruits unlobed, oblong, 4-6 mm long,

3-3.5 mm diameter, with dense, sessile stellate

trichomes. Seeds ellipsoid, 3-5.2 mm long, 1.7-

3 mm wide, 1.4-2.2 mm thick, cream and tan-

brown mottled, ventral surface bifacial, dorsal

surface rounded, micropylar ridge 2.5-4 mm

long; ecarunculate. Fig. 24.

Additional specimens'. United States of America.

California. Near Oroville, Butte County, Jul 1937,

Copeland 1604 (BRI); Santa Cruz Island, Pelican Bay,

Jul 1930, Clokey 4993 (BRI); Mandeville Canyon,

Santa Monica Mountains, Los Angeles County, Aug

1929, Clokey 4599 & Templeton (BRI); Tenaja Ranger

Station area, Rancho California, 7 miles NW of

Murietta, southwest Riverside County, Oct 1971, Grove

8 (BRI); Saratoga summit, Santa Clara County, Sep

1941, Rose 41457 (BRI). Australia. New South

Wales. Trangie district, Feb 1939, Glenfield Vet.

Research Station (NSW); Trangie district, Feb 1942,

s.coll. [AQ207216] (BRI); Trangie Expt. farm, Jan

1943, May s.n. (NSW); near Corowa, Apr 1965,

Mulham s.n. (NSW); Shire of Corowa, Apr 1969,

Rodway (MEL601784). Victoria. Granya, 36°07’S,

147°19’E, Apr 1984, Roberts (MEL662901).

Distribution and habitat : Croton setigerus is

native to California, U.S.A. and is naturalised in

New South Wales and Victoria in agricultural

areas where it is a weed of cultivated ground

(Map 9). Jeanes (1999) mentioned that it was

also naturalised in Western Australia and South

Australia, but I have not seen specimens from

those states.

Notes : Croton setigerus has been usually

placed in the monotypic genus Eremocarpus,

most recently by Radcliffe-Smith (2001).

Webster (1993a, 1994) has advocated the

reduction of Eremocarpus to a monotypic

Austrobaileya 6 (3): 349-436

section of Croton and this was followed by

Radcliffe-Smith & Govaerts (1997), but at

subgeneric level. Croton setigerus is very

different from the species of Croton familiar to

me from Australia and adjacent regions, primarily

in the 1-locular female flower (thought to be an

adaptation to wind pollination by Webster

1993a) and fruit and the ecarunculate seed.

Webster’s sectional classification requires a

rigorous examination utilizing both

morphological and molecular data, much beyond

the scope of the current work, hence his

reduction of Eremocarpus is followed here,

although with reservations.

Phenology: In Australia, flowering occurs from

October to January, fruiting from January to

April.

Etymology: The specific name is derived from

the Latin seta (bristle) and -ger (carrying or

bearing) and probably alludes to the dense

stellate indumentum of this plant.

25. Croton simulans RI.Forst. sp. nov., affinis

C. capitis-york autem lamina foliorum

venis lateralibus leniter (vice valde)

effectis et squamis densis superpositus

peltatis mixtis trichomatis stellatis remotis

usque sparsis (vice non nisi squamis

peltatis sparsis usque densis) subtus

praeditis, nectariis extrafloralibus basi

laminae foliorum dispositis et tantum

subter videndis (vice mode infra basin

laminae dispositis et videndis et supra et

subter), floribus masculinis pedicellis

longioribus (5-6 mm vice 1.4-1.6 mm)

petalis longioribus (1.8-2 mm vice 1.4-1.5

mm) differt. Typus: Queensland. Cook

District: Timber Reserve 14, Parish of

Kesteven, 28 November 1991, B. Hyland

14377 (holo: QRS; iso: BRI).

Shrub to 5 m high, monoecious, deciduous,

perennial. Indumentum silver. Branchlets

rounded, with dense peltate scales and

scattered stellate trichomes, glabrescents.

Stipules minute, triangular, <0.3 mm long and

0.3 mm wide, with scattered peltate scales.

Leaves discolorous, petiolate; petioles 3-10 mm

long, c. 1 mm wide, with dense peltate scales

and stellate trichomes when young,

glabrescent; lamina elliptic-ovate to

oblanceolate, chartaceous, 10-150 mm long,

Forster, Croton in Australia

413

Fig. 25. Croton simulans. A. branchlet. x 0.5. B. undersurface of leaf, x 1. C. base of leaf lamina showing

extrafloral nectaries, x 6. D. node showing stipule, x 6. E. inflorescence with female flowers towards base and male

flowers in upper two-thirds, x 1. F. male flower, x 6. G. female flower, x 6. A-C from Forster PIF24072 (BRI);

D-Gfrom Hyland 25813 (BRI). Del. W. Smith.

414

4-60 mm wide, penninerved with 8-12 lateral

veins per side of midrib, tertiary reticulate veins

obscure; upper surface grey-green, lateral veins

indistinct, with sparse to dense (overlapping)

peltate scales, glabresent; lower surface silver

to silver-green, lateral veins weakly developed,

with sparse to dense (overlapping) peltate

scales & scattered stellate trichomes, neither

scabrid nor velutinous; margins entire or weakly

sinuate, foliar glands prominent; tip obtuse,

acute to short-acuminate; base cuneate to

cordate; extrafloral nectaries 2, at leaf lamina

base, sessile or stipitate to 1.2 mm long,

ellipsoid, 0.3-0.4 mm long, 0.2-0.3 mm wide,

visible only from below. Inflorescence up to 10

mm long, unbranched, unisexual or

androgynous, pedunculate up to 10 mm; axis

with dense peltate scales & stellate trichomes;

bracts lanceolate, c. 1 mm long and 0.7 mm wide,

with dense peltate scales. Male flowers c. 3 mm

long, 4-5 mm diameter, clustered on

inflorescence in glomerules of 1-3 flowers, or

spaced to 5 mm apart; pedicels 5-6 mm long, c.

0.5 mm wide, with dense peltate scales & peltate

trichomes; sepals valvate, 5, lanceolate-ovate,

1.8-2 mm long, 1-1.2 mm wide, lanate in upper

half; petals 5, lanceolate-ovate, 1.8-2 mm long,

0.5-0.8 mm wide, lanate in upper half; stamens

10-12, filaments filiform, 2-2.2 mm long, c. 0.1

mm wide, glabrous, anthers oblong, 0.8-1 mm

long, 0.4-0.5 mm wide. Female flowers 2-2.5 mm

long, 2.5-3 mm diameter, held singly and spaced

up to 10 mm apart; pedicels 4-6 mm long, c.

1 mm diameter, with dense peltate scales and

peltate trichomes; sepals valvate, 5, lanceolate-

ovate, 1.8-2 mm long, 1.5-1.7 mm wide, with

dense peltate scales; petals absent; styles 3,

obloid, up to 1.2 mm long and 0.2 mm wide, bifid

for up to 0.7 mm long, glabrous; ovary 3-locular,

1.8-2 mm long, 1.8-2 mm diameter, with dense

ferruginous, stellate trichomes. Fruits and seeds

not seen. Fig. 25.

Additional specimens : Queensland. Cook District:

T.R.14, Parish of Kesteven, Nov 1991, Hyland 25813

RFK (BRI, QRS). Cultivated: Station road, Ipswich by

M.C.Tucker (ex plant collected on Leo Creek road,

eastern fall of Mcllwraith Range), Mar 1999, Forster

PIF24072 (BRI).

Distribution & habitat: Croton simulans is

thus far known from only along the Leo Creek

road in the western side of the Mcllwraith Range

on Cape York Peninsula (Map 3). Plants grow

as understorey shrubs in araucarian microphyll

vineforest on granite.

Austrobaileya 6 (3): 349-436

Phenology: Flowers have been recorded in

November. Fruit would be expected between

January and February.

Notes: Croton simulans is superficially similar

to C. capitis-york and may well be a sister-taxon

to that species but is immediately

distinguishable in the field by the more silver

appearance of the seasonally deciduous foliage.

Croton simulans differs from C. capitis-york in

the undersurface of the leaf lamina having

weakly developed lateral veins (versus strongly

developed), a mixture of dense (overlapping)

peltate scales and scattered to sparse stellate

trichomes (versus sparse to dense peltate scales

only); the extrafloral nectaries situated at the

base of the lamina and visible only from below

(versus situated just below the lamina base and

visible both from above and below); the male

flowers with longer pedicels (5-6 mm versus

1.4-1.5 mm) and longer petals (1.8-2 mm versus

1.4-1.5 mm).

Etymology: The specific epithet simulans

(imitating or resembling) is formed directly from

Latin and refers to the superficial similarity of

this species to Croton capitis-york.

26. Croton stigmatosus F.Muell., Fragm. 4:140

(1864); C. phebalioides var. stigmatosus

(F.Muell.) Domin, Biblioth. Bot. 89: 880

(1928). Type: Queensland. Moreton

District: Moreton Bay, 1845, Leichhardt

[sheet with female specimen] (lecto [here

chosen]: P n.v., photo at BRI!).

Croton stigmatosus Muell.Arg., Linnaea 34:

107 (1865), nom. illeg.’, Oxydectes

stigmatosus (Muell.Arg.) Kuntze, Rev.

Gen. PI. 2:613 (1891). Type: Queensland.

Moreton District: Moreton Bay, 1845,

Leichhardt [2 sheets seen, one is the

lectotype sheet of C. stigmatosus

F.Muell.] (syn: P n.v., photo at BRI!); New

South Wales. Clarence River, [ Beckler\ F.

Mueller (syn: G-DC n.v., fiche at BRI!;

isosyn: P n.v., photo at BRI!).

Croton phebalioides var. hirsuta F.M.Bail.,

Queensland FI. 5: 1436 (1902). Type:

Queensland. Moreton District: Taylor’s

Range, near Brisbane, Bailey (holo: n.v.

at BRI, presumed lost).

Illustrations: Floyd (1989: 143); James &

Harden (1990:420); Hauser (1992:89).

Forster, Croton in Australia

415

Fig. 26. Croton stigmatosus A. fruiting branchlet. x 0.6. B. undersurface of leaf, x 1. C. base of leaf lamina showing

extrafloral nectaries, x 8. D. node showing stipule, x 8. E. inflorescence with female and male flowers, x 3. F. male

flower, x 8. G. female flower, x 6. H & I fruit, x 3. J. mamillate process with stellate hairs, x 20. K. seed, x 6. All

from Forster PIF28042 (BRI). Del. W. Smith.

416

Shrub or small tree to 18 m high, monoecious,

evergreen, perennial. Bark lenticellate, blaze

flesh-yellow, wood cream-yellow. Indumentum

silver. Branchlets ± rounded, with dense stellate

trichomes when young, glabrescent. Stipules

subulate, 1.5-7 mm long, c. 0.5 mm wide, entire

and with dense stellate trichomes. Leaves

alternate, discolorous, petiolate; petioles 3-25

mm long, 1-1.2 mm wide, with dense peltate

scales and stellate trichomes; lamina cuneate-

obovate, elliptic or ovate, 13-160 mm long, 5-

60 mm wide, penninerved with 12-15 lateral

veins per side of midrib, tertiary reticulate veins

obscure; upper surface matt dark-green, lateral

veins weakly visible, with sparse stellate

trichomes and sparse peltate scales; lower

surface silver-white, lateral veins prominent,

with dense stellate trichomes and dense peltate

scales, neither scabrid nor velutinous; margins

denticulate with 14-24 small teeth up to 0.5 mm

long, foliar glands prominent; tip acute to

acuminate; base cordate, cuneate, retuse;

extrafloral nectaries 2 at base of leaf lamina,

stipitate to 1.8 mm long, circular, 0.4-0.5 mm

long, 0.4-0.5 mm wide, visible above and below.

Inflorescence up to 180 mm long, unbranched,

usually bisexual and androgynous but

occasionally with male and female flowers mixed

in same glomerules, pedunculate up to 32 mm;

axis with dense stellate trichomes and dense

peltate scales; bracts linear-lanceolate, 1.5-2 mm

long, 0.3-0.5 mm wide, with dense stellate

trichomes. Male flowers 2-3.5 mm long, 2.5-5

mm diameter, clustered towards top of

inflorescence in glomerules of 2-7 flowers or

held singly and spaced up to 8 mm apart;

pedicels 2-4 mm long, 0.4-0.5 mm wide, with

dense stellate trichomes; sepals valvate, 5,

lanceolate-ovate, ovate or obovate, 1.5-2.2 mm

long, 0.4-1.4 mm wide, with dense stellate

trichomes; petals 5, obovate, 1.5-2.2 mm long,

0.3-0.8 mm wide, lanate in upper half; stamens

10-12, filaments ± filif orm, 1.5-2 mm long, c. 0.1

mm wide, with dense simple trichomes at base,

anthers oblong, 0.6-0.8 mm long, 0.5-0.7 mm

wide. Female flowers 2.5-5 mm long, 3-8 mm

diameter, usually held singly and spaced up to

7 mm apart; pedicels 5-8 mm long, c. 1 mm

diameter, with dense stellate trichomes and

dense peltate scales; sepals valvate, 5, obovate

to ovate, 2.8-4 mm long, 1.5-3 mm wide, with

dense stellate trichomes; petals absent; styles

3, linear, 1.5-3.5 mm long, multifid, twice divided

Austrobaileya 6 (3): 349-436

for 1.3-3.2 mm long, connate at base for c. 0.2

mm, papillose and with sparse stellate trichomes

on proximal two-thirds; ovary 3-locular, 2-3 mm

long, 2.5-3.5 mm diameter, with dense, stalked

stellate trichomes. Fruits trilobate, depressed-

globose, 7-8 mm long, 7-10 mm diameter, with

dense, stalked stellate trichomes on fleshy

mamillate processes. Seeds ± ovoid, 5.5-6.5 mm

long, 3.8-5 mm wide, 2.4-3.2 mm thick, black-

brown, ventral surface bifacial, dorsal surface

rounded, micropylar ridge 3.5-4 mm long;

caruncle oblong-rectangular, 1-1.5 mm long,

1.3-2.5 mm wide, cream. Fig. 26.

Selected additional specimens: Queensland. Port

Curtis District: T.R. 353, W of Many Peaks, 24°32’S,

151°16’E, Nov 1995, Bean 9140 & Turpin (BRI);

Shoalwater Bay, small island in archipelago in Pearl

Bay, 22°25’S, 150°43’E, Jun 1999, Brushe JB2006 &

Plumb (BRI); Pine Creek, S.F. 391 Bulburin, off lower

reaches of Granite Creek, 24°37’S, 151°33’E, Dec 1994,

Forster PIF16029 et al. (BRI); S.F. 67 Bulburin, May

1985, Gibson 734 (BRI, NSW). Wide Bay District: S.F.

234, SW of Cooroy, 26°29’S, 152°49’E, Apr 1993,

Bean 5903 (BRI); Imbil, Mar 1918, Weatherhead

[AQ202204] (BRI). Darling Downs District: Gully W

of Swan Creek, 7.5 km NE of Swanfels & 11 km SW of

Cunningham’s Gap, 28°07’S, 152°19’E, Jan 1989, Bird

[AQ455783] (BRI). Moreton District: Slopes of Mt

Chingee, S of Rathdowney, 28°18’S, 152°26’E, Nov

2001, Bean 18015 (BRI, CANB, MEE, NSW); Rosen’s

Lookout, Beechmont, Mar 1977, Elsol [AQ194906]

(BRI); CSR Land, Ormeau, 27°47’S, 153°13’E, Dec

2001, Forster PIF28042 & Leiper (A, BISH, BRI, DNA,

L, MEL, MO, NSW, WIS, Z); Upper Brookfield,

Brisbane, Feb 1978, Jessup 43 (BRI); Beechmont Ridge,

Beechmont, Macpherson Range, Oct 1969, Schodde

5592 (BRI, CANB, MEL); Sunday Creek, Lamington

N.P., Smith [AQ379568] (BRI); Blackall Range, Dec

1916, White [AQ202197] (BRI); Tamborine Mt, May

1940, White 11440 (BRI). New South Wales.

Lismore, Mar 1898, Baker [MEL231575] (MEL);

Wiangaree S.F., Jan 1981, Bird [AQ345018] (BRI); 23

km NW of Kyogle, Toonumbar forest road, Toonumbar

S.F., 28°29’S, 152°48’E, Dec 1991, Halford Q824 (BRI,

MEL, NSW); c. 2 miles [3.3 km] SW of Wiangaree,

Oct 1966, Hayes 2558 et al. (BRI); Palm Gully forest

road. Long Creek, between Roseberry & Queensland

border, 28°23’S, 152°56’E, Apr 1981, Jessup 323 (BRI).

Distribution and habitat : Croton stigmatosus

is restricted to south-eastern Queensland from

Shoalwater Bay in the north to the north-eastern

comer of New South Wales (Map 2). It has been

recorded from eleven 1° grid squares. Plants

grow in complex notophyll vineforest that is

often dominated by Argyrodendron spp. on

volcanic soils.

Phenology: Flowering occurs from September

to May; fruiting occurs from September to May.

Forster, Croton in Australia

Notes: F. Mueller (1864) cited four syntypes in

the protologue for Croton stigmatosus , viz. “Ad

flumen Richmond River, Dr Beckler; ad sinum

Moreton Bay, Dr Leichhardt, F.M.; ad sinum

Broad sound, Bowman; ad flumen Fitzroy River

et montem Mueller Australiae orientalis tropicae,

Dallachy”. At P there are two sheets with

specimens collected by Leichhardt from

Moreton Bay, both apparently in 1845. One of

these has buds, the other has female flowers.

There are also sheets with collections

apparently by F. Mueller (no locality) and

Beckler ‘Clarence River’. There is a Bowman

collection from Broad Sound at

MEL (MEL231578). This last collection does

represent Croton stigmatosus , but the present

day Broad Sound is well to the north of where

the species is known to occur. I have been

unable to locate the collections by Dallachy or

Beckler (Richmond River). The name Croton

stigmatosus F.Muell. is lectotypified with the

Leichhardt collection at P that has female

flowers.

To further complicate the nomenclature

of this species, the name Croton stigmatosus

was independently used by J. Mueller (1865)

with the protologue stating “In Nova Hollandiae

orientali ad Clarence River (F.Muell.! in hb.DC.),

in Moreton-Bay (Leichhard! inhb.Mus.Paris)”.

The Leichhardt collection is the same one that

I have used to lectotypify the name

C. stigmatosus F.Muell. The Clarence River

collection appears to have been made by Beckler

and is present at both G-DC and P.

Croton stigmatosus is sometimes

confused with large leaved forms of

C. phebalioides. This latter species grows in

drier closed-forest communities and differs most

noticeably in the ± entire to weakly denticulate

foliage with poorly developed lateral venation.

Conservation status : Croton stigmatosus is

infrequently collected but is not uncommon

throughout its known range. It is present in at

least three conservation reserves in south¬

eastern Queensland and two in New South

Wales (Floyd 1989). No conservation coding is

necessary.

Etymology : The specific epithet is derived from

the Latin stigmatosus and means having well

developed stigmas.

417

27. Croton stockeri (Airy Shaw) Airy Shaw,

Kew Bull. 35: 622 (1981); Croton wassi-

kussae Croizat var. stockeri Airy Shaw,

Muelleria4:229 (1980). Type: Queensland.

Cook District: between Rocky River

and Massy Creek, 13°40’S, 143°25’E,

13 September 1973, G.C. Stocker 1076

(holo: QRS; iso: BRI, CANB).

Shrub to 4m high, monoecious, evergreen or

seasonally deciduous, perennial. Indumentum

orange-brown. Branchlets rounded, with dense

stellate trichomes. Stipules lanceolate, 3-6 mm

long, 1-1.8 mm wide, entire and with dense

stellate trichomes. Leaves alternate, discolorous,

petiolate; petioles 5-12 mm long, c. 1 mm wide,

with dense stellate trichomes; lamina ovate to

elliptic, 12-80 mm long, 10-50 mm wide,

penninerved with 9-11 lateral veins per side of

midrib, tertiary reticulate veins obscure; upper

surface matt green, venation not visible, with

sparse stellate trichomes; lower surface orange-

brown, lateral veins weakly prominent, with

dense, stellate trichomes, velutinous; margins

sinuate to denticulate with 8-30 teeth up to 0.3

mm long, foliar glands prominent; tip acute to

rounded; base cordate; extrafloral nectaries 2

at lamina base, sessile, ellipsoid, 0.5-0.9 mm

long, 0.3-0.5 mm wide, visible only below.

Inflorescence up to 80 mm long, unbranched,

androgynous, pedunculate up to 5 mm; axis with

dense stellate trichomes; bracts linear-

lanceolate, 1.7-2 mm long, c. 0.4 mm wide, with

dense stellate trichomes. Male flowers c. 2 mm

long and 3 mm diameter, densely clustered in

glomerules of 4-6 flowers towards the top of

the inflorescence; pedicels c. 3.5 mm long and

0.2 mm wide, with dense stellate trichomes;

sepals valvate, 5, lanceolate-ovate, c. 1.8 mm

long and 1 mm wide, with dense stellate

trichomes; petals 5, oblanceolate, 2-2.2 mm long,

0.4-0.5 mm wide, lanate in upper half; stamens

11-12, filaments filif orm, 1.8-2.1 mm long, c. 0.2

mm wide, glabrous; anthers oblong, c. 0.8 mm

long and 0.5 mm wide. Female flowers 4.5-5 mm

long, 4.5-5 mm diameter, often mixed in same

glomerule as males or held singly and spaced

up to 5 mm apart; pedicels c. 3 mm long and 1

mm diameter, with dense stellate trichomes;

sepals valvate, 5, lanceolate-ovate, c. 2.8 mm

long and 1.5 mm wide, with dense stellate

trichomes; petals absent; styles 3, linear, 2.5-3

mm long, bifid for 1-1.5 mm, glabrous, connate

418

Austrobaileya 6 (3): 349-436

Fig. 27. Croton stockeri. A. budding branchlet. x 0.8. B. undersurface of leaf, x 1. C. base of leaf lamina showing

extrafloral nectaries, x 4. D. node showing stipule, x 6. E. female flower, x 6. F. male flower, x 6. A & D from

Clarkson 3631 (BRI); B,C,F from Tucker s.n. (BRI); E from Stocker 1076 (BRI). Del. W. Smith.

Forster, Croton in Australia

at base for c. 0.2 mm; ovary 3-locular, 3-3.8 mm

long, c. 4 mm diameter, with dense stellate

trichomes. Fruits and seed not seen. Fig. 27.

Additional specimens: Queensland. Cook District: c.

10 km N of upper crossing of Massy Creek N of Silver

Plains on eastern fall of Mcllwraith Range, 13°51’S,

145°28’E, Aug 1978, Clarkson 2450 (BRI, CANB);

3 km N of Upper crossing of Massy Creek on Silver

Plains Station, 13°53’S, 143°31’E, Nov 1980, Clarkson

3631 (BRI, QRS); Scrubby Creek, between the Rocky

& the Chester River, Silver Plains Station, 13°46’S,

143°30’E, Dec 1990, Fell DGF2285 (QRS); 6 km W

of the Rocky River mouth, Silver Plains Holding,

13°46’S, 143°29’E, Aug 1993, Fell DGF3484 et al.

(BRI, DNA); 4 km NNE of Massy Creek Crossing,

Silver Plains Station, 13°53’S, 143°30’E, Jun 1992,

Forster PIF10567 et al. (BRI, DNA, MEF, QRS);

3 km N of Massy Creek Crossing, Silver Plains Station,

13°53’S, 143°31’E, Jun 1992, Forster PIF10580 et al.

(A, BRI, DNA, K, F, MEF, QRS); Silver Plains, S of

Scrubby Creek & W of Colmer Point, 13°46’S,

143°29’E, Jun 1995, Forster PIF17042 (BRI, MEF,

QRS); Cultivated (ex Silver Plains Station, same site as

Forster PIF17042), Jan 2002, Tucker s.n. (BRI).

Distribution and habitat : Croton stockeri is

endemic to Australia and is restricted to a single

1° grid square along with nine other species. All

known populations occur on Silver Plains

Station on the eastern fall of the Mcllwraith

Range on far northern Cape York Peninsula

(Map 3). Plants grow in deciduous vinethicket

on stabilised white sand dunes.

Phenology : Flowers have been collected only

once, in September. It may be presumed that

the main flowering period is from September to

January with fruiting two to three months later.

Buds are retained on the plants for much of the

remaining year.

Notes: Croton stockeri is a distinctive Croton

because of the heavily velutinous foliage

covered in orange-brown stellate trichomes.

Conservation status: The few known

populations of Croton stockeri have no

conservation security all being located on

Aboriginal land. This species is currently listed

as Rare under Queensland Government

legislation.

Etymology: The specific epithet honours Geoff

Stocker, plant ecologist, who made the first

collections of this plant while employed by the

CSIRO.

419

28. Croton tomentellus F.Muell., Fragm. 4:141

(1864). Type: Northern Territory. Upper

Victoria River, on rocks, January 1856,

F. Mueller [MEL231573] (holo: MEL).

Croton tomentellus Muell.Arg., Linnaea 34:

108 (1865), nom. illeg.; Oxydectes

tomentella (Muell.Arg.) Kuntze, Rev. Gen.

PI. 2:613 (1891). Type: Northern Territory.

Arnhem’s Land, F. Mueller (holo: G-DC

n.v., fiche at BRI).

Illustrations: Wheeler (1992:598, Fig. 182c);

Dunlop etal. (1995:214, Fig. 71).

Shrub to 4 high, monoecious, seasonally

deciduous, perennial. Indumentum clear.

Branchlets rounded, with sparse to dense

stellate trichomes, glabrescent. Stipules

lanceolate, 1.2-1.8 mm long, 0.3-0.5 mm wide,

entire and with dense stellate trichomes. Leaves

alternate, discolorous, petiolate; petioles 12-

55 mm long, 1-2 mm wide, with dense stellate

trichomes; lamina ovate, 20-250 mm long, 15-

110 mm wide, ± palminerved with 3-5 veins

from the base, 10-14 lateral veins per side of

midrib, and tertiary reticulate veins; upper

surface matt green, venation not visible, with

scattered to dense stellate trichomes; lower

surface silver, lateral and tertiary venation

weakly developed, with dense, velutinous

stellate trichomes, often glabrescent, neither

scabrid nor velutinous; margins denticulate or

weakly crenate with 40-54 teeth up to 0.5 mm

long, foliar glands prominent; tip acute to short

acuminate; base cordate, cuneate or rounded;

extrafloral nectaries 2 at lamina base, sessile,

circular to ellipsoid, 0.4-0.7 mm long, 0.4-0.7

mm wide, visible on both surfaces. Inflorescence

up to 130 mm long, unbranched, androgynous,

pedunculate up to 20 mm; axis with sparse to

dense stellate trichomes; bracts lanceolate, 1-2

mm long, 0.3-0.5 mm wide, with sparse to dense

stellate trichomes. Male flowers 2-4.5 mm long,

3.5-5 mm diameter, densely clustered in

glomerules of 1-5 flowers towards the top of

the inflorescence; pedicels 2-5.2 mm long and

c. 0.5 mm wide, with sparse to dense stellate

trichomes; sepals valvate, 5, lanceolate-ovate,

2-2.3 mm long, 0.8-1.8 mm wide, with sparse

stellate trichomes; petals 5, oblanceolate, 3-3.2

mm long, 0.7-1 mm wide, lanate in upper half;

stamens 11 or 12, filaments filiform, 2-2.5 mm

long, c. 0.2 mm wide, glabrous, anthers oblong,

420

Austrobaileya 6 (3): 349-436

F

IA\|

u

u

Fig. 28. Croton tomentellus. A. flowering branchlet. x 0.4. B. node showing stipule, x 6. C. base of leaf lamina

showing extrafloral nectaries, x 6. D. inflorescence with female flowers towards base, male flowers in upper two-

thirds, some glomerules with both sexes, x 1. E. male flower, x 8. F. female flower, x 8. G & H. fruit x 4.1. seed, x 8.

A,D,E,F from Brock 739 (BRI); B,C,I from Byrnes 1204 (BRI); G,H from Brock 656 (BRI). Del. W. Smith.

Forster, Croton in Australia

0.7-1 mm long, 0.6-0.7 mm wide. Female flowers

2.5-3 mm long, 3^\ mm diameter, often mixed in

same glomerule as males or held singly and

spaced up to 5 mm apart; pedicels 1.2-2.2 mm

long, c. 0.8 mm diameter, with sparse stellate

trichomes; sepals valvate, 5, lanceolate-ovate,

1.8-2.3 mm long, 1-1.8 mm wide, with sparse

stellate trichomes; petals absent; styles 3, linear,

2-2.7 mm long, bifid for 1.2-1.4 mm, glabrous;

ovary 3-locular, 1.5-2.2 mm long 1.5-2.2 mm

diameter, with dense, + sessile stellate trichomes.

Fruits trilobate, globose, 4.5-6 mm long, 5-5.5

mm diameter, with sparse, ± sessile stellate

trichomes. Seed + ovoid, 3.5-4.4 mm long, 3-

3.3 mm wide, 2.3-2.8 mm thick, dorsal surface

rounded, ventral surface bifacial, micropylar line

2-2.8 mm long; caruncle ovate, c. 0.7 mm long

and 0.7 mm wide, cream. Fig. 28.

Selected additional specimens : Western Australia.

Upper Neville Creek in Harding Range, Eastern Walcott

inlet, 16°17’S, 124°59’E, May 1983, Fell DGF32 (BRI,

PERTH); Savion Gorge, 16°30’S, 125°16’E, Feb 1989,

Hyland 13842 (QRS); Mitchell Plateau, 14°33’S,

125°48’E, Dec 1982, Kenneally 8636 (CANB,

PERTH); S of Ninbing Homestead site, N of Kununurra,

Jun 1969, Lullfitz 691102-21 & McKenzie (PERTH);

c. 15 km W of Kalumburu on road to Truscott on Lip

of Poompangala Hill, 14°17’S, 126°34’E, Dec 1992,

Mitchell 2812 (BRI); Walsh Point - Point Warrender,

14°34’S, 125°45’E, May 1981, Tracey 13955 (BRI);

Lone Dingo between Mitchell Plateau Mining Camp &

Point Warrender, 14°35’S, 125°43’E, May 1981, Tracey

13956 (BRI). Northern Territory. Timber Creek,

15°40’S, 130°30’E, Mar 1989, Brock 656 (BRI, DNA);

Mt Bundy, 12°52’S, 131°36’E, Nov 1990, Brock 739

& Russell-Smith (BRI, DNA, CANB, MEL, QRS);

Bullita Station, Gregory N.P., 16°03’S, 130°23’E, Feb

1986, Clark 292 & Wightman (DNA); c. 67 miles [111.7

km] NE of Maranboy Police Station, Mar 1965,

Lazarides & Adams 98 (CANB, MEL); Mt Goyder,

12°52’S, 131°41’E, May 1987, Russell-Smith 2358 &

Lucas (DNA); 4 km W Umbakumba, Groote Eylandt,

13°52’S, 136°47’E, Jul 1987, Russell-Smith 2740 &

Lucas (DNA); Groote Eylandt, 8 km SW Umbakumba,

13°55’S, 136°43’E, Jul 1987, Russell-Smith 2764 &

Lucas (DNA); Groote Eylandt, 6 km S Umbakumba,

13°54’S, 136°49’E, Jul 1987, Russell-Smith 2814 &

Lucas (DNA); Cutta Cutta, Guy Cave, 14°35’S,

132°27’E, Dec 1988, Russell-Smith 6506 & Lucas

(BRI, DNA); 4 km SE Mt Harris, Kakadu, 13°18’S,

131°57’E, Jan 1989, Russell-Smith 6605 & Lucas

(DNA); 20 km S Jasper Gorge, 16°13’S, 130°43’E, Mar

1989, Russell-Smith 7715 (DNA); Headwaters of Big

Horse Creek, 15°43’S, 130°25’E, Mar 1989, Russell-

Smith 7779 (BRI, DNA); Guy Cave area, 16 Mile Cave

Reserve S of Katherine, May 1978, Tracey 14049

(BRI); Malgala, Groote Eylandt, 13°52’S, 132°26’E,

Nov 1976, Waddy 617 (DNA).

Distribution and habitat : Croton tomentellus

is endemic to northern Australia where it occurs

421

in a number of disjunct populations in the

Northern Territory and Western Australia over

a total of fourteen 1° grid squares (Map 9). Plants

grow in deciduous vinethicket on granite,

laterised basalt or sandstone substrates. There

is also one record from open woodland on

basalt.

Phenology : Flowering occurs from November

to March following storm rains; fruiting occurs

from November to April.

Notes: This species was included by Webster

(1993a) in Croton section Croton , but does not

have the characters given for that section, e.g.

11 or 12 stamens (versus 15-35).

Conservation status: Croton tomentellus is

widespread and present in Kakadu and Gregory

National Parks. It is not considered rare or

threatened.

Etymology: The specific epithet is derived from

the Latin tomentellus and means minutely

tomentose, perhaps alluding to the indumentum

on the lower leaf surface or the fruit.

29. Croton triacros F.Muell., Fragm. 6: 185

(1868). Type: Queensland. North Kennedy

District: Rockingham’s Bay, Dallachy

(lecto [herechosen]: MEL231564).

Shrub to 5 m high, monoecious, evergreen,

perennial. Indumentum ferruginous. Branchlets

rounded, with scattered peltate trichomes when

young, glabrescent. Stipules lanceolate, 4-4.5

mm long, c. 0.5 mm wide, entire and with

scattered to sparse peltate trichomes. Leaves

alternate, discolorous, petiolate; petioles 3-36

mm long, c. 1 mm wide, with sparse peltate

trichomes; lamina elliptic, broadly elliptic or

oblanceolate, 75-185 mm long, 40-70 mm wide,

penninerved with 9-11 lateral veins per side of

midrib and tertiary reticulate veins; upper surface

dark green, venation weakly visible, with

scattered peltate trichomes when young; lower

surface pale green, lateral and tertiary venation

weakly prominent, glabrous apart from a few

scattered peltate trichomes, neither scabrid nor

velutinous; margins sinuate or very weakly

denticulate with 20-30 teeth up to 0.3 mm long,

foliar glands prominent; tip acute to short

acuminate; base cuneate; extrafloral nectaries 2

at lamina base sessile, ellipsoid, 0.7-1.3 mm

422

Austrobaileya 6 (3): 349-436

Fig. 29. Croton Macros. A. fruiting branchlet. x 0.4. B. base of leaf lamina showing extrafloral nectaries, x 8. C.

node showing stipules, x 8. D. inflorescence with female flowers in lower half and male flowers in upper half, x 2.

E. female flower, x 12. F. male flower, x 8. G & H. fruits, x 4. I. seed, x 8. A-E from Jago 3060 (BRI); F from

Forster PIF18189 (BRI); G-I from Forster PIF13081 (BRI). Del. W. Smith.

Forster, Croton in Australia

long, 0.4-0.7 mm wide, visible below only.

Inflorescence up to 110 mm long, unbranched,

androgynous, pedunculate up to 10 mm; axis

glabrous or with a few scattered peltate

trichomes; bracts lanceolate, 0.5-1 mm long, 0.2-

0.3 mm wide, glabrous. Male flowers 2-2.5 mm

long, 2-3.5 mm diameter, held singly or in 2-4-

flowered glomerules up to 4 mm apart towards

the top of the inflorescence; pedicels 1.8-4 mm

long and c. 0.2 mm wide, glabrous; sepals

valvate, 5, lanceolate-ovate, 1.6-2 mm long, 0.9-

1.2 mm wide, glabrous, lanate in upper half;

petals 5, oblanceolate, 1.8-2 mm long, 0.6-0.8

mm wide, lanate; stamens 10-11, filaments

filif orm, 1-2 mm long, c. 0.1 mm wide, glabrous,

anthers oblong, 0.4-0.6 mm long, c. 0.3 mm wide.

Female flowers c. 2 mm long and 2 mm diameter,

often mixed in pairs in same glomerule as males

or held singly and spaced up to 13 mm apart;

pedicels 1.5-3 mm long, 0.5-0.6 mm diameter,

with scattered peltate trichomes; sepals valvate,

5, lanceolate, 1.4-2.3 mm long, 0.8-1.3 mm wide,

glabrous, or with scattered peltate trichomes;

petals absent; styles 3, linear, 1.8-2.2 mm long,

bifid for 0.7-1 mm, shortly connate at base,

glabrous; ovary 3-locular, 1.2-2 mm long, 1.2-2

mm diameter, with sparse, + sessile peltate

trichomes. Fruits trilobate, globose, 4.5-7 mm

long, 5-7 mm diameter, with scattered, ± sessile

peltate trichomes. Seed + ovoid to obloid, 3-5

mm long, 2.3^4- mm wide, 1.8-3.5 mm thick, dorsal

surface rounded, ventral surface bifacial,

micropylar line 2.5-3.5 mm long; caruncle ±

crescent shaped, 0.5-0.8 mm long, 0.7-1.2 mm

wide, cream. Fig. 29.

Selected additional specimens: Queensland. Cook

District: c. 15 miles [25 km] NNW of Daintree, 16°04’S,

145°14’E, Nov 1967, Boyland 466 & Gillieatt (BRI,

CANB); T.R. 14, Mcllwraith Range, head of Peach

Creek, 13°44’S, 143°20’E, Dec 1990, Fell DGF2279

(QRS); 36.5 km along road to Leo Creek Mine,

Mcllwraith Range, 13°44’S, 143°20’E, Jun 1992,

Forster PIF10299 & Tucker (BRI, QRS); T.R. 14, Leo

Creek Mine area, Mcllwraith Range, 13°44’S,

143°22’E, Jun 1992, Forster PIF10116 et al. (BRI,

QRS); S.F. 185 Danbulla, 7 km SW of Hoop Pine

Triangle, 17°09’S, 145°35’E, Jan 1993, Forster

PIF13081 & Bean (BRI, MEL, NSW); Pinnacle Track,

2 km W of Karnak, 16°23’S, 145°18’E, Jul 1994,

Forster PIF15529 et al. (BRI, QRS); Tully Falls Weir

road, 17°46’S, 145°33’E, Nov 1995, Forster PIF18199

& Spokes (BRI, MEL, QRS); S.F. 185 Danbulla, Tinaroo

Dam, 17°09’S, 145°33’E, Jan 2002, Forster PIF28160

et al. (A, BRI, L, MEL, NY, WIS); T.R. 9, Lankelly

Creek, 13°55’S, 143°20’E, Sep 1971, Hyland 5382

423

(BRI, QRS); Mt Carter, 13°00’S, 143°15’E, Sep 1974,

Hyland 7532 (BRI, QRS); T.R. 14 Kesteven, 13°43’S,

143°20’E, Oct 1981, Hyland 11138 (QRS); Harvey

Creek, 10 km N of Babinda, 17°50’S, 145°54’E, Jan

1994, Jago 3060 (BRI); Harvey’s Creek, Russell River,

1887, Sayer s.n. (MEL); c. 12.8 km SW of Atherton

on ranges near Moomin, Sep 1950, Smith 4663 (BRI);

Baileys Creek area, on bank of Hutchinson Creek, Oct

1962, Smith 11586 (BRI); McDowall Range, 16°06’S,

145°17’E, Oct 1973, Tracey 14540 (BRI); Lankelly

Creek on western fall of Mcllwraith Range, 13°55’S,

143°15’E, Oct 1969, Webb & Tracey 9625 (BRI); Spear

Creek, Mt Danbulla, 16°43’S, 145°24’E, Oct 1973,

Webb & Tracey 12027 (BRI). North Kennedy District:

S.F. 268 Mt Spec, 3 km along Ewan road past Paluma

Dam turnoff, 19°01’S, 146°08’E, Jan 1992, Forster

PIF9493 (BRI, DNA, K, L, MEL, QRS); Mt Spec near

Bambaroo, Nov 1933, Francis [AQ202219] (BRI).

Distribution and habitat : Croton triacros is

endemic to north-eastern Queensland where it

is disjunct with most populations in the “Wet

Tropics”, and a northern population in the Leo

Creek area of the Mcllwraith Range (Map 7).

The species occurs in six 1° grid squares. Plants

grow in araucarian or complex notophyll

vineforests, usually on granite substrates.

Phenology : Flowering occurs from October to

April following storms or seasonal rains, fruiting

occurs from October to May.

Notes: There are two sheets at MEL (MEL231567

and MEL231564) that are suitable as a lectotype

of Croton triacros. Both are undated and the

labels state that the material originates from

Rockingham’s Bay and was collected by

Dallachy. The first of these is male flowering

material and the latter has female flowers and

fruit. It is unclear whether the two sheets were

collected at the same time, hence the better sheet

(MEL231564) is selected as lectotype of the

name C. triacros.

Conservation status : Common and present in

conservation reserves at Crystal Creek and

Daintree National Parks.

Etymology: Obscure.

30. Croton verreauxii (‘verreauxia’) Baill., Et.

Gen. Euphorb. 357 (1858); Oxydectes

verrauxii (Baill.) Kuntze, Rev. Gen. PI. 2:

613 (1891). Type: New South Wales. Camp

in Heaven, Verreaux 59 (holo: P n.v., photo

at BRI).

Croton verreauxii var. genuinus Muell.Arg.,

Linnaea 34:47 (1865), nom. inval. Type: as

for C. verreauxii Baill.

424

Croton verreauxii var. longifolius Muell.Arg.,

Linnaea 34:47 (1865). Type: New South

Wales. Clarence River, Beckler (holo: B

n.v.).

Illustrations: Williams (1987:83); Floyd (1989:

144); James & Harden (1990:418); Hauser

(1992:181).

Shrub to 5 m high, monoecious, evergreen,

perennial. Indumentum silver. Branchlets +

rounded, with scattered to sparse peltate scales

when young, glabrescent. Stipules linear-

lanceolate, 1.5-2 mm long, c. 0.6 mm wide, entire

and with sparse peltate scales. Leaves alternate,

discolorous, petiolate; petioles 1-36 mm long,

0.8-1 mm wide, with scattered peltate scales

when young, glabrescent; lamina elliptic to

lanceolate, 15-120 mm long, 5-40 mm wide,

penninerved with 12-13 lateral veins per side

of midrib, tertiary reticulate veins obscure to

poorly developed; upper surface glossy dark

green, venation obscure, glabrous; lower

surface pale green, lateral veins poorly

developed, with scattered peltate scales, neither

scabrid nor velutinous; margins denticulate to

weakly crenate with 11-24 teeth up to 0.5 mm

long, foliar glands prominent; tip acute to

acuminate; base cuneate; extrafloral nectaries 2

at lamina base, stipitate up to 0.3 mm long,

circular to ellipsoid, 0.3-0.6 mm long, 0.2-0.4

mm wide, visible from above and below.

Inflorescence up to 170 mm long, androgynous,

pedunculate up to 10 mm; axis glabrous or with

scattered peltate scales; bracts lanceolate to

triangular, 0.6-1 mm long, 0.2-0.3 mm wide,

glabrous or with a few scattered peltate scales

near top. Male flowers 2-2.6 mm long, 1.5-2.2

mm diameter, in clusters of 2 or 3 flowers per

glomerule, spaced up to 5 mm apart; pedicels

2-3.5 mm long, c. 0.2 mm wide, glabrous; sepals

valvate, 5, lanceolate-ovate, 1-1.8 mm long, 0.5-

0.9 mm wide, lanate; petals 5, oblanceolate, 1-

1.8 mm long, 0.4-0.6 mm wide, lanate; stamens

10-12, filaments filiform, 0.6-1 mm long, c. 0.1

mm wide, glabrous, anthers oblong, 0.3-0.6 mm

long, 0.3-0.6 mm wide. Female flowers c. 2 mm

long, 2.5-3 mm diameter, held singly and spaced

up to 7 mm apart; pedicels 1-2 mm long, 0.5-0.7

mm diameter, with scattered peltate scales;

sepals valvate, 5, lanceolate, 1-1.8 mmlong, 0.4-

0.8 mm wide, glabrous or lanate at tip; petals

absent; styles 3, linear, 2-2.8 mm long, bifid for

1-1.8 mm long, connate at base for c. 0.1 mm,

Austrobaileya 6 (3): 349-436

glabrous; ovary 3-locular, 1.4-1.8 mmlong, 1.5-

1.8 mm diameter, with dense, sessile, yellow

peltate scales. Fruits trilobate, depressed-

globose, 4-6 mm long, 5-6.5 mm diameter, with

sparse, sessile, yellow peltate scales. Seeds ±

ovoid, 4-4.5 mm long, 2.9-3.5 mm wide, 2.2-3

mm thick, brown with slight cream mottling,

ventral surface + rounded, dorsal surface

rounded, micropylar ridge 3-4 mm long;

caruncle crescent shaped, 0.5-0.8 mm long, 1-

1.3 mm wide, cream. Fig. 30.

Selected additional specimens’. Queensland. Wide

Bay District: S.F. 234, SW of Cooroy, 26°28’S,

152°49’E, Dec 1993, Bean 7144 (BRI, MEL, MO);

Imbil, S.F. 135, Brooloo, Western L.A., 26°30’S,

152°39’E, Oct 1982, McDonald 3749 & Williams

(BRI, NSW). Darling Downs District: The Head, NE

of Killamey, Dec 1984, Bird [AQ396387] (BRI); The

Head, Main Range N.P., 28°14’S, 152°57’E, Jan 2001,

Forster PIF28064 & Leiper (A, BRI, K, MEL, WIS).

Moreton District: Petrie Creek, W of Woombye,

26°40’S, 152°55’E, Nov 1989, Bean 1188 (BRI);

Upper end of Duck Creek road, near O’Reilly’s,

Lamington Plateau, 28°12’S, 153°07’E, May 2000,

Forster PIF25612 & Booth (A, BRI, L, MEL); The

Ranch, foot of Wilson’s Peak, Nov 1935, Michael

2241 (BRI); Sarabah Range, c. 10 miles [16.7 km] S

of Canungra, Oct 1969, Schodde 5584 (BRI, CANB);

Beechmont Ridge, Beechmont, McPherson Range,

Oct 1969, Schodde 5591 (BRI, CANB); Peecheys

Scrub, Dec 1887, Simmonds [AQ202226](BRI);

Tamborine Mt, Jan 1916, White [AQ202220] (BRI);

Lamington N.P., Dec 1937, White 11401 (BRI). New

South Wales, c. 10 miles [16.7 km] WSW of Dungog

on the road to Gresford, Nov 1970, Blaxell 3357 &

Coveny (BRI); Wyong, Nov 1916, Boorman

[AQ202242](BRI); Border Fence, Moss Gardens,

28°17’S, 152°27’E, Jan 1990, Forster PIF6215 et al.

(BRI, QRS); 23 km NW of Kyogle, Toonumbar Forest

road, Toonumbar S.F., 28°29’S, 152°48’E, Dec 1991,

Halford Q821 (BRI, MEL); Edinburgh Castle, 8 km

SSE of Woodenbong, 28°27’S, 152°38’E, Dec 1992,

Halford Q1561 (BRI); Minyon Falls, Whian Whian,

Sep 1966, Jones [AQ202248] (BRI); Yaamba, Oct

1947, King [AQ202247] (BRI); 22 miles [36.7 km]

NE of Singleton, Mar 1960, Story 7163 (BRI, QRS);

Upper Williams River near Salisbury, Mar 1938, White

11608 (BRI). Victoria. Cult, at “Dunedin”, Tyers

from a cutting from... Back Creek N of Noorinbee,

Apr 1979, Galbraith [MEL1527573] (MEL).

Distribution and habitat : Croton verreauxii

is endemic to eastern Australia where it occurs

from south-eastern Queensland through eastern

New South Wales south to Illawarra (Floyd

1989) over a total of fifteen 1° grid squares

(Map 7). A doubtful locality record in north¬

eastern Victoria (cited above) has been

discounted by Jeanes (1999) who stated that

attempts to relocate the plant in the field have

been unsuccessful. Plants grow in complex

notophyll vineforest and microphyll moss/fern

Forster, Croton in Australia

425

Fig. 30. Croton verreauxii. A. flowering branchlet. x 0.5. B. base of leaf lamina showing extrafloral nectaries, x 8.

C. node showing stipule, x 8. D. inflorescence with female flowers in lower half and male flowers in upper half, x 1.

E. male flower, x 8. F. female flower, x 8. G & H. fruit, x 4. I. seed, x 8. All from Forster PIF25612 (BRI).

Del. W.Smith.

426

thickets on volcanic soils, or may be found in

the ecotonal areas between closed-forest and

open forest dominated by eucalypts.

Phenology : Flowering occurs from August to

March and fruiting occurs from October to April.

Notes: The spelling of the epithet for this species

was originally given as ‘verreauxia’, but as the

species was named after Verreaux, the spelling

should be ‘ verreauxii’.

Croton verreauxii was included in Croton

section Tiglium by Webster (1993a) but does

not agree with character states for this section,

e.g. peltate scales (versus stellate trichomes).

Govaerts et al. (2000) have recently

erroneously labelled an old illustration from

Seeman (1867) as C. verreauxii based on Fijian

material. As noted by Smith (1981), this is

referable to the Fijian endemic C. microtiglium.

Conservation status : Common. Present in at

least three conservation reserves in Queensland

(Forster et al. 1991) including Lamington and

Main Range National Parks. Recorded from nine

conservation reserves in New South Wales

(Floyd 1989).

Etymology: The specific epithet honours

J.P Verreaux (1807-1873), a French man resident

in Tasmania and one-time botanical collector.

31. Croton waterhouseae P.I.Forst., sp. nov.

affinis C. multicauli autem dentibus

marginis foliorumpluribus (32-40 vice lb-

28), pedicellis florium masculinorum

multo longioribus (10-12 mm vice 1.5-7

mm), staminibus in floribus masculinis

pluribus (32-38 vice 11-24) differt.

Typus: Queensland. Cook District:

Gabba Island, Torres Strait, 13 January

1998, B.M.Waterhouse BMW4775 &

J.Grimshaw (holo: BRI).

Shrub to 4 m high, monoecious, evergreen (?),

perennial. Indumentum silver to silver-

ferruginous. Branchlets ± rounded, with sparse

to dense sessile and shortly stalked stellate

trichomes, glabrescent. Stipules linear, 4-6 mm

long, c. 0.8 mm wide, with dense sessile and

shortly stalked stellate trichomes. Leaves

alternate, discolorous, petiolate; petioles 5-25

mm long, 1-1.2 mm wide, with dense sessile and

shortly stalked stellate trichomes; lamina elliptic

to obovate, 35-90 mm long, 20-50 mm wide,

Austrobaileya 6 (3): 349-436

palminerved with 2 nerves from base per side of

midrib and 4 lateral nerves per side of midrib,

and distinct tertiary reticulate veins; upper

surface dark green, venation weakly visible, with

sparse, sessile stellate trichomes mainly on

veins; lower surface grey-silver, lateral and

interlateral veins prominent, with dense sessile

and stalked stellate trichomes, velutinous;

margins weakly crenate, with 32-38 teeth to 2

mm long, foliar glands scattered and prominent;

tip acute; base cuneate to truncate; extrafloral

nectaries 2(3) at base of lamina, shortly stipitate

to 1 mm long, ellipsoid, 1.4-1.8 mm long, 1-1.2

mm wide, visible above and below.

Inflorescence up to 110 mm long, androgynous,

pedunculate up to 25 mm long, axis with sparse

to dense sessile and stalked stellate trichomes;

bracts linear-lanceolate, 0.8-1 mm long, 0.4-0.5

mm wide, with dense sessile stellate trichomes.

Male flowers 4-5 mm long, 6-7 mm diameter, in

sparse glomerules of 1 or 2 flowers in upper 4/5

of inflorescence; pedicels 10-12 mm long, c. 0.3

mm diameter, with dense sessile and stalked

stellate trichomes; sepals valvate, 5, lanceolate-

ovate, 2-2.3 mm long, c. 0.8 mm wide, with lanate

tip and with dense sessile to stalked stellate

trichomes; petals 5, obovate, 2-2.2 mm long and

c. 0.5 mm wide, lanate around entire edge;

stamens 32-38; filaments filiform, 2.5-3.5 mm

long and c. 0.2 mm wide, with dense, simple

trichomes at base; anthers oblong, 0.8-1 mm

long and c. 0.5 mm wide. Female flowers 2.8-3

mm long, c. 4 mm wide, held singly and spaced

up to 11 mm apart; pedicels 4.5-5 mm long, 0.8-

1 mm wide, with dense sessile stellate trichomes;

sepals valvate, 5, lanceolate-ovate, c. 2 mm long

and 1.5 mm wide, with dense sessile stellate

trichomes; petals absent; styles 3, linear-

flabellate, c. 2.5 mm long, bifid for c. 2 mm, with

scattered sessile stellate trichomes in lower

third; ovary 3-locular, c. 3 mm long and 2 mm

wide, with dense sessile stellate trichomes.

Fruits and seeds not seen. Fig. 31.

Distribution & habitat: Croton waterhouseae

is known only from the type locality at present.

Gabba Island is a continental granitic island

(Map 7). No information about the habitat was

available.

Phenology: The species probably flowers from

December through to February, with fruits

several months later. The only known specimen

is flowering and collected in January.

Forster, Croton in Australia

427

Fig. 31. Croton waterhouseae. A. habit of flowering branchlet. x 0.6. B. undersurface of leaf, x 1. C. base of leaf

lamina showing extrafloral nectaries, x 6. D. node showing stipule, x 6. E. inflorescence with all male flowers,

x 1.5. F. male flower, x 6. G. female flower, x 6. All from Waterhouse All 5 (BRI). Del. W. Smith.

428

Notes : Croton waterhouseae is closely allied

to the complex of species centred around

C. arnhemicus. It is perhaps most closely

related to Croton multicaulis, but differs from

that species in the greater number of leaf margin

teeth (32-40 per leaf, versus 16-28), the much

longer male flower pedicels (10-12 mm versus

1.5-7 mm ) and the greater number of stamens in

the male flowers (32-38 versus 11-24). Croton

arnhemicus is immediately distinguishable in

the much greater number of leaf margin teeth

(60-100 versus 32-40) and the generally

scabrous foliage (versus softly velutinous).

From both of these species it would also appear

that Croton waterhouseae may differ (more

specimens are required to be sure) in the

disposition of the veins that emanate from the

base of the lamina and that are immediately

adjacent to the midrib. In Croton waterhouseae

these veins steeply ascend beside the midrib at

an angle of no more than 30° terminating well

over half way along the leaf lamina. In both

Croton arnhemicus and C. multicaulis, as well

as the related C. aridus and C. minimus, these

same veins strongly diverge way from the midrib

at an angle of more than 45° and usually

terminate no more than half way along the leaf

lamina.

Conservation status : Unknown at this stage.

Gabba Island is uninhabitated.

Etymology : Named for Barbara Waterhouse,

NAQS botanist with the Australian Quarantine

Inspection Service, and collector of several

thousand specimens for the Queensland

Herbarium from northern Australia and adjacent

Malesia.

Excluded names and species

1. Croton argyratus Blume, Bijdr. 602 (1826).

Notes: This name was misapplied to the

Australian endemic Croton schultzii (see notes

there).

2. Croton opponens F.Muell. ex Benth., FI.

Austral. 6:125 (1873).

Notes: Base name for Bertya opponens (F. Muell.

ex Benth.) Guymer (Guymer 1985; Halford &

Henderson 2002).

Austrobaileya 6 (3): 349-436

3. Croton phebalioides var. hispida J.Simmonds,

Proc. Roy. Soc. Queensland 6: 68 (1889).

nom. nud. Type: not designated.

Notes: There is no diagnosis or type for this

name.

4. Croton prunifolius Airy Shaw, nom. illeg.

non Geiseler (1807), Kew Bull. 33: 56

(1978); Croton coccymelophyllus Radcl.-

Sm. & Govaerts, Kew Bull. 52:186 (1997).

Notes: The name Croton prunifolius was

misapplied by Airy Shaw (1981) to a sterile

collection of C. habrophyllus from Western

Australia. Because the name Croton prunifolius

Airy Shaw was illegimate, Radcliffe-Smith &

Govaerts (1997) consequently renamed this

species as Croton coccymelophyllus Radcl.-Sm.

& Govaerts. As a result this species was also

recorded with a ? from N. Western Australia in

Govaerts et al. (2000), a splendid example of

misinformation and error perpetuation. Croton

coccymelophyllus appears to be distributed in

parts of Malesia, such as the Lessa Sunda

Islands, Maluku and New Guinea.

5. Croton quadripartitus Labill. (as

‘quadripartitum’), Nov. Holl. PI. Sp. 2:73

(1806).

Notes: Base name for Adriana quadripartita

(Labill.) Muell.Arg. (Airy Shaw 1980:593).

6. Croton rosmarinifolius A.Cunn. (as

‘rosmarinifolium’) in Field, Geographical

Mem. New South Wales 355 (1825).

Notes: Base name for Ricinocarpos

rosmarinifolius (A.Cunn.) Benth.

7. Croton stigmatosus var. eurybioides Baill.,

Adansonia6: 301 (1866). nom. nud. Type:

not designated.

Notes: There is no diagnosis and no type for

this name.

8. Croton storckii (Muell.Arg,) A.C.Sm., Bull.

Bishop Mus. 141:83(1936).

Notes: This name was misapplied to Australian

material by Airy Shaw (1980b, 1981). The

populations concerned are referrable to Croton

mutabilis.

Forster, Croton in Australia

9. Croton urticoides A.Cunn. in Field,

Geographical Mem. New South Wales 355

(1825).

This is Adriana urticoides , see Appendix.

10. Croton viscosus Labill., Nov. Holl. PI. Sp. 2:

72(1806).

Notes: Base name for Beyeria viscosa (Labill.)

Miq.

Acknowledgements

The excellent artwork was executed by W. Smith

(BRI). Field work and the collection of

specimens over many years were facilitated with

the assistance of A.R. Bean, L.H. Bird, R. Booth,

P.D. Bostock, F. Carter, R. Fensham, A. Ford,

D.A. Halford, R.L. Jago, R. Jensen, G Kenning,

G Leiper, DJ. & I.M. Liddle, C. Lyons, P. Machin,

W.J.F. McDonald, D. Orford, J. Russell-Smith

& D. Lucas, G. & N. Sankowsky, PR. Sharpe,

G.Smyrell and M.C. Tucker. The Directors or

Curators of the cited herbaria allowed access to

specimens either on loan or in situ at the

institutions. L.W.Jessup (BRI) and PS.Short

(MEL) while Australian Botanical Liaison

Officers at Kew (U.K.) located and arranged for

photographs to be made of various specimens

at BM, K and P. The diagnoses were translated

into Latin by L.A.Craven (CANB) with the

exception of C. mamillatus, C. simulans and

C. waterhouseae that were prepared by

L.Pedley (BRI). Comments on the manuscript

were provided by H.-J. Esser and an anonymous

referee. The project was funded by the

Australian Biological Resource Study as part

of preferred objective research on the

Euphorbiaceae for the ‘Flora of Australia’ during

1992-1994. Additional funds for travel in the

‘Wet Tropics’ were provided by a grant for ‘Rare

or Endangered Euphorbiaceae of the Wet

Tropics’ from the Wet Tropics Management

Authority in 1993-1994.

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Appendix

Adriana urticoides (A.Cunn.) Guymer, comb. nov.

Croton urticoides A.Cunn., Field Geographical

Memoirs on NSW 355 (April 1825).

Type: Cox’s and Macquarie Rivers,

A. Cunningham, Oct-Dee 1822 (Chelsea

Physic Library, London, U.K. (n.v.), see

Mabberley (1978)).

Adriana tomentosa Gaudich., Ann. Sciences

Nat. 5:223 (8 July 1825). Type: ‘Habitatin

Nova-Hollandia (Orientali.) Baie de Chiens

Marins, Uranis, C. Gaudichaud’ (lecto:

P \fide Gross & Whalen 1996).

Two varieties were recognised by Gross and

Whalen (1996) based on their research of

morphological characters. As these varieties

are worthy of recognition, new combinations

are provided for them below.

Adriana urticoides var. urticoides

Adriana tomentosa Gaudich. var. tomentosa

See Gross and Whalen (1996) for a complete list

of synonyms.

Austrobaileya 6 (3): 349-436

Adriana urticoides (A.Cunn.) Guymer var.

hookeri (F.Muell.) Guymer, comb. nov.

Adriana tomentosa Gaudich. var. hookeri

(F.Muell.) C.L.Gross & M.A.Whalen,

Austral. System. Bot. 9:765 (1996);

Trachycarpon hookeri F.Muell., Trans.

Proc. Phil. Soc. Victoria 1:16 (1854). Type:

“On sand ridges along the Murray,

towards the junction of the Darling and

Murrumbidgee”, F. Mueller (lecto: E (n.v.),

fide Gross & Whalen 1996).

See Gross and Whalen (1996) for a complete list

of synonyms.

Mueller Argoviensis in A. de Candolle’s

Prodromus (1866) was the first to recognise that

Croton urticoides was a species of Adriana

and referred to it as Adriana acerifolia Cunn.

ex Hook. var. genuina nom. inval. in his list of

excluded names at the end of his treatment of

Croton (p. 699). However, he did not include

Croton urticoides as a synonym of any name

under Adriana in his treatment of that genus in

the same publication (p. 889). Index Kewensis

(1895) reported Croton urticoides as Adriana

glabrata and more recently the University of

Wisconsin on their Croton website has

incorrectly referred Croton urticoides to

Adriana tomentosa var. tomentosa.

The type of Croton urticoides was

collected by Allan Cunningham and is

presumably held in the ‘book’ herbarium of

Robert Heward’s in the Chelsea Physic Garden’s

library in London. Mabberley, in Taxon 27:

489-491 (1978), reported the discovery of

Heward’s ‘book’ herbarium entitled “Specimens

described in Field’s memoirs as a specimen of

the botany of the Blue Mountains” in this

library.

Forster, Croton in Australia

433

115 120 125 130 135 140 145 150 155

Map 2. Distribution of Croton in Australia. A C. aridus, ★ C. arnhemicus, • C. stigmatosus.

Map 3. Distribution of Croton in Australia. A C. schultzii, ■ C. brachypus, ★ C. multicaulis subsp. velutinus,

# C. acronychioides, ▼ C. choristadenius, & C. simulans, & C. stockeri.

434

Austrobaileya 6 (3): 349-436

Map 4. Distribution of Croton in Australia. ▲ C. bymesii, ■ C. dockrillii, ★ C. rarus, • C. minimus.

Map 5. Distribution of Croton in Australia. ▲ C. habrophyllus, ★ C. capitis-york, # C. densivestilus,

■ C. magneticus, ▼ C. mamillatus.

Forster, Croton in Australia

435

Map 6. Distribution of Croton in Australia. ★ C. insularis, • C. armstrongii.

115 120 125 130 135 140 145 150 155

Map 7. Distribution of Croton in Australia. • C. caudatus, ★ C. triacros, A C. verreauxii, ▼ C. waterhousecie.

436

Austrobaileya 6 (3): 349-436

Map 8. Distribution of Croton in Australia. ▲ C. multicaulis subsp. multicaulis, ★ C. phebalioides.

115 120 125 130 135 140 145 150 155

Map 9. Distribution of Croton in Australia. • C. tomentellus, ▲ C. mutabilis, ★ C. capitatus, ■ C. setigerus,

▼ C. glandulosus.

Phebalium distans P.I.Forst. (Rutaceae), a new and endangered species from

south-eastern Queensland, and reinstatement of P. longifolium S.T.Blake

Paul I. Forster

Summary

Forster, Paul I. Phebalium distans P.I.Forst. (Rutaceae), a new and endangered species from

south-eastern Queensland, and reinstatement of P. longifolium S.T.Blake. Austrobaileya 6(3):

437-444 (2003). Phebalium distans P.I.Forst. is named as a new species and distinguished from

P longifolium S.T.Blake (newly resurrected at specific rank) and P. squamulosum Vent. It is

known from ten extant populations in small remnants of semi-evergreen vine thickets in south¬

eastern Queensland and is considered as endangered due to the low number of populations with few

individuals. Both P. distans and P. longifolium are described and illustrated. A key to the species of

Phebalium in Queensland is provided.

Key words: Phebalium, Phebalium distans, Phebalium longifolium, Phebalium squamulosum,

Queensland - flora

Paul I. Forster, Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic

Gardens Mt Coot-tha, Mt Coot-tha Road, Toowong, Queensland 4066, Australia

Introduction

A revision of the species of Phebalium Vent,

occurring in Queensland was provided by

Wilson (1970) as part of his overall account of

the genus in Australia. More recently he

(Wilson 1998) revisited this genus and

transferred a number of species into the allied

genera Leionema (F.Muell.) Paul G. Wilson,

Nematolepis Turcz. and Rhadinothamnus Paul

G. Wilson. This new circumscription of

Phebalium meant that the genus consisted of

c. 25 species with 5 species recognised to occur

in Queensland (Forster 2002).

Wilson (1970) considered that some of the

“species” of Phebalium comprised complexes

of subspecies that were markedly disjunct, both

ecologically and geographically. This was

particularly the case with the taxa included under

P. squamulosum Vent, and he recognised some

ten subspecies for this species. Three of these

proposed subspecies, P. squamulosum subsp.

squamulosum, P. squamulosum subsp.

longifolium (S.T.Blake) Paul G. Wilson and

P. squamulosum subsp. gracile Paul G. Wilson

have been recognised for Queensland (Ross

1983,1994; Forster 1997,2002). It is generally

considered that subspecies should differ in only

Accepted for publication 3 February 2003

a few characters and that intermediate

populations should exist to demonstrate

continuity of character states (e.g. Stebbins

(1950) states “subspecies....connected....by a

series of intergrading forms” or Stace (1989)

states “a population of several biotypes

forming a more or less distinct regional facies

of a species.a geographical race, ecotype,

topodeme or genoecodeme”). This does not

seem to be the case for some of the taxa included

in P squamulosum, at least in Queensland,

where no obvious intermediates exist, there are

clear character state differences and the taxa

are markedly allopatric. Bruyns (2002) has

provided a succinct species concept in “species

are generally taken....as groups of populations

that differ in at least two persistent, “good”

characters” and subspecies are “geographically

complementary taxa that differ in only one

“reasonably reliable” character”. This has

always, and continues to be, the approach that

I have taken in the delimitation of taxa and these

species and subspecies concepts are applied

herein. This species concept equates broadly

with the ‘diagnostic species concept’ of Judd

et al. (2002).

In the current paper, a narrower

circumscription for Phebalium squamulosum is

advocated with P. squamulosum subsp.

438

longifolium reinstated to species rank as

P. longifolium S.T.Blake as listed in Forster

(2002). In addition, one new species (P. distans)

in this complex is described. These three species

differ in more than three character states one

from another. The morphological characters that

form the discontinuities in this group of taxa

are easily discernible and include habit (shrub

or tree), the form and composition of

indumentum (stellate trichomes, lepidote

scales), whether the branchlets are glandular-

tuberculate, the shape of the leaf lamina and

especially the length/width ratio, the form of

the leaf margin (flat or recurved), whether the

calyx is glandular-tubercular, corolla size, shape

and colour and seed size. All of the taxa thus

delimited are markedly allopatric, with the

northern P. longifolium having the greatest

disjunction from the others.

Materials & Methods

This paper is based on collections in the

Queensland Herbarium (BRI) augmented by

field observations and collections by the author

in Queensland and northern New South Wales.

Floral descriptions are based on material

preserved in spirit or by reconstitution in boiling

water. A full description of P. longifolium is

provided for comparative purposes.

Taxonomy

Phebalium distans P.I.Forst. sp. nov. a

P. squamuloso subsp. squamuloso habitu

arboris (non fruticis), lamina folii lineari

(anguste oblanceolata usque anguste

elliptica comparate), ratione longitudinis

/latitudinis 7.7-15.5 (non4.2-6.6), margine

recurvata (plana comparate), petalis

cremeis (non vivide flavis) differt.Typus:

Queensland. Burnett District: [5*] Mt

Walla, Walla Range, 5 km SW of Coalstoun

Lakes, 13 September 2002, PI. Forster

PIF28831 (holo: BRI [2 sheets + spirit];

iso: A, AD, DNA, HO, L, MEL, MO, NE,

NSW, NY, WELT, Z distribuendi)

Phebalium squamulosum subsp.

squamulosum auct. non Vent, pro parte

(Wilson 1970; Forster 1997,2002; Forster

etal. 1991).

Small tree (rarely a shrub) up to 8 m high, up to

15 cm dbh, bark rough-flaky, grey; blaze cream-

Austrobaileya 6 (3): 437-444

yellow with strong aromatic scent; wood yellow.

Indumentum (unless otherwise stated) on

foliage and reproductive parts of overlapping

lepidote trichomes that are silver to ferruginous-

silver giving the covered surface this colour.

Branchlets sparsely glandular-tuberculate, with

a dense covering of trichomes, glabrescent.

Leaves petiolate, strongly aromatic when

crushed; petioles 1.7-3 mm long, 0.5-0.8 mm

wide, with a sunken midrib and with dense

trichomes; lamina chartaceous, linear, 14-62 mm

long, 1.5-4.5 mm wide (length/width ratio 7.7-

15.5); margins entire or somewhat sinuate to

minutely crenate near apex, recurved; adaxial

surface with sunken midrib, glossy dark-green,

sparsely glandular, glabrous; abaxial surface

with strongly raised midrib, densely covered in

trichomes; tip apiculate to shortly acuminate;

base attenuate. Inflorescences pedunculate,

terminal umbels. Flowers 4-4.5 mm long, 3-4

mm wide; pedicels 4-5 mm long, c. 0.5 mm

diameter, with dense trichomes; mature bud

shape turbinate; calyx shortly subturbinate, 0.8-

1 mm long, 1.7-1.8 mm diameter, adaxially

glabrous, abaxially strongly glandular-

tuberculate and with sparse trichomes, lobes

broadly triangular, c. 0.3 mm long and 0.8 mm

wide, irregularly dentate; petals elliptic, 3-3.2

mm long, 1-1.8 mm wide, adaxially glabrous,

cream, abaxially with dense trichomes apart from

c. 0.2 mm around margin that is devoid of

trichomes; stamens 10, filaments 3.5-5 mm long,

c. 0.1 mm diameter, filiform, glabrous, anthers

oblong, 0.7-0.8 mm long, 0.4-0.5 mm wide; ovary

spherical, c. 1 mm high, with dense trichomes;

style 3-3.2 mm long, c. 0.3 mm diameter, with

scattered multifid stellate trichomes in lower half,

stigma capitate, papillate, c. 0.2 mm long and 0.3

mm wide. Cocci erect, 3.5—4 mm long, 2.5-3 mm

wide, glandular, truncate at suture. Seed

somewhat reniform, longitudinally compressed,

2.2-2.5 mmlong, 1.3-1.5 mm wide, longitudinally

corrugate, grey-black. Fig. 1.

Additional specimens examined : Queensland.

Burnett District: [1*] Spencers Road, 7 km E of

Wooroolin, Sep 2002, Forster PIF28870 & Smyrell

(BRI, MEL); [2*] Klass & Townes Road, 3.5 km SE of

Memerambi, Sep 2002, Forster PIF28873 & Smyrell

(BRI, MEL, NE, NSW); [3*] Kingaroy Heights Park

& Environmental Area, 3 km SE of Kingaroy, Dec

2002, Forster PIF29129 & Smyrell (BRI); [4*]

Couchmans road, 5 km N of Kingaroy, Dec 2002,

Forster PIF29137 & Smyrell (BRI); Kingaroy, Mar

1933, Lang [AQ152703] (BRI); [5*] 3 km SW of

Forster, Phebalium distans

439

Fig. 1. Phebalium distans. A. flowering stem, x 1. B. undersurface of leaf showing recurved margin, x 1.5. C.

flower, x 12. D. style, x 12. E. calyx showing markedly glandular-tuberculate surface and sparse covering of

lepidote trichomes. x 12. F. external view of petal, x 12. G. coccus, x 6. H. lateral view of seed, x 12. A from Leiper

(AQ678817) (BRI); B, F-H from Forster PIF24927 (BRI); C-E from Forster PIF28831 (BRI). Del. W. Smith.

440

Coalstoun Lakes, Walla Range, Aug 1990, Randall 613

(BRI); between Kingaroy & Memerambi, Mar 1986,

Schilling [AQ399826] (BRI); Wooroolin, s.dat. [?

Apr 1914], Simmonds [AQ152702] (BRI); [5*]

Coalstoun Lakes, Biggenden, Jun 1994, Thomas &

Sinclair [AQ636504] (BRI). Moreton District: [6*]

Scanlon Scrub, Mt Berryman area, Aug 1990, Bird &

Orford [AQ473152] (AD, BISH, BRI, CANB, DNA,

MEL, MO, NSW, PERTH); [6*] Mt Berryman area,

20 km S of Laidley, Dick Scanlon Scrub near Neumanns

Lookout, Nov 1991, Bird [AQ590868] (BRI); [7*]

Berlin Road, Mt Berryman area, 15 km SW of Laidley,

Mar 1992, Bird [AQ541907] (BRI; CANB, PERTH

n.v.); [8*] 1.5 km E of Mt Berryman, 10 km SSW of

Laidley, Aug 1985, Forster PIF2109 & Bird (BRI);

[7*] Welk Remnant, Mt Berryman, Sep 1999, Forster

PIF24927 & Booth (AD, BRI, MEL, QRS); ditto loc..

May 2002, Forster PIF28697 & Endress (A, BRI, L,

MEL, NE, NSW, Z); ditto loc., Sep 1999, Leiper

[AQ678817] (BRI, MEL, NSW).

* extant vouchered populations numbered 1-8, different

collectors have named these sites in various ways.

Notes: Phebalium distans was first collected

by J.H. Simmonds near Kingaroy, probably in

April 1914, as all other collections by him from

the Burnett were in this month. Until recently,

further collections of this species have been

spasmodically added to the herbarium record,

mainly from the Mt Berryman area in the Lockyer

Valley.

This species was included in the broad

concept of P. squamulosum advocated by

Wilson (1970). The collections by Lang and

Simmonds were examined by Paul Wilson in 1965

and he noted on the determinant slips (as

P. squamulosum var. squamulosum ) that they

were “approaching var. longifolium”.

Although individuals of P distans may

be initially shrublike, they will eventually form

small trees up to 8 m high with a stem up to 15

cm dbh. This tree habit is quite unique in the

genus Phebalium. The flowers of P. distans are

always cream (ageing cream-fawn). It is always

found in semi-evergreen vine thicket on red

volcanic soils or communities adjacent to this

vegetation type. It would appear to be allied to

both P. squamulosum and P longifolium but

cannot be considered as intermediate between

the two.

In comparison P. squamulosum subsp.

squamulosum is a small shrub (never a tree),

with narrow oblanceolate to narrow elliptic leaf

laminae (versus linear) with a length/width ratio

Austrobaileya 6 (3): 437-444

of 4.2-6.6 (versus 7.7-15.5) with a flat margin

(versus recurved) and the flowers are vivid

yellow. Some collections identified and

distributed as P. squamulosum subsp.

squamulosum from the Dorrigo area in northern

New South Wales (the oblanceolate-leaved

form illustrated in Weston & Porteners (1991))

have cream flowers and a mixture of lepidote

trichomes and stellate trichomes on the foliage,

whereas the forms at Mt Ballow (Border Ranges)

and Girraween N.P. in Queensland only have

lepidote trichomes. This variation in

P. squamulosum subsp. squamulosum requires

resolution, but is beyond the scope of the

current paper. Phebalium distans has lepidote

trichomes only on the foliage. In Queensland

Phebalium squamulosum subsp. squamulosum

has been found in heathland, shrubland or

woodland in rocky areas based on adamellite,

granite, rhyolite or trachyte substrates. Both

P. squamulosum and P. distans have a calyx that

is markedly glandular-tuberculate.

Phebalium longifolium is also a shrub,

with branchlets that are not glandular-

tuberculate. It has narrow-elliptic leaves with a

length/width ratio of 5-7.2, leaf laminae margins

with a flat edge, a calyx that is not markedly

glandular-tuberculate and seeds that are 1.8-2

mm long x 1-1.3 mm wide (versus 2.2-2.5 x 1.3-

1.5 mm). Both P. distans and P. longifolium

have cream flowers.

The only other taxon in this complex that

occurs in Queensland is Phebalium

squamulosum subsp. gracile. This taxon is a

small shrub (never a tree) with linear-oblong

leaf laminae (versus linear) that are markedly

shorter (5-25 mm long versus 14-62 mm) with a

length/width ratio of 3-7.7 (versus 7.7-15.5) and

yellow flowers with shorter petals (2-3 mm

versus 3-3.2 mm long). Whether or not

P. squamulosum subsp. gracile is worth

recognition at specific level requires further

study, particularly in relation to the other

proposed subspecies of P. squamulosum that

occur in New South Wales (Wilson 1970). As

an entity it is certainly quite distinct

from P. squamulosum subsp. squamulosum,

P. distans and P. longifolium.

Distribution and habitat : Phebalium distans

is currently known from ten populations, with

Forster, Phebalium distans

two of these unvouchered (viz. Forster et

al. 1991). Five of these are in close proximity to

one another at Mt Berryman. Four are near

Kingaroy, and the tenth most northerly one at

Mt Walla, is near Coalstoun Lakes. All except

the Mt Walla population, are on red volcanic

soils with semi-evergreen vine thicket. At Mt

Walla P. distans occurs in semi-evergreen vine

thicket, but the soil varies from red volcanic

where the population is at the base of the

mountain to rubble derived from rhyolitic

ignimbrite at the most elevated parts of the

population.

Conservation status: Phebalium distans is

considered to be endangered in habitat for

several reasons. Firstly less than 1000

individuals are known to exist with the extant

populations having the following estimated

number of plants (population number cited

above: number of plants estimated/area of

remnant in ha) (1: 7/<0.5; 2:3/<0.5:3:14/<0.5;

4:20/< 0.5; 5: <200/2; 6: < 200/30; 7: < 100/2; 8:

< 50/4). Several additional, but unvouchered

localities in the Mt Berryman area (Sites 174 &

178 in Forster et al 1991) have an unknown

number of plants, however the small sizes of

the remnants (5 & 3 ha respectively) means they

are unlikely to harbour a large number of

individuals.

Most of the suitable habitat for this

species in the Coalstoun Lakes, Kingaroy and

Lockyer valley areas was cleared for agriculture

in the 20 th century. Intensive survey of vine

thicket and vineforest remnants in these three

areas has been conducted since the mid 1980’s

(viz. Forster et al. 1991; Forster unpubl.,

W.J.F.McDonald unpubl.) and major

populations of P. distans cannot be considered

to have been overlooked throughout its known

range. Recent intensive survey (2002) of

roadside remnants in the Kingaroy Shire

revealed a further three populations

(populations 1,2,4). Population 1,2 and 4 are in

small roadside remnants and the other

populations are in more secure remnants (e.g.

Welk Remnant - population 7), freehold land

(populations 5, 6, 8) or on shire council land

with no active management strategies

(population 3). The large population 5 at Walla

Range (which co-occurs with the Endangered

Pomaderris clivicola E.M.Ross) has significant

441

intrusions through the stand of naturalised

pasture grasses such as Panicum maximum and

environmental weeds such as Lantana camara

that will act as a fire wick on the steep slope.

As with all Phebalium species, P. distans

has small seed that are shed locally from the

capsular fruit with little apparent long-range

dispersal ability. The disjunct occurrence of this

species in south-eastern Queensland suggests

P. distans was much more widespread in the

past, but is currently resticted due to the loss

of suitable habitat.

Under the IUCN (2001) risk categories,

P distans may be categorised as Endangered

under the criteria of B la, b (i, ii, iii, iv, v), 2b (i, ii,

iii, iv, v), C 2a (i).

Etymology: The specific epithet is derived from

the Latin distans (scattered) and refers to the

scattered extant populations of this species.

Phebalium longifolium S.T.Blake, Proc. Roy.

Soc. Queensland 70: 44 (1959). Type:

Queensland. North Kennedy District:

About W of Ingham, near Wallaman Falls,

14 August 1954, S.T. Blake 18809 (holo:

BRI [AQ318496]).

Shrub to 3 m high. Indumentum (unless

otherwise stated) on foliage and reproductive

parts of overlapping lepidote trichomes that are

golden-ferruginous to silver-ferruginous, giving

the covered surface this colour. Branchlets not

glandular-tuberculate, with a dense covering of

trichomes, glabrescent. Leaves petiolate,

aromatic when crushed; petioles 1.7-3 mm long,

0.7-0.8 mm wide, with a sunken midrib and with

dense trichomes; lamina chartaceous, narrow-

elliptic to narrow-oblanceolate, 15-80 mm long,

2.5-10 mm wide (length/width ratio 5-7.2);

margins entire or somewhat sinuate to minutely

dentate for entire length, thickened, flat; adaxial

surface with sunken midrib, glossy dark-green,

glabrous or sometimes with sparse trichomes

on margin, sparsely glandular-tuberculate;

abaxial surface with prominently raised midrib,

densely covered in trichomes; tip acute; base

attenuate. Inflorescences pedunculate, terminal

umbels. Flowers 4-5.5 mm long, 4-6 mm

diameter; pedicels 5-12 mm long, 0.5-0.8 mm

diameter, with dense trichomes; mature bud

shape turbinate; calyx shortly subturbinate, 0.7-

442

1.4 mm long, 2.2-3 mm diameter, adaxially

glabrous, abaxially with dense trichomes, not

glandular-tuberculate or only weakly so, lobes

broadly triangular, 0.3-0.5 mm long, 1-1.2 mm

wide, irregularly dentate; petals elliptic, 3-3.2

mm long, 1.5-1.8 mm wide, adaxially glabrous,

cream, abaxially with dense trichomes apart from

c. 0.2 mm margin that is devoid of trichomes;

stamens 10, filaments 3-5 mm long, c. 0.1 mm

diameter, filif orm, glabrous; anthers oblong, 0.7-

1 mm long, 0.4-0.5 mm wide; ovary spherical, c.

1 mm high, with dense trichomes; style 2.8-3.3

mm long, 0.1-0.2 mm diameter, with scattered

multifid stellate trichomes in lower half, stigma

capitate, papillate, c. 0.2 mm long and 0.3 mm

wide. Cocci erect, 3-3.5 mm long, 2.2-2.6 mm

wide, truncate at suture. Seeds somewhat

reniform, longitudinally compressed, 1.8-2.2 mm

long, 1-1.3 mm wide, longitudinally corrugate,

grey-black. Fig. 2.

Selected specimens examined: Queensland. Cook

District: Longlands Gap, S.F. 194, Jun 1995, Forster

PIF16776 (BRI, MEL, QRS); ditto loc., Sep 2001,

Forster PIF27518 et al. (A, BRI, K, L, MET,); S.F. 194

Mt Baldy, Oct 1999, Forster PIF25088 & Booth (AD,

BRI, MEL, QRS); Herberton Range S.F., Apr 1998,

Jago 4720 (BRI). North Kennedy District: Bluewater

S.F., 55 km NW of Townsville, Nov 1991, Bean 3790

Austrobaileya 6 (3): 437-444

(BRI; BISH, CANB n.v.): Birthday Falls, Paluma Range,

Sep 1966, Birch [AQ152608] (BRI); W side of Paluma

- Hidden Valley road, c. 10 km W of Paluma, Jul 1992,

Jobson 1730 et al. (BRI; MEL, NSW n.v.); S.F, 344, c.

27 km W of Kennedy township, May 1976, Thorsborne

218 (BRI).

Distribution and habitat : Phebalium

longifolium is endemic to the “Wet Tropics” of

north-eastern Queensland in several disjunct

populations from the Herberton Range in the

north to Paluma in the south. It inhabits the

edges of wet rainforest in the ecotone to

adjacent eucalypt open forest, usually on basalt

or metamorphic substrates.

Notes: This species is easily distinguished from

P. squamulosum by the long narrow-elliptic leaf

laminae with an attenuate base (versus cuneate),

the cream flowers (versus yellow) with a non-

glandular-tuberculate (or only weakly so) calyx

and with narrower petals (1.5-1.8 mm wide

versus 1.8-2 mm wide).

Conservation status: Phebalium longifolium

is well represented in State Forests (S.F. 194,

268, 344) and National Parks (Mt Spec)

throughout its range and is not considered

threatened.

Key to the species of Phebalium found in Queensland. N.B. P. distans is usually a tree,

but is also keyed out as a shrub for encounters with small individuals of this species.

1. Flowers large with petals 4.5-9 mm long.2

Flowers small with petals 2^1 mm long.4

2. Calyx with short lobes < 1 mm long; corolla yellow.P. whitei

Calyx with well developed lobes 1-5 mm long; corolla white or pink.3

3. Calyx 5-6 mm diameter, cupular, 6-8 lobed; petals pink. P.nottii

Calyx 2-3 mm diameter, obturbinate, 5-lobed; petals white or pink. P. woombye

4. Tree.P. distans

Shrub.5

5. Leaf lamina linear to oblong-cuneate.6

Leaf lamina narrow-elliptic to narrow-oblanceolate.8

6. Leaf margin undulate, markedly glandular. P. glandulosum

Leaf margin entire or somewhat sinuate to minutely crenate near the apex,.7

Forster, Phebalium distans

443

Fig. 2. Phebalium longifolium. A. flowering stem, x 1. B . undersurface of leaf showing flat margin, x 1.5. C.

flower, x 6. D. style, x 12. E. calyx showing poor development of glandular-tuberculate surface and dense coverage

of lepidote trichomes. x 12. F. external view of petal, x 12. G. coccus, x 6. H. seed, x 12. A, B, G, H from Forster

PIF25088 (BRI); C-F from Jago 4720 (BRI). Del W. Smith.

444

Austrobaileya 6 (3): 437-444

7. Leaf lamina short, 5-25 mm long, with a length/width ratio of 3-7.7;

corolla yellow, petals 2-3 mm long. P. squamulosum subsp. gracile

Leaf lamina long, 14-62 mm long, with a length/width ratio 7.7-15.5; corolla

cream, petals 3-3.2 mm long.P. distans

8. Leaf lamina with a cuneate base; corolla yellow, petals 1.8-2 mm wide

. P. squamulosum subsp. squamulosum

Leaf lamina with an attenuate base; corolla cream, petals 1.5-1.8 mm wide

. P. longifolium

Acknowledgements

I wish to thank W.M. Powell for access to

Mt Walla; W.Smith (BRI) for the illustrations;

L.Pedley for the Latin diagnosis, and L.H.Bird,

R.Booth (BRI), F. Carter (QPWS), R.Jensen,

G.Leiper, P.Machin and G. Smyrell for assistance

with fieldwork where Phebalium material was

collected.

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A synopsis of Racosperma C. Mart. (Leguminosae: Mimosoideae)

Les Pedley

Summary

Pedley, L. (2003). A synopsis of Racosperma C.Mart. (Leguminosae: Mimosoideae). Austrobaileya

6(3): 445-496. The history of the genus Racosperma, which is based on Acacia (sect.) Phyllodineae

DC., is briefly outlined. Names of 976 species of Racosperma (with 130 names of infraspecific

taxa, autonyms excluded) are listed. Of these, only ten species (one represented by one of two

subspecies) do not occur in Australia. Transfers of 834 taxa (711 to species, 58 to subspecies and

65 to variety) are made to Racosperma the vast majority from Acacia, 15 from Mimosa and one

from Adiantum. Taxonomy closely follows the treatment of Acacia in the Flora of Australia,

vols 11A & 11B (Orchard & Wilson 2001) and reference is made to the treatment of each species

in the Flora. Acacia cyclops, A. stowardii and A. vincentii are treated as synonyms of Racosperma

eglandulosum, R. sibiricum, and R. deltoideum respectively. Acacia juncifolia subsp. serpentinicola

is raised to specific rank, Racosperma serpentinicola ; as is Acacia wickhamii subsp. parviphyllodinea

to Racosperma calligerum (nom. nov.), with A. wickhamii subsp. viscidula as a synonym of it.

Acacia clivicola, usually considered conspecific with A. stowardii, is reinstated as R. clivicola',

A. cupularis is referred to R. ligulatum var. minus and A. curvinervia to R. juliferum subsp.

curvinervium. R. sophorae is considered to be a species distinct from R. longifolium. A. desmondii

and A. racospermoides are the same as R. nelsonii and R. paniculatum respectively.

Key words: Racosperma, Acacia, Australia, nomenclature, new combinations.

Les Pedley, cl- Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic

Gardens Mt Coot-tha, Mt Coot-tha Road, Toowong, Queensland 4066, Australia

Introduction

The name Racosperma was first used by C.F.P.

von Martius in a catalogue of plants growing in

the Royal Botanical Garden, Munich 1 (Martius

1829). He listed species of Acacia in two groups,

namely ‘ Acacia genuinae / (Omnes ip 2 ,

pleraeque C.)’ and ‘Acaciae phyllodineae De

C. (Genus dist.: Racosperma Mart.) /(Pleraeq.

N. Holl., omn. ip TJ. Species in each group

were arranged alphabetically. Since he included

Racosperma in Acacia, it must be taken that

Martius did not accept it as a distinct genus at

that time, and the 21 specific epithets listed under

it cannot be considered as having been

transferred from Acacia to Racosperma.

Martius (1835) again used the name, this time

quite definitely as a generic name. In an

alphabetical list it was placed between Pyrus

and Ranunculus. He listed three species, the

names of two of which are now considered

synonyms of Racosperma paradoxum. The

other, Racosperma sophorae, is the only

currently accepted name that can be considered

as being validly published in Racosperma by

Martius. 3

In reviewing the classification of Acacia

sens. lat. (Pedley 1986a) I reinstated the generic

name Racosperma. I used the name in a number

of subsequent papers (Pedley 1986b, 1987a,

1987b, 1987c, 1987d, 1988), but refrained from

transferring the names of all Australian species

to Racosperma. Over a considerable period I

had more or less confined my studies to the

Queensland flora and considered it fitting that

the necessary new combinations be made by

workers whose areas of expertise covered other

1 Not Monaco as I stated previously (Pedley 1986a).

2 An approximation to the Linnean symbol meaning woody, see Stearn (1992: 351).

3 Previously I also carelessly, wrongly attributed the following names to Martius: Racosperma falcatum,

R. hispidulum, R. melanoxylon, R. strictum (Pedley 1987c), R. longifolium, R. paradoxum, R. verticillatum

(Pedley 1987d). In all cases I cited Martius (1835), which is incorrect. They were listed by Martius (1829),

but not in the later publication.

Accepted for publication 14 March 2003

446

parts of Australia. Those expected

combinations were not forthcoming. In fact,

opposition to the adoption of the name

Racosperma was widespread and strong. An

extreme example of the views of the interested

lay community was given by Anon. (1993) who

proclaimed inter alia that “the name

Racosperma is an abomination”. A more

measured, less emotional, but nevertheless

hostile, response was that of Kanis (1986) who

wrote: “Recently it was proposed by Pedley

(1986) to promote Vassal’s subgenera to genera

in their own right. Among other things this will

mean the transfer of more than 90% of the

Australian species to Racosperma Mart., a bold

step indeed. However some insiders fear that

this might not be the last word at generic level”.

In the intervening years I have seen nothing of

substance to support the fears mentioned. The

validity of the name was also questioned, mainly

covertly, but Chappill & Maslin (1995) spelled

out the problem as they saw it: “The basic point

of contention is whether Martius’ (1835)

publication can be indirectly connected to his

earlier (Martius 1829) listing of the name where

he indicated that it should be based on Acacia

(section) Phyllodineae DC. (fide Maslin 1988).

If Racosperma is not validly published then the

name must date from Pedley (1986) and will

therefore need to be conserved against earlier

names”. They went on to discuss the

consequences and possible pitfalls of such

conservation.

In view of the lack of endorsement of

Racosperma and for convenience in managing

BRI collections which include many taxa that

do not occur in Queensland, I transferred names

of species which I had described originally as

Racosperma to Acacia (Pedley 1990) and

subsequently referred species described by me

as new to the latter genus. In a short

introduction to the 1990 paper, I explained why

those actions should not be construed as

implying that I did not accept Racosperma as a

distinct genus. Neither do I doubt the validity

of the name Racosperma. Martius’s 1835 use

of it was an indirect reference to his 1829 use,

and meets the requirements of Art. 32.4 of the

International Code of Botanical Nomenclature

(St Louis Code) (Greuter et al. 2000), and is

covered by Ex. 4 appended to the article. Instead

of Racosperma I could have adopted one of

several names proposed by Rafinesque (1838).

Austrobaileya 6 (3): 445-496 (2003)

No one seems to have any difficulty in tracing

Racosperma from Martius’s publication of the

name in 1835 back to his 1829 publication, and

its conservation is not necessary.

The situation has now changed. More

recent studies of Chappill & Maslin (1995),

Grimes (1999), Robinson & Harrison (2000), and

particularly the molecular work of Miller &

Bayer (2000, 2001) provide strong grounds for

the recognition of the ‘Australian acacias’ (that

is, Acacia subg. Phyllodineae (DC.) Seringe;

Acacia subg. Heterophyllum Vassal) as a genus

distinct from Acacia. The recognition of

Racospermyces, a rust fungus related to

Uromyces, restricted to Australian and extra-

Australian species Walker (2001), also adds

weight to the mycological evidence previously

presented (Pedley 1986a) for the distinctiveness

of Racosperma. Though I have a first-hand

knowledge of possibly only one-third of the

Australian acacias, in the words of that well

known dissident, Croizat (1958: 202), “the time

is right here to call a spade a spade and to stop

floating in molasses”. Now is an appropriate

time to complete the transfer of all taxa to

Racosperma. Since the genus is virtually

restricted to Australia publication of Flora of

Australia vol. 11A & 11B (Orchard & Wilson

2001) has made this possible. It has brought

together descriptions of all species accepted

by the contributors to the Flora up to about

the end of 2000. Importantly, in the compilation

of a check-list such as this, the Flora provides

basionyms and references to their protologues.

It is now feasible to compile a list of up-to-date

species of Racosperma without the need to

check a mass of bibliographic references. I have,

however, consulted protologues where

possible.

On the whole I have followed the

taxonomy presented in the Flora, but have

diverged in a few cases where my views differ

from those presented by the authors.

Taxonomists working on acacias from Australia

have used both ‘variety’ and, since the mid-

1960s, ‘subspecies’ as infraspecific categories;

some one, some the other, and some (myself

included) both. It is sometimes difficult to

distinguish their concepts of one from the other;

as Benson (1962) stated, “interpretations of the

two ranks vary; what a particular botanist

interprets as one is exactly what another person

447

Pedley, A synopsis of Racosperma

accepts as the other.” By and large, I have

retained the categories accepted by the Flora

authors, but, in some species that I know well, I

have reduced subspecies to variety.

Notes on text

The name accepted in Racosperma is followed

on the next line, if it is a comb, nov., by the

basionym. Infraspecific taxa are listed under

the relevant species. Autonyms are not listed,

except in the case of Racosperma spirorbis

subsp. spirorbis, one of the taxa which does

not occur in Australia.

An asterisk (*) indicates that the taxon

does not occur in Australia. Literature

references in addition to the reference to the

basionym are given for extra-Australian taxa.

The number in italics following a section

mark (§) refers to the species number in the Flora

of Australia. An obelus (f) indicates that the

taxon is not included in the Flora, usually

because it has been described since its

publication. If a species is merely mentioned in

the text of the Flora, without reference to its

description, usually because of its later

publication or because its name was treated as

a synonym of a taxon that is included in the

Flora, then it is referred (by the use of sub) to

that taxon.

Brief notes have been added in the places

where the taxonomy of the Flora has not been

followed. Some changes will be discussed in

subsequent papers.

Racosperma abbatianum (Pedley) Pedley, comb,

nov. f

Acacia abbatiana Pedley, Austrobaileya 5:

313(1999)

Racosperma abbreviatum (Maslin) Pedley,

comb. nov. §747

Acacia abbreviata Maslin, J. Adelaide Bot.

Gard. 2:301 (1980).

Racosperma abruptum (Maiden & Blakely)

Pedley, comb. nov. §476

Acacia abrupta Maiden & Blakely, J. Roy.

Soc. W. Australia 13:6 (1927).

Racosperma acanthaster (Maslin) Pedley,

comb. nov. §314

Acacia acanthaster Maslin, Nuytsia 12: 312

(1999).

Racosperma acanthocladum (F.Muell.) Pedley,

comb.nov. §311

Acacia acanthoclada F.Muell., Fragm. 3:127

(1863).

R. acanthocladum subsp. glaucescens (Maslin)

Pedley, comb. nov. §3 lib

Acacia acanthoclada subsp. glaucescens

Maslin, Nuytsia 12: 314 (1999).

Racosperma acelleratum (Maiden & Blakely)

Pedley, comb. nov. §551

Acacia acellerata Maiden & Blakely, J. Roy.

Soc. W. Australia 13:2 (1928).

Racosperma acinaceum (Lindl.) Pedley, comb,

nov. §446

Acacia acinacea Lindl., in T.L.Mitchell, Three

Exped. Australia 2:265 (1838).

Racosperma aciphyllum (Benth.) Pedley, comb,

nov. §854

Acacia aciphylla Benth., Linnaea 26: 627

(1855).

Racosperma acomum (Maslin) Pedley, comb,

nov. §436

Acacia acoma Maslin, Nuytsia 12:315 (1999).

Racosperma acradenium (F.Muell.) Pedley,

Austrobaileya 2: 344 (1987). §688

Racosperma acrionastes (Pedley) Pedley, comb,

nov. §121

Acacia acrionastes Pedley, Austrobaileya 3:

297(1990).

Racosperma acuarium (W.V.Fitzg.) Pedley,

comb. nov. §299

Acacia acuaria W.V.Fitzg., J. W. Australian

Nat. Hist. Soc. 7 (1904).

Racosperma aculeatissimum (J.F.McBr.)

Pedley, comb. nov. §289

Acacia aculeatissima J.F.McBr., Contrib. Gray

Herb. 59:6(1919).

Racosperma aculeiforme (Maslin) Pedley,

comb. nov. §397

Acacia aculeiformis Maslin, Nuytsia 12: 317

(1999).

Racosperma acuminatum (Benth.) Pedley,

comb.nov. §866

Acacia acuminata Benth., London J. Bot. 1: 373

(1842).

See also Racosperma burkittii.

448

Racosperma acutatum (W.V.Fitzg.) Pedley,

comb.nov. §337

Acacia acutata W.V.Fitzg., J. W. Australian

Nat. Hist. Soc. 6(1904).

Racosperma adenogonium Pedley,

Austrobaileya2:316 (1987). §618

Racosperma adinophyllum (Maslin) Pedley,

comb.nov. §405

Acacia adinophylla Maslin, Nuytsia 12: 318

(1999).

Racosperma adnatum (F.Muell.) Pedley, comb,

nov. §541

Acacia adnata F.Muell., Chem. & Druggist

Australas. Suppl. 5 (51): 26 (1882).

Racosperma adoxum (Pedley) Pedley, comb. nov.

§913

Acacia adoxa Pedley, Contrib. Queensland

Herb. 11:6(1972).

R. adoxum var. subglabrum (Pedley) Pedley,

comb.nov. §913b

Acacia adoxa var. sub glabra Pedley, Contrib.

Queensland Herb. 11:7 (1972).

Racosperma adsurgens (Maiden & Blakely)

Pedley, Austrobaileya 2: 344 (1987).§S79

Racosperma aduncum (A.Cunn. ex G.Don)

Pedley, Austrobaileya 2:344 (1987).§720

Racosperma aemulum (Maslin) Pedley, comb,

nov. §367

Acacia aemula Maslin, Nuytsia 10:169 (1995).

R. aemulum subsp. muricatum (Maslin) Pedley

comb.nov. §367b

Acacia aemulum subsp. muricata Maslin,

Nuytsia 10:171 (1995).

Racosperma aestivale (E.Pritzel) Pedley, comb,

nov. §104

Acacia aestivalis E.Pritzel, Bot. Jahr. Syst. 35:

300(1904).

Racosperma alatum (R.Br.) Pedley, comb. nov.

§252

Acacia alata R. Br. in W.T. Aiton: Hortus Kew.

ed. 2.5:464(1813).

Austrobaileya 6 (3): 445-496 (2003)

R. alatum var. biglandulosum (Benth.) Pedley,

comb.nov. §252b

Acacia alata var. biglandulosa Benth., FI.

Austral. 2:321 (1864).

R. alatum var. platypterum (Lindl.) Pedley, comb,

etstat. nov. 252c

Acaciaplatyptera Lindl., Bot. Reg. 27: misc.

3(1841).

R. alatum var. tetranthum (Maslin) Pedley, comb,

nov. §252d

Acacia alata var. tetrantha Maslin, Nuytsia

10:157(1995).

Racosperma alcockii (Maslin & Whibley)

Pedley, comb. nov. §85

Acacia alcockii Maslin & Whibley, Nuytsia

6:19(1987).

Racosperma alexandri (Maslin) Pedley, comb,

nov. §197

Acacia alexandri Maslin, Nuytsia 8: 288

(1992).

Racosperma allenianum (Maiden) Pedley,

Austrobaileya 2:344 (1987). §188

Racosperma alpinum (F.Muell.) Pedley, comb,

nov. §896

Acacia alpina F.Muell., Fragm. 3:129 (1863).

Racosperma amandae (GJ.Leach) Pedley, comb,

nov. §641

Acacia amandae GJ.Leach (as ‘amanda’), FI.

Australia 1 IB: 488 (2001).

The specific epithet amanda was published

deliberately as a noun in apposition with the

generic name. Article 23 of the International

Code of Botanical Nomenclature permits such

usage, and further states that the epithet may

be taken from any source whatever. On the

other hand, Recommendation 23A states that

the names of persons used in specific epithets

should take the form of nouns in the genitive

or of adjectives. Since, in practice,

Recommendations often have the force of

Articles and since ‘scientific names of

taxonomic groups are treated as Latin regardless

of their derivation’ (Principle IV), I consider the

449

Pedley, A synopsis of Racosperma

lack of a genitive inflection to be an error to be

corrected. See also Racosperma dorotheae and

R. veronicae.

A complication in the present case is that

amanda, as well as being a common enough

English-language given name, is also the

gerundive of the Latin amo, and could be, and

has been when occurring in other genera,

declined like any other adjectival epithet.

Racosperma amblygonum (A.Cunn. ex Benth.)

Pedley, Austrobaileya 2: 344 (1987). §302

Racosperma amblyophyllum (F.Muell.) Pedley,

comb.nov. §99

Acacia amblyophylla F.Muell., South. Sci.

Rec. 2 (7): 149(1882).

Racosperma amentiferum (F.Muell.) Pedley,

comb.nov. §741

Acacia amentifera F.Muell., J. Proc. Linn Soc.,

Bot.3:141(1859).

Racosperma ammobium (Maconochie) Pedley,

Bot. J. Linn. Soc. 92:247 (1986). §801

Racosperma ammophilum (Pedley) Pedley,

Austrobaileya 2: 344 (1987). §607

Racosperma amoenum (H.L.Wendl.) Pedley,

comb.nov. §72

Acacia amoena H.L.Wendl., Comm. Acac.

Aphyll. 41.4 (1820).

Racosperma ampliatum (R.S.Cowan & Maslin)

Pedley, comb. nov. §848

Acacia ampliata R.S.Cowan & Maslin,

Nuytsia 10:16 (1995).

Racosperma ampliceps (Maslin) Pedley, comb,

nov. §219

Acacia ampliceps Maslin, Nuytsia 1: 315

(1974).

Racosperma amputatum (Maslin) Pedley, comb,

nov. §940

Acacia amputata Maslin, Nuytsia 12: 493

(1999).

Racosperma amycticum (R.S.Cowan & Maslin)

Pedley, comb. nov. §569

Acacia amyctica R.S.Cowan & Maslin,

Nuytsia 10:222(1995).

Racosperma anarthon (Maslin) Pedley, comb,

nov. §949

Acacia anarthos Maslin, Nuytsia 2: 354

(1979).

Racosperma anasillum (A. S. George) Pedley,

comb. nov. §917

Acacia anasilla A.S.George, J. Roy. Soc. W.

Australia 82:67 (1999).

Racosperma anastemum (Maslin) Pedley, comb,

nov. § 846

Acacia anastema Maslin, Nuytsia 4: 383.

(1983).

Racosperma anaticeps (Tindale) Pedley, comb,

nov. §642

Acacia anaticeps Tindale, Contrib. New

South Wales Natl Herb. 4:269 (1972).

Racosperma anceps (DC.) Pedley, comb. nov.

Acacia anceps DC., Prodr. 2:451 (1825).

Racosperma ancistrocarpum (Maiden &

Blakely) Pedley, Austrobaileya 2: 344

(1987). §758

Racosperma ancistrophyllum (C.R.P. Andrews)

Pedley, comb. nov. §568

Acacia ancistrophylla C.R.P.Andrews, J. W.

Australia Nat. Hist. Soc. 40 (1904).

R. ancistrophyllum var. lissophyllum (J.M.

Black) Pedley, comb. nov. §568b

Acacia sclerophylla var. lissophylla

J.M.Black, Trans. & Proc. Roy. Soc. S.

Australia 47:369 (1923).

R. ancistrophyllum var. perarcuatum

(R.S.Cowan & Maslin) Pedley, comb. nov.

§568c

Acacia ancistrophylla var. perarcuata

R.S.Cowan & Maslin, Nuytsia 10: 227

(1995).

450

Racosperma andrewsii (W.V. Fitzg.) Pedley,

comb.nov. §353

Acacia andrewsii W.V.Fitzg., J. W. Australia

Nat. Hist. Soc. 6(1904).

Racosperma aneurum (F.Muell. ex Benth.)

Pedley, Austrobaileya2: 344 ( \9%1).§839

R. aneurum var. argenteum (Pedley) Pedley,

comb.nov. §839j

Acacia aneura var. argentea Pedley, FI.

Australia 1 IB: 490 (2001).

R. aneurum var. coniferum (Randell) Pedley,

comb.nov. §839a

Acacia aneura var. conifera Randell, J.

Adelaide Bot. Gard. 14:122(1992).

R. aneurum var. fuligineum (Pedley) Pedley,

comb.nov. §839i

Acacia aneura var. fuliginea Pedley, FI.

Australia 1 IB: 489 (2001).

R. aneurum var. intermedium (Pedley) Pedley,

comb.nov. §839g

Acacia aneura var. intermedia Pedley, FI.

Australia 1 IB: 489 (2001).

R. aneurum var. macrocarpum (Randell) Pedley,

comb.nov. §839b

Acacia aneura var. macrocarpa Randell, J.

Adelaide Bot. Gard. 14:121 (1992).

R. aneurum var.microcarpum (Pedley) Pedley,

comb.nov. §839c

Acacia aneura var. microcarpa Pedley, FI.

Australia 1 IB: 489 (2001).

R. aneurum var. majus (Pedley) Pedley, comb,

nov. §839f

Acacia aneura var. major Pedley, FI. Australia

1 IB: 489 (2001).

R. aneurum var. pilbaranum (Pedley) Pedley,

comb.nov. §839e

Acacia aneura var. pilbarana Pedley, FI.

Australia 1 IB: 489 (2001).

R. aneurum var. tenue (Pedley) Pedley, comb,

nov. §839d

Acacia aneura var. tenuis Pedley, FI. Australia

1 IB: 489 (2001).

Austrobaileya 6 (3): 445-496 (2003)

Racosperma anfractuosum (Maslin) Pedley,

comb.nov. §887

Acacia anfractuosa Maslin, Nuytsia 2: 96

(1976).

Racosperma angustum (Maiden & Blakely)

Pedley, Austrobaileya 2: 344 (1987). §129

Racosperma anomalum (C.A.Gardner ex Court)

Pedley, comb. nov. §249

Acacia anomala C.A.Gardner ex Court,

Nuytsia 2:168(1978).

Racosperma anthochaerum (Maslin) Pedley,

comb.nov. §212

Acacia anthochaera Maslin, Nuytsia 10: 183

(1995).

Racosperma aphanocladum (Maslin) Pedley,

comb.nov. §198

Acacia aphanoclada Maslin, Nuytsia 8: 290

(1992).

Racosperma aphyllum (Maslin) Pedley, comb,

nov. §250

Acacia aphylla Maslin, Nuytsia 1: 320 (1974).

Racosperma applanatum (Maslin) Pedley, comb,

nov. §254

Acacia applanata Maslin, Nuytsia 10: 158

(1995).

Racosperma apreptum (Pedley) Pedley,

Austrobaileya 2:344 (1987). §815

Racosperma apricum (Maslin & A. R.Chapm.)

Pedley, comb. nov. §874

Acacia aprica Maslin & A.R.Chapm., Nuytsia

12:471(1999).

Racosperma arafuricum (Tindale & Kodela)

Pedley, comb. nov. §612

Acacia arafurica Tindale & Kodela, Telopea

5:53(1992).

Racosperma araneosum (Whibley) Pedley,

comb.nov. §89

Acacia araneosa Whibley, Contrib. Herb.

Austral. 14:1 (1976).

Racosperma arbianum (Pedley) Pedley, comb,

nov. §163

Acacia arbiana Pedley, Austrobaileya 5: 307

(1999).

451

Pedley, A synopsis of Racosperma

Racosperma arcuatile (R.S.Cowan & Maslin)

Pedley, comb. nov. §873

Acacia arcuatilis R.S.Cowan & Maslin,

Nuytsia 12:472 (1999).

Racosperma areolatum (M.W.McDonald)

Pedley, comb. nov. f

Acacia areolata M.W.McDonald, Austral.

SystBot. 16:142, t.2 (2003).

Racosperma argutifolium (Maslin) Pedley,

comb. nov. §361

Acacia argutifolia Maslin, Nuytsia 2:98

(1976).

Racosperma argyraeum (Tindale) Pedley,

Austrobaileya 2: 344 (1987). §763

Racosperma argyrodendron (Domin) Pedley,

Austrobaileya 2: 345 (1987). §611

Racosperma argyrophyllum (Hook.) Pedley,

comb. nov. §176

Acacia argyrophylla Hook., Bot. Mag. 74: t.

4384(1848).

Racosperma argyrotrichum (Pedley) Pedley,

comb. nov. §585

Acacia argyrotricha Pedley, Austrobaileya

5:310(1999).

Racosperma aridum (Benth.) Pedley, comb. nov.

§733

Acacia arida Benth., London J. Bot. 1: 370

(1842).

Racosperma aristulatum (Maslin) Pedley,

comb. nov. §318

Acacia aristulata Maslin, Nuytsia 12: 320

(1999).

Racosperma armillatum Pedley, Austrobaileya

2:325(1987). §624

Racosperma armitii (F.Muell. ex Maiden)

Pedley, Austrobaileya2: 345 (1987). §700

Racosperma arrectum (Maslin) Pedley, comb,

nov. §750

Acacia arrecta Maslin, Nuytsia 4: 73 (1982).

Racosperma ascendens (Maslin) Pedley, comb,

nov. §477

Acacia ascendens Maslin, Nuytsia 7: 223

(1990).

Racosperma asepalum (Maslin) Pedley, comb,

nov. §380

Acacia asepala Maslin, Nuytsia 12: 321

(1999).

Racosperma ashbyae (Maslin) Pedley, comb,

nov. §40

Acacia ashbyae Maslin, Nuytsia 1:321 (1974).

Racosperma asparagoides (A.Cunn.) Pedley,

comb. nov. §293

Acacia asparagoides A.Cunn. in B.Field,

Geogr. Mem. New South Wales. 343 (1825).

Racosperma asperulaceum (F.Muell.) Pedley,

Austrobaileya 2: 345 (1987). §925

Racosperma asperum (Lindl.) Pedley, comb. nov.

§451

Acacia aspera Lindl., in T.L.Mitchell, Three

Exped. Australia 2:138(1838).

Racosperma assimile (S.Moore) Pedley, comb,

nov. §534

Acacia assimilis S. Moore, J. Linn. Soc., Bot.

45:172(1920).

R. assimile subsp. atroviride (R.S.Cowan &

Maslin) Pedley, comb. nov. §534b

Acacia assimilis subsp. atroviridis

R.S.Cowan & Maslin, Nuytsia 10: 240

(1995).

Racosperma ataxiphyllum (Benth.) Pedley,

comb. nov. §356

Acacia ataxiphylla Benth., Linnaea 26: 605

(1855).

R. ataxiphyllum subsp. magnum (Maslin)

Pedley, comb. nov. §356b

Acacia ataxiphylla subsp. magna Maslin,

Nuytsia 12: 324(1999).

Racosperma atkinsianum (Maslin) Pedley,

comb. nov. §814

Acacia atkinsiana Maslin, Nuytsia 4: 75

(1982).

452

Racosperma atopum (Pedley) Pedley, comb. nov.

§845

Acacia atopa Pedley, FI. Australia 11B: 490

( 2001 ).

Racosperma atrox (Kodela) Pedley, comb. nov.

Acacia atrox Kodela, Telopea 9:315 (2001). f

Racosperma attenuatum (Maiden & Blakely)

Pedley, Austrobaileya 2: 345 (1987). § 70

Racosperma aulacocarpum (A.Cunn. ex Benth.)

Pedley, Austrobaileya 2: 345 ( \9%1).§670

R. aulacocarpum var. fruticosum (C.White)

Pedley, Austrobailey 2:345 (1987) syn.nov.

Racosperma aulacophyllum (R.S.Cowan &

Maslin) Pedley, comb. nov. §533

Acacia aulacophylla R.S.Cowan & Maslin,

Nuytsia 10:241(1995).

Racosperma auratiflorum (R.S.Cowan &

Maslin) Pedley, comb. nov. §502

Acacia auratiflora R.S.Cowan & Maslin,

Nuytsia 12:414 (1999).

Racosperma aureocrinitum (Conn & Tame)

Pedley, comb. nov. §172

Acacia aureocrinita Conn & Tame, Austral.

Syst.Bot.9:851(1996).

Racosperma auricomum (Maslin) Pedley, comb,

nov. §643

Acacia auricoma Maslin, J. Adelaide Bot.

Gard.2:303(1980).

Racosperma auriculiforme (A.Cunn. ex Benth.)

Pedley, Bot. J. Linn. Soc. 92:247 (1986).

§671

Racosperma auripilum (R.S. Co wan & Maslin)

Pedley, comb. nov. §593

Acacia auripila R.S.Cowan & Maslin,

Nuytsia 12:415 (1999).

Racosperma auronitens (Lindl.) Pedley, comb,

nov. §373

Acacia auronitens Lindl., Sketch Veg. Swan

R. xv (1839).

Racosperma ausfeldii (Regel) Pedley, comb. nov.

§457

Acacia ausfeldii Regel, Index Sem. Hort.

Petrop. 106(1867).

Austrobaileya 6 (3): 445-496 (2003)

Racosperma awestonii (R.S. Co wan & Maslin)

Pedley, comb. nov. §499

Acacia awestonii R.S.Cowan & Maslin,

Nuytsia 7:185 (1990).

Racosperma axillare (Benth.) Pedley, Bot. J.

Linn. Soc. 92:248 (1986). §908

Racosperma ayersianum (Maconochie) Pedley,

comb. nov. §840

Acacia ayersiana Maconochie, J. Adelaide

BotGard. 1:182(1978).

Racosperma baeuerlenii (R.T.Baker) Pedley,

Austrobaileya 2:345 (1987). §466

Racosperma baileyanum (F.Muell.) Pedley,

Austrobaileya 2:345 (1987). §30

Racosperma bakeri (Maiden) Pedley,

Austrobaileya 2:345 (1987). §656

Racosperma balsameum (R.S.Cowan & Maslin)

Pedley, comb. nov. §578

Acacia balsamea R.S.Cowan & Maslin,

Nuytsia 12:417 (1999).

Racosperma bancroftiorum (Maiden) Pedley (as

‘bancroftii’), Austrobaileya 2: 345 (1987).

§65

Racosperma barakulense (Pedley) Pedley,

comb. nov. §sub 280

Acacia barakulensis Pedley, Austrobaileya

5:308(1999).

Racosperma barattense (J.M.Black) Pedley,

comb. nov. §469

Acacia barattensis J.M. Black, Trans. & Proc.

Roy. Soc. S. Australia 56:42 (1932).

Racosperma barbinerve (Benth.) Pedley, comb,

nov. §364

Acacia barbinervis Benth., London J. Bot. 1:

326(1842).

R. barbinerve subsp. boreale (Maslin) Pedley,

comb. nov. §364b

Acacia barbinervis subsp. borealis Maslin,

Nuytsia 12: 327(1999).

Racosperma barringtonense (Tindale) Pedley,

comb. nov. §134

Acacia barringtonensis Tindale, Telopea 1:

72(1975).

Pedley, A synopsis of Racosperma

Racosperma basedowii (Maiden) Pedley, comb,

nov. §309

Acacia basedowii Maiden, J. & Proc. Roy.

Soc. New South Wales 53:197 (1920).

Racosperma baueri (Benth.) Pedley,

Austrobaileya 2: 345 (1987). §158

R. baueri subsp. asperum (Maiden & Betche)

Pedley, comb, et stat. nov. §158b

Acacia baueri var. aspera Maiden & Betche,

Census New South Wales Plants 90

(1916).

Racosperma baxteri (Benth.) Pedley, comb. nov.

§354

Acacia baxteri Benth (as ‘Bagsteri’), London

J. Bot. 1:347 (1842).

Racosperma beauverdianum (Ewart & Sharman)

Pedley, comb. nov. §851

Acacia beauverdiana Ewart & Sharman,

Proc. Roy. Soc. Victoria n. ser. 28: 230

(1916).

Racosperma beckleri (Tindale) Pedley, Bot. J.

Linn. Soc. 92:248 (1986). §180

Racosperma benthamii (Meisn.) Pedley, comb,

nov. §545

Acacia benthamii Meisn., in J.G.C.Lehmann,

PI. Preiss. 1:11(1844).

Racosperma betchei (Maiden & Blakely)

Pedley, Austrobaileya 2: 345 (1987). §119

Racosperma bidentatum (Benth.) Pedley, comb,

nov. §316

Acacia bidentata Benth., London J. Bot. X:

333(1842).

Racosperma bifarium (Maslin) Pedley, comb,

nov. §442

Acacia bifaria Maslin, Nuytsia 10:160 (1995).

Racosperma biflorum (R.Br.) Pedley, comb. nov.

§337

Acacia biflora R. Br., in W.T. Aiton, Hortus

Kew. ed 2.5:463 (1813).

Racosperma binatum (Maslin) Pedley, comb,

nov. §417

Acacia binata Maslin, Nuytsia 2: 202 (1978).

Racosperma binervatum (DC.) Pedley,

Austrobaileya2: 345 (1987). §57

453

Racosperma binervium (Wendl.) Pedley, comb,

nov. §775

Mimosa binervia Wendl., Bot. Beobot. 56

(1798).

Racosperma bivenosum (DC.) Pedley, comb. nov.

§220

Acacia bivenosa DC., Prodr. 2:452 (1825).

Racosperma blakei (Pedley) Pedley,

Austrobaileya2: 345 (1987). §778

R. blakei subsp. diphyllum (Tindale) Pedley,

Austrobaileya2: 345 (1987). §778b

Racosperma blakelyi (Maiden) Pedley, comb,

nov. §236

Acacia blakelyi Maiden, J. & Proc. Roy. Soc.

New South Wales 51:246 (1917).

Racosperma blaxellii (Maslin) Pedley, comb,

nov. §420

Acacia blaxellii Maslin, Nuytsia 12: 328

(1999).

Racosperma blayanum (Tindale & Court) Pedley,

comb. nov. §17

Acacia blayana Tindale & Court, Telopea 4:

109(1990).

Racosperma boormanii (Maiden) Pedley, comb,

nov. §150

Acacia boormanii Maiden, J. & Proc. Roy.

Soc. New South Wales 49:489 (1916).

Racosperma botrydion (Maslin) Pedley, comb,

nov. §402

Acacia botrydion Maslin, Nuytsia 4: 30

(1982).

Racosperma brachybotryum (Benth.) Pedley,

comb. nov. §175

Acacia brachybotrya Benth., London J. Bot.

1:347(1842).

Racosperma brachycarpum (Pedley) Pedley,

Austrobaileya 2: 345 (1987). §290

Racosperma brachycladum (W. V.Litzg.) Pedley,

comb. nov. §325

Acacia brachyclada W.V.Litzg., J. Bot. 50: 20

(1912).

Racosperma brachyphyllum (Benth.) Pedley,

comb. nov. §493

Acacia brachyphylla Benth., Linnaea 26: 615

(1855).

454

R. brachyphyllum var. recurvatum (R.S.Cowan

& Maslin) Pedley, comb. nov. §493b

Acacia brachyphylla var. recurvata

R.S.Cowan & Maslin, Nuytsia9:72 (1993).

Racosperma brachypodum (Maslin) Pedley,

comb. nov. §478

Acacia brachypoda Maslin, Nuytsia 7: 225

(1990).

Racosperma brachystachyum (Benth.) Pedley,

Austrobaileya 2:345 (1987). §836

Racosperma bracteolatum (Maslin) Pedley,

comb. nov. §443

Acacia bracteolata Maslin, Nuytsia 12: 329

(1999).

Racosperma brassii (Pedley) Pedley,

Austrobaileya 2:345 (1987). §690

Racosperma brockii (Tindale & Kodela) Pedley,

comb. nov. §712

Acacia brockii Tindale & Kodela, Telopea 5:

62(1992).

Racosperma brownei (Poir.) Pedley,

Austrobaileya 2:346 (1987). §291

Racosperma brownianum (H.L.Wendl.) Pedley,

comb. nov. §934

Acacia browniana H.L.Wendl., Flora 2: 139

(1819).

R. brownianum var. endlicheri (Meisn.) Pedley,

comb, etstat. nov. §934c

Acacia endlicheri Meisn., in J.G.C.Lehmann,

PI. Preiss. 1:21(1844).

R. brownianum var. glaucescens (Maslin)

Pedley, comb. nov. §934e

Acacia browniana var. glaucescens Maslin,

Nuytsia 2:356(1979).

R. brownianum var. intermedium (E.Pritzel)

Pedley, comb. nov. §934d

Acacia strigosa var. intermedia E. Pritzel, Bot.

Jahr.Syst. 35:312(1904).

R. brownianum var. obscurum (A.DC.) Pedley,

comb, etstat. nov. §934b

Acacia obscura A. DC., Mem. Soc. Phys.

Geneve 6:605 (1834).

Austrobaileya 6 (3): 445-496 (2003)

Racosperma brumale (Maslin) Pedley, comb,

nov. §100

Acacia brumalis Maslin, Nuytsia 10: 185

(1995).

Racosperma brunioides (A.Cunn. ex GDon)

Pedley, Austrobaileya2: 346 (1987).§7<57

R. brunioides subsp. graniticum (Pedley)

Pedley, Austrobaileya 2: 346 (1987).

§161b

Racosperma bulgaense (Tindale & S. Davies)

Pedley, comb. nov. §777

Acacia bulgaensis Tindale & S.Davies,

Austral. Syst. Bot. 5:645 (1992).

Racosperma burbidgeae (Pedley) Pedley,

Austrobaileya 2: 346 (1987). §281

Racosperma burdekense (Pedley) Pedley, comb,

nov. §sub 666

Acacia burdekensis Pedley, Austrobaileya 5:

313(1999).

Racosperma burkittii (F.Muell. ex Benth.)

Pedley, comb. nov. §sub 866

Acacia burkittii F.Muell. ex Benth., FI.

Austral. 2:400 (1864).

Kodela & Tindale (1998) and the NSW

contributors to the Flora of Australia treated

Acacia burkittii as a subspecies of A.

acuminata (866). Maslin in WATTLE, citing

an unpublished report on variation in the

‘Acacia acuminata (Jam) group’, treated the

two as distinct species.

Racosperma burrowii (Maiden) Pedley,

Austrobaileya 2:346 (1987). §776

Racosperma buxifolium (A.Cunn.) Pedley,

Austrobaileya 2: 346 (1987). 157

R. buxifolium subsp. pubiflorum (Pedley)

Pedley, Austrobaileya 2: 346 (1987).

§157b

Racosperma bynoeanum (Benth.) Pedley, comb,

nov. §522

Acacia bynoeana Benth., Finnaea 26: 614

(1855).

Pedley, A synopsis of Racosperma

Racosperma caerulescens (Maslin & Court)

Pedley, comb. nov. §61

Acacia caerulescens Maslin & Court,

Muelleria7:131 (1989).

Racosperma caesariatum (R.S.Cowan &

Maslin) Pedley, comb. nov. §496

Acacia caesariata R.S.Cowan & Maslin,

Nuytsia7:210 (1990).

Racosperma caesiellum (Maiden & Blakely)

Pedley, comb. nov. §132

Acacia caesiella Maiden & Blakely, J. & Proc.

Roy. Soc. New South Wales 60:180 (1927).

Racosperma calamifolium (Sweet ex Lindl.)

Pedley, comb. nov. §81

Acacia calamifolia Sweet ex Lindl., Bot. Reg.

10: t. 839 (1824).

Racosperma calanthum (Pedley) Pedley,

Austrobaileya2: 346 (1987). §277

Racosperma calcaratum (Maiden & Blakely)

Pedley, comb. nov. §381

Acacia calcarata Maiden & Blakely, J. Roy.

Soc. W. Australia 13:2 (1928).

Racosperma calcicola (Forde & Ising) Pedley,

Austrobaileya 2: 346 (1987). §606

Racosperma caleyi (A.Cunn. ex Benth.) Pedley,

comb. nov. §117

Acacia caleyi A.Cunn. ex Benth., London J.

Bot. 1:347 (1842).

Racosperma calligerum Pedley, nom. et stat.

nov. §756c

Acacia wickhamii subsp. parviphyllodinea

Tindale, Kodela & D.Keith, FI. Australia

11B: 488 (2001) A. calligera F. Muell, J.

Proc. Linn. Soc., Bot. 3: 141 (1859), pro

syn., nom. invalid.

A. wickhamii subsp. viscidula (F.Muell.)

Tindale, Kodela & D. Keith, FI. Australia

11B: 488 (2001); syn. nov. A. wickhamii

var. viscidula F. Muell., J. Proc. Linn. Soc.

3:141 (1859), syn. nov.

As discussed at some length elsewhere (Pedley

2002), Bentham, in editing Mueller’s (1859)

paper, relegated some of his names to

synonymy. Most of these have since been

validated. Here I have taken up Mueller’s

455

specific epithet but based the name on Acacia

wickhamii subsp. parviphyllodinea, type

material of which has been widely distributed

among Australian herbaria. Acacia wickhamii

subsp. viscidula may warrant formal recognition

but is not specifically distinct from R.

calligerum.

Racosperma calyculatum (ACunn. ex Benth.)

Pedley, Austrobaileya 2: 346 (1987). §768

Racosperma cambagei (R.T.Baker) Pedley,

Austrobaileya 2: 346 (1987). §609

Racosperma camptocladum (C.R.P. Andrews)

Pedley, comb. nov. §215

Acacia camptoclada C.R.P.Andrews, J. W.

Australia Nat. Hist. Soc. 39 (May 1904).

Racosperma campylophyllum (Benth.) Pedley,

comb. nov. §554

Acacia campylophylla Benth., Linnaea 26:

605(1855).

Racosperma cangaiense (Tindale & Kodela)

Pedley, comb. nov. §21

Acacia cangaiensis Tindale & Kodela,

Austral. Syst. Bot. 4: 582 (1991).

Racosperma canum (Maiden) Pedley,

Austrobaileya 2: 346 (1987). §604

Racosperma capillare (A.S.George) Pedley,

comb. nov. §918

Acacia capillaris A.S. George, J. Roy. Soc.

W. Australia 82:69(1999).

Racosperma cardiophyllum (A.Cunn. ex Benth.)

Pedley, comb. nov. §33

Acacia cardiophylla A.Cunn. ex Benth.,

London J. Bot. 1:385(1842).

Racosperma carens (Maslin) Pedley, comb. nov.

Acacia carens Maslin, Nuytsia 10:172 (1995).

Racosperma carneorum (Maiden) Pedley,

comb. nov. §186

Acacia carneorum Maiden (as ‘carnei’), J. &

Proc. Roy. Soc. New South Wales 49:470

(1916).

Racosperma carnulosum (Maslin) Pedley,

comb. nov. §423

Acacia carnulosa Maslin, Nuytsia 12: 331

(1999).

456

Racosperma caroleae (Pedley) Pedley,

Austrobaileya 2:346 (1987). § 799

Racosperma cassiculum (R.S.Cowan & Maslin)

Pedley, comb. nov. § 503

Acacia cassicula R.S.Cowan & Maslin,

Nuytsia7:187(1990).

Racosperma castanostegium (Maslin) Pedley,

comb. nov. §238

Acacia castanostegia Maslin, Nuytsia 12: 332

(1999).

Racosperma cataractae (Tindale & Kodela)

Pedley, comb. nov. § 759

Acacia cataractae Tindale & Kodela, Telopea

5:56(1992).

Racosperma catenulatum (C.T. White) Pedley,

Austrobaileya 2:346 (1987). §843

Racosperma caveale (R.S.Cowan & Maslin)

Pedley, comb. nov. §375

Acacia cavealis R.S.Cowan & Maslin,

Nuytsia 12:454 (1999).

Racosperma cedroides (Benth.) Pedley, comb,

nov. §359

Acacia cedroides Benth., Linnaea 26: 615

(1855).

Racosperma celastrifolium (Benth.) Pedley,

comb. nov. §258

Acacia celastrifolia Benth., London J. Bot.

1:349(1842).

Racosperma celsum (Tindale) Pedley, comb. nov.

§sub 670

Acacia celsa Tindale, Austral. Syst. Bot. 13:

34(2000).

Racosperma centrinervium (Maiden & Blakely)

Pedley, Austrobaileya 2:346. (1987).

§sub 450

The name may be a synonym of R. lineatum,

but the transfer to Racosperma is warranted

pending further investigation.

Racosperma cerastes (Maslin) Pedley, comb,

nov. §387

Acacia cerastes Maslin, Nuytsia 10: 173

(1995).

Austrobaileya 6 (3): 445-496 (2003)

Racosperma chalkeri (Maiden) Pedley, comb,

nov. §74

Acacia chalkeri Maiden, J. & Proc. Roy. Soc.

New South Wales 49:482 (1916).

Racosperma chamaeleon (Maslin) Pedley,

comb. nov. §101

Acacia chamaeleon Maslin, Nuytsia 10: 189

(1995).

Racosperma chapmanii (R.S.Cowan & Maslin)

Pedley, comb. nov. §552

Acacia chapmanii R.S.Cowan & Maslin,

Nuytsia 12:455 (1999).

R. chapmanii subsp. australe (R.S.Cowan &

Maslin) Pedley, comb. nov. §552b

Acacia chapmanii subsp. australis

R.S.Cowan & Maslin, Nuytsia 12: 457

(1999).

Racosperma chartaceum (Maslin) Pedley,

comb. nov. §204

Acacia chartacea Maslin, Nuytsia 8: 293

(1992).

Racosperma cheelii (Blakely) Pedley, comb. nov.

§781

Acacia cheelii Blakely, Proc. Linn. Soc. New

South Wales ser. 2.42:441 (1917).

Racosperma chinchillense (Tindale) Pedley,

Austrobaileya 2:346 (1987). §20

Racosperma chippendalei (Pedley) Pedley,

Austrobaileya 2:346 (1987). §915

Racosperma chisholmii (F.M.Bailey) Pedley,

Austrobaileya 2:346 (1987). §725

Racosperma chrysellum (Maiden & Blakely)

Pedley, comb. nov. §103

Acacia chrysella Maiden & Blakely, J. Roy.

Soc. W. Australia 13:16 (1928).

Racosperma chrysocephalum (Maslin) Pedley,

comb. nov. §336

Acacia chrysocephala Maslin, Nuytsia 2: 304

(1978).'

Racosperma chrysochaetum (Maslin) Pedley,

comb. nov. §718

Acacia chrysochaeta Maslin, Nuytsia 4: 367

(1983).

Pedley, A synopsis of Racosperma

Racosperma chrysopodum (Maiden & Blakely)

Pedley, comb. nov. §505

Acacia chrysopoda Maiden & Blakely, J. Roy.

Soc. W. Australia 13:10 (1928).

Racosperma chrysotrichum (Tindale) Pedley,

comb. nov. §34

Acacia chrysotricha Tindale, Contrib. New

South Wales Natl Herb. 4: 20 (1966).

Racosperma cincinnatum (F.Muell.) Pedley,

Austrobaileya 2: 347 (1987). §673

Racosperma citrinoviride (Tindale & Maslin)

Pedley, comb. nov. §830

Acacia citrinoviridis Tindale & Maslin,

Nuytsia2:86(1976).

Racosperma clandullense (Conn & Tame)

Pedley, comb. nov. § 171

Acacia clandullensis Conn & Tame, Austral.

Syst. Bot.9: 849(1996).

Racosperma clelandii (Pedley) Pedley, comb,

nov. §837

Acacia clelandii Pedley, FI. Australia 1 IB: 488

( 2001 ).

Racosperma clivicola (Pedley) Pedley, comb,

nov. §sub 818

Acacia clivicola Pedley, Contrib. Queensland

Herb. 15:7(1974).

Racosperma clunies-rossiae (Maiden) Pedley,

comb. nov. §133

Acacia clunies-rossiae Maiden, J. & Proc.

Roy. Soc. New South Wales 49:486 (1916).

Racosperma clydonophorum (Maslin) Pedley,

comb. nov. §256

Acacia clydonophora Maslin, Nuytsia 10: 87

(1995).

Racosperma cochleare (Labill.) Pedley, comb,

nov. §544

Mimosa cochlearis Labill., Nov. Holl. PI. 2:

85. t. 234(1807).

Racosperma cochlocarpum (Meisn.) Pedley,

comb. nov. §884

Acacia cochlocarpa Meisn., Bot. Zeitung

(Berlin) 13:10(1855).

R. cochlocarpum subsp. velutinosum (Maslin

& A.R. Chapm.) Pedley, comb. nov.

§884b

Acacia cochlocarpa Meisn. subsp.

velutinosa Maslin & A.R.Chapm., Nuytsia

12:475(1999).

457

Racosperma cognatum (Domin) Pedley, comb,

nov. §464

Acacia cognata Domin, Biblioth. Bot. 89: 260

(1926).

Racosperma colei (Maslin & L. A.J.Thomson)

Pedley, comb. nov. §682

Acacia colei Maslin & L.A.J.Thomson,

Austral. Syst. Bot. 5:737 (1992).

Racosperma colletioides (Benth.) Pedley, comb,

nov. §547

Acacia colletioides Benth., London J. Bot. 1:

336(1842).

Racosperma comans (W.V.Fitzg.) Pedley, comb,

nov. §542

Acacia comans W.V.Fitzg., J. W. Australia

Nat. Hist. Soc. 5 (1904).

Racosperma complanatum (A.Cunn. ex Benth.)

Pedley, Austrobaileya 2: 347 (1987). §631

Racosperma concolorans (Maslin) Pedley,

comb. nov. §3 83

Acacia concolorans Maslin, Nuytsia 12: 334

(1999).

Racosperma concurrens (Pedley) Pedley, Bot.

J. Linn. Soc. 92:248 (1986). §665

Racosperma confertum (A.Cunn. ex Benth)

Pedley, Austrobaileya 2: 347 (1987). §164

Racosperma confluens (Maiden & Blakely)

Pedley, Bot. J. Linn. Soc. 92:248 (1986).

§95

^Racosperma confusum (Merr.) Pedley, Bot. J.

Linn. Soc. 92:248 (1986). f

Acacia confusa Merr., Philip. J. Sci. 5 (Bot.):

27 (1920); Pedley, Contrib. Queensland

Herb. 18: 12 (1975); I. Nielsen, FI.

Malesiana ser. 1.11:61 (1992).

Racosperma congestum (Benth.) Pedley, comb,

nov. §395

Acacia congesta Benth, London J. Bot. 1: 327

(1842).

R. congestum subsp. cliftonianum (W.V. Fitzg.)

Pedley, comb, et stat. nov. §395b

Acacia cliftoniana W.V. Fitzg., J. W. Australia

Nat. Hist. Soc. 10(1904).

458

R. congestum subsp. wonganense (Maslin)

Pedley, comb. nov. §395c

Acacia congesta subsp. wonganensis

Maslin, Nuytsia 12: 338 (1999).

Racosperma conjunctifolium (F. Muell.) Pedley,

Austrobaileya2: 347 (1987). §740

Racosperma connianum (Maslin) Pedley, comb,

nov. §804

Acacia conniana Maslin, Nuytsia 5: 323

(1985).

Racosperma consanguineum (R.S.Cowan &

Maslin) Pedley, comb. nov. §538

Acacia consanginea R.S. Co wan & Maslin,

Nuytsia 10:243(1995).

Racosperma consobrinum (R.S.Cowan &

Maslin) Pedley, comb. nov. § 50 7

Acacia consobrina R.S.Cowan & Maslin,

Nuytsia 7:189(1990).

Racosperma conspersum (F.Muell.) Pedley,

comb. nov. §705

Acacia conspersa F. Muell., J. Proc. Linn. Soc.,

Bot. 3:140(1859).

Racosperma constablei (Tindale) Pedley, comb,

nov. §48

Acacia constablei Tindale, Telopea 1: 429

(1980).

Racosperma continuum (Benth.) Pedley, comb,

nov. §307

Acacia continua Benth., FI. Austral. 2: 322

(1864).

Racosperma convallium (Pedley) Pedley, comb,

nov. §650

Acacia convallium Pedley, Austrobaileya 5:

312(1999).

Racosperma coolgardiense (Maiden) Pedley,

comb. nov. §847

Acacia coolgardiensis Maiden, J. & Proc.

Roy. Soc. New South Wales 53:211 (1920).

R. coolgardiense subsp. effusum (R.S.Cowan

& Maslin) Pedley, comb. nov. §847b

Acacia coolgardiensis subsp. effusa

R.S.Cowan & Maslin, Nuytsia 10: 21

(1995).

Austrobaileya 6 (3): 445-496 (2003)

R. coolgardiense subsp. latius (R.S.Cowan &

Maslin) Pedley, comb. nov. §847c

Acacia coolgardiensis subsp. latior

R.S.Cowan & Maslin, Nuytsia 10: 22

(1995).

Racosperma coriaceum (DC.) Pedley, Bot. J.

Linn. Soc. 92:248 (1986). §591

R. coriaceum subsp. pendens (R.S. Cowan &

Maslin) Pedley, comb. nov. §591 b

Acacia coriacea subsp. pendens R.S.Cowan

& Maslin, Nuytsia 9: 86 (1993).

R. coriaceum subsp. sericophyllum (F.Muell.)

Pedley, comb, et stat. nov. §591 c

Acacia sericophylla F. Muell., J. Proc. Linn.

Soc., Bot. 3:122 (1859).

Racosperma costatum (Benth.) Pedley, comb,

nov. §363

Acacia costata Benth., London J. Bot. 1: 339

(1842).

Racosperma costinianum (Tindale) Pedley,

comb.nov. §136

Acacia costiniana Tindale, Telopea 1: 441

(1980).

Racosperma courtii (Tindale & Herscovitch)

Pedley, comb. nov. §904

Acacia courtii Tindale & Herscovitch,

Telopea 4:115 (1990).

Racosperma covenyi (Tindale) Pedley, comb,

nov. §138

Acacia covenyi Tindale, Telopea 1:432 (1980).

Racosperma cowanianum (Maslin) Pedley,

comb.nov. §473

Acacia cowaniana Maslin. Nuytsia 7: 226

(1990).

Racosperma cowleanum (Tate) Pedley,

Austrobaileya 2:347 (1987). §684

R. oligophlebum (Pedley) Pedley,

Austrobaileya 2:353 (1987), syn. nov.

Racosperma cracente (R.S.Cowan & Maslin)

Pedley, comb. nov. §878

Acacia cracentis R.S.Cowan & Maslin,

Nuytsia 12:476 (1999).

Pedley, A synopsis of Racosperma

Racosperma craspedocarpum (F.Muell.) Pedley,

comb.nov. §842

Acacia craspedocarpa F.Muell., Chemist &

Druggist Australas. 2:73 (1887).

Racosperma crassicarpum (A.Cunn. ex Benth.)

Pedley, Austrobaileya 2: 347 (1987). §669

Racosperma crassistipulum (Benth.) Pedley,

comb.nov. §329

Acacia crassistipula Benth., London J. Bot.

1:326(1842).

Racosperma crassiusculum (H.L.Wendl.)

Pedley, comb, no v. §233

Acacia crassiuscula H.L.Wendl., Comm.

Acac. Aphyll. 5,31 (1820).

Racosperma crassuloides (Maslin) Pedley,

comb.nov. §416

Acacia crassuloides Maslin, Nuytsia 2: 203

(1978).

Racosperma crassum (Pedley) Pedley,

Austrobaileya 2: 347 (1987). §664

R. crassum subsp. longicomum (Pedley) Pedley,

Austrobaileya 2: 347 (1987). §664b

Racosperma cremiflorum (Conn & Tame)

Pedley, comb. nov. § 173

Acacia cremiflora Conn & Tame, Austral.

Syst. Bot. 9: 853(1996).

Racosperma crenulatum (R.S.Cowan & Maslin)

Pedley, comb. nov. §808

Acacia crenulata R.S.Cowan & Maslin,

Nuytsia 12:418 (1999).

Racosperma cretaceum (Maslin & Whibley)

Pedley, comb. nov. §90

Acacia cretacea Maslin & Whibley, Nuytsia

6:27(1987).

Racosperma cretatum (Pedley) Pedley,

Austrobaileya 2: 347 (1987). §659

Racosperma crispulum (Benth.) Pedley, comb,

nov. §330

Acacia crispula Benth., Linnaea 26: 606

(1855).

Racosperma crombiei (C.T. White) Pedley, Bot.

J. Linn. Soc. 92:247 (1986). §184

459

Racosperma cultriforme (A.Cunn. ex G.Don.)

Pedley, Austrobaileya 2: 347 (1987). §153

Racosperma cummingianum (Maslin) Pedley,

comb.nov. §371

Acacia cummingiana Maslin, Nuytsia 10:175

(1995).

Racosperma cuneifolium (Maslin) Pedley,

comb. nov. §398

Acacia cuneifolia Maslin, Nuytsia 12: 339

(1999).

Racosperma curranii (Maiden) Pedley,

Austrobaileya 2: 347 (1987). §809

Racosperma curvatum (Maslin) Pedley, comb,

nov. §553

Acacia curvata Maslin, Nuytsia 2:148 (1977).

Racosperma cuspidifolium (Maslin) Pedley,

comb.nov. §206

Acacia cuspidifolia Maslin, Nuytsia 4: 79

(1982).

Racosperma cuthbertsonii (Luehm.) Pedley,

comb.nov. §823

Acacia cuthbertsonii Luehm., Victorian

Naturalist 13:117(1897).

R. cuthbertsonii subsp. lineare (R.S.Cowan &

Maslin) Pedley, comb. nov. §823b

Acacia cuthbertsonii subsp. linearis

R.S.Cowan & Maslin, Nuytsia 10: 25

(1995).

Racosperma cylindricum (R.S.Cowan &

Maslin) Pedley, comb. nov. §892

Acacia cylindrica R.S.Cowan & Maslin,

Nuytsia 10: 31 (1995).

Racosperma cyperophyllum (F. Muell. ex Benth.)

Pedley, Austrobaileya 2: 347 (1987). §810

R. cyperophyllum var. omearanum (Maslin)

Pedley, comb. nov. §810b

Acacia cyperophylla var. omearana Maslin.

W. Austral. Naturalist 18:186 (1991).

Racosperma dacrydioides (Tindale) Pedley,

comb. nov. §717

Acacia dacrydioides Tindale, Telopea 1: 77

(1975).

460

Racosperma dallachianum (F.Muell,) Pedley,

comb.nov. §901

Acacia dallachiana F.Muell., Fragm. 1: 7

(1858).

Racosperma dangarense (Tindale & Kodela)

Pedley, comb. nov. §50

Acacia dangarensis Tindale & Kodela,

Austral. Syst. Bot. 4:586 (1991).

Racosperma daviesii (Bartolome) Pedley, comb,

nov. §sub 452

Acacia daviesii Bartolome, Austral. Syst. Bot.

15:472(2002).

Racosperma daviesioides (C.A.Gardner) Pedley,

comb.nov. §386

Acacia daviesioides C.A.Gardner, J. Roy. Soc.

W. Australia 27:173(1942).

Racosperma daweanum (Maslin) Pedley, comb,

nov. §829

Acacia daweana Maslin, Nuytsia4: 82 (1982).

Racosperma dawsonii (R.T.Baker) Pedley,

Austrobaileya2: 347 (1987). §480

Racosperma dealbatum (Link) Pedley,

Austrobaileya 2:358 (1987). §52

R. dealbatum subsp. subalpinum (Tindale &

Kodela) Pedley, comb. nov. §52b

Acacia subdealbata subsp. subalpina

Tindale & Kodela, Telopea 9: 319 (2001).

Racosperma deanei (R.T.Baker) Pedley, Bot. J.

Linn. Soc. 92:248 (1986). §40

R. deanei subsp. paucijugum (F.Muell. ex N.A.

Wakef.) Pedley, comb, et stat. nov. §40b

Acacia paucijuga F. Muell. ex N.A. Wakef.,

Victorian Naturalist 72:93 (1955).

Racosperma debile (Tindale) Pedley,

Austrobaileya 2:347 (1987). §16

Racosperma declinatum (R.S.Cowan & Maslin)

Pedley, comb. nov. §495

Acacia declinata R.S.Cowan & Maslin, W.

Austral. Naturalist 18:79 (1990).

Racosperma decorum (Rchb.) Pedley,

Austrobaileya 2:347 (1987). §116

Racosperma decurrens (Willd.) Pedley,

Austrobaileya 2:358 (1987). §49

Austrobaileya 6 (3): 445-496 (2003)

Racosperma deficiens (Maslin) Pedley, comb,

nov. §409

Acacia deficiens Maslin, Nuytsia 12: 340

(1999).

Racosperma deflexum (Maiden & Blakely)

Pedley, comb. nov. §486

Acacia deflexa Maiden & Blakely, J. Roy. Soc.

W. Australia 13:18 (1928).

Racosperma delibratum (A.Cunn. ex Benth.)

Pedley, comb. nov. §736

Acacia delibrata A.Cunn. ex Benth., London

J. Bot. 1:374(1842).

Racosperma delicatulum (Tindale & Kodela)

Pedley, comb. nov. § 744

Acacia delicatula Tindale & Kodela, Telopea

5:58(1992).

Racosperma delphinum (Maslin) Pedley, comb,

nov. §332

Acacia delphina Maslin, Nuytsia 2: 322

(1978).

Racosperma deltoideum (A.Cunn. ex GDon)

Pedley,Austrobaileya2: 315 (1987).§677

Acacia vincentii R.S.Cowan & Maslin,

Nuytsia 7:207 (1990), syn. nov.

Acacia vincentii R.S.Cowan & Maslin (no. 614)

is based on the aberrant specimen of

Racosperma deltoideum noted by Pedley

(1987a).

R. deltoideum subsp. amplum (R.S.Cowan &

Maslin) Pedley, comb. nov. §617b

Acacia deltoideum subsp. ampla R.S.Cowan

& Maslin, Nuytsia 7: 206 (1990)

Racosperma demissum (R.S.Cowan & Maslin)

Pedley, comb. nov. § 83 3

Acacia demissa R.S.Cowan & Maslin,

Nuytsia 10:25 (1995).

Racosperma dempsteri (F. Muell.) Pedley, comb,

nov. §202

Acacia dempsteri F.Muell., Fragm. 11: 65

(1879).

Racosperma densiflorum (Morrison) Pedley,

comb.nov. §575

Acacia densiflora Morrison, Scott. Bot. Rev.

1:96(1912).

Pedley, A synopsis of Racosperma

Racosperma denticulosum (F.Muell.) Pedley,

comb.nov. §893

Acacia denticulosa F.Muell., Fragm. 10: 32

(1876).

Racosperma dentiferum (Benth.) Pedley, comb,

nov. §392

Acacia dentifera Benth., Botanist 4: t. 179

(1848).

Racosperma depressum (Maslin) Pedley, comb,

nov. §939

Acacia depressa Maslin, Nuytsia 1: 422

(1975).

Racosperma dermatophyllum (Benth.) Pedley,

comb.nov. §433

Acacia dermatophylla Benth., FI. Australia

2:346(1864).

Racosperma desertorum (Maiden & Blakely)

Pedley, comb. nov. §891

Acacia desertorum Maiden & Blakely, J. Roy.

Soc. W. Australia 13: 24 (1927).

R. desertorum var. nudipes (R.S.Cowan &

Maslin) Pedley, comb. nov. §891 b

Acacia desertorum var. nudipes R.S.Cowan

& Maslin, Nuytsia 10: 34 (1995).

Racosperma deuteroneurum (Pedley) Pedley,

Austrobaileya2: 347 (1987). §77

Racosperma diaphanum (R.S.Cowan & Maslin)

Pedley, comb. nov. §421

Acacia diaphana R.S.Cowan & Maslin,

Nuytsia 12:420 (1999).

Racosperma diaphyllodineum (Maslin) Pedley,

comb.nov. §415

Acacia diaphyllodinea Maslin, Nuytsia 2:

206(1978).'

Racosperma dictyoneurum (E. Pritzel) Pedley,

comb. nov. §500

Acacia dictyoneura E.Pritzel, Bot. Jahr. Syst.

35:303(1904).

Racosperma dictyophlebum (F.Muell.) Pedley,

Austrobaileya 2: 347 (1987). §269

Racosperma didymum (A.R.Chapm. & Maslin)

Pedley, comb. nov. §229

Acacia didymum A.R.Chapm. & Maslin,

Nuytsia 8:268 (1992).

461

Racosperma dielsii (E.Pritzel) Pedley, comb. nov.

§561

Acacia dielsii E.Pritzel, Bot. Jahr. Syst. 35:

294(1904).

Racosperma dietrichianum (F.Muell.) Pedley,

Austrobaileya 2: 347 (1987). §191

Racosperma difficile (Maiden) Pedley,

Austrobaileya 2: 348 (1987). §692

Racosperma difforme (R.T.Baker) Pedley, comb,

nov. §110

Acacia dijformis R.T.Baker, Proc. Linn. Soc.

New South Wales 22:154 (1897).

Racosperma dilatatum (Benth.) Pedley, comb,

nov. §366

Acacia dilatata Benth., Linnaea 26: 608

(1855).

Racosperma dimidiatum (Benth.) Pedley,

Austrobaileya 2: 348 (1987). §657

Racosperma diminutum (Maslin) Pedley, comb,

nov. §408

Acacia diminuta Maslin, Nuytsia 12: 342

(1999).

Racosperma disparrimum (M.W. McDonald &

Maslin) Pedley, comb. nov. §sub 670

Acacia disparrima M.W.McDonald &

Maslin, Austral. Syst. Bot. 13:46 (2000).

R. disparrimum subsp. calidestre (M.W.

McDonald & Maslin) Pedley, comb. nov.

§sub 670

Acacia disparrima subsp. calidestris

M.W.McDonald & Maslin, Austral. Syst.

Bot. 13:52(2000).

Racosperma dissimile (M.W.McDonald)

Pedley, comb. nov. §sub 691

Acacia dissimilis M.W.McDonald, Austral.

Syst. Bot. 16:147. t.6 (2003).

Racosperma dissonum (R.S.Cowan & Maslin)

Pedley, comb. nov. §572

Acacia dissona R.S.Cowan & Maslin,

Nuytsia 10:209(1995).

R. dissonum var. indolorium (R.S.Cowan &

Maslin) Pedley, comb. nov. §572b

Acacia dissona var. indoloria R.S.Cowan &

Maslin, Nuytsia 10: 211 (1995).

462

Racosperma distans (Maslin) Pedley, comb. nov.

§794

Acacia distans Maslin, Nuytsia 4: 386 (1983).