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AUSTROBHILEYA
A Journal of Plant Systematics
Queensland Herbarium
Queensland Department of Environment and Heritage
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Editorial Committee E.M. Ross (editor) R.J.F. Henderson (technical advisor)
Word Processing Cover design & Graphics Y.C. Smith W.A. Smith
Austrobaileya Vol. 1, No. | was published on | December 1977
Vol. 3, No. 3 was published on 27 September 1991 Austrobaileya is published once per year. Exchange: This journal will be distributed on the basis of exchange.
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ISSN 0155-4131 ©Queensland Government 1992
Austrobaileya is the journal of the Queensland Herbarium and is devoted to publication of results of sound research and of informed discussion on plant systematics, with special emphasis on Queensland plants.
_ Opinions expressed by authors are their own and do not necessarily represent the policies or views of the Queensland Herbarium.
Austrobaileya 3(4): 569-766 (1992)
CONTENTS
A Taxonomic Revision of Cerbera L. alae in Australia and Papuasia Paul I. Forster 5! , Ss te, Eee nee
A New Species of icine is Rich. c IRS ERaEE from North i eeasar einai B, Gray..
Studies in Australian Grasses: 5* New Species of and New Combinations for Queensland Panicoid Grasses Bryan K. Simon .. ‘a
Perotis Aiton (Gramineae) in Australia and Southeast Asia J.F. Veldkamp & H. van Steenbergen ro,
Studies in Euphorbiaceae A.L. Juss., sens. lat. 2*. A Revision of Neoroepera Muell. Arg. & F. Muell, (Oldfieldioideae Kohler & Webster, Caletieae Muell. Arg.)
Rodney J.F. Henderson
Taxonomic Studies on the Genus Hoya R. Br. (Asclepiadaceae) in Papuasia,
{-5 Paul I. Forster & David J. Liddle... ..
The Genus Leptospermum Forst. et Forst. f. (Myrtaceae) in Northern Australia and Malesia A.R. Bean... .. ....
The Circumscription of Adiantum diaphanum Blume (Adiantaceae), The filmy Maidenhair Fern Peter D. Bostock
Notes on Lycopodiella Holub in North-eastern Queensland R.J. Chinnock ES win kis itd aay) Yael me eae
Studies in Australian Grasses 6*. Alexfloydia, Cliffordiochloa and Dallwatsonia, three New Panicoid Grass Genera from Eastern Australia BYvatt I SIMO . 5. se cud lee tn, de fate ete re Le fw et ala a
Review of the Genus Oldenlandia L. anneal) and Related Genera in Australia AVA) PS -ETATOBEE lot cies ete phe nto eels miserseoe ote Lange com oma reine oa
A New Paspalum L. (Gramineae) from New Caledonia and Vanuatu Bernard Toutain . en eee
Five New Species of Plectranthus L. Herit cL SACRE from Queensland Paul I. Forster oe eek re ee ee ie el eee
Notes on Australian aaah cine eisai J.F. Veldkamp ‘7
Acacia pedleyi (Fabaceae, Mimosoideae), A New Species from Central-Eastern Queensland Mary D. Tindale & Phillip G. Kodela
Chromosome numbers of Australian species of Ipomoea L. atuiiauante, D.E. Yen, P.M. Gaffey & D.J. Coates J income 'y pes cunts tun sneuaed
569
581
385
609
615
627
643
661
665
669
683
723
7129
741
745
749
(continued)
sarang”
RRS yr ne ey inh re cry unt te hci Manat SE nage Scan mn Re oc amaaene ect nem rere AA
ee ne
NOTES
Lectotypification of Lepinia solomonensis Hemsley (Apocynaceae) with notes on distribution Paul I. Forster
Parsonsia alboflavescens (Dennst.) Mabb., (Apocynaceae), a new record from Northern Territory, Australia Paul I. Forster
Notes on the Naturalised Flora of Queensland, 2 Paul I. Forster 2 oa sabe oS outro Pesos
Book Review
757
159
76] 765
Austrobaileya 3(4): 569-579 (1992) 569
A TAXONOMIC REVISION OF CERBERA L. (APOCYNACEAE) IN AUSTRALIA AND PAPUASIA
Paul I. Forster
Queensland Herbarium, Meiers Road, Indooroopilly, Qld 4068, Australia
Summary
The genus Cerbera L. (Apocynaceae) is revised for Australia and Papuasia (Irian Jaya, Papua New Guinea and Solomon Islands). Four species are recognised and described with a key for their identification given. C. manghas L. and C. floribunda Schumann are widespread in the region whereas C, inflata S.T. Blake and C. durmnicola sp. nov. are endemic to Australia. Ethnobotanical use of C. floribunda and C. manghas in the region is reviewed.
Introduction
The genus Cerbera was described by Linnaeus (1753) who recognised three species, namely C. manghas L. C. thevetia L. and C. ahouai L. C. thevetia and C. ahouai are now considered to be Cascabela thevetia (L.) Lippold (syn. Thevetia peruviana (Pers.) Schumann) and Thevetia ahouai (L.) DC. respectively. Cerbera manghas 1s now widely considered the lectotype of Cerbera (Lippold 1980; Boiteau 1981; Smith 1988). Subsequent to Linnaeus, additional species have been added to Cerbera, and until recently eight or nine species were considered to belong in the genus (Lippold 1980, Smith 1988). However, Lippold (1980) in her review did not refer to Markgraf (1972) or Fosberg et al. (1977) wherein several of these species were referred to Kopsia Blume or Neisosperma Raf. The genus Cerbera up to now therefore comprises C. manghas, C. floribunda Schumann, C. odollam Gaertner, C. dilatata Markgraf, C. inflata S.T. Blake, and C. micrantha (Kan- chiru) Pichon.
As defined by Lippold (1980), Boiteau (1981) and Smith (1988), Cerbera is most closely allied to genera such as Cerberiopsis Vieillard ex Pancher & Sebert from New Caledonia and Thevetia L. and Cascabela Rafinesque from South America. It differs from Cerberiopsis in the indehiscent, fleshy polycarpous fruit lacking albumen (Boiteau 1981), from Thevetia most noticeably in the lack of a floral disk, the indehiscent, fleshy polycarpous fruit with a fleshy exocarp, reticulate spongy mesocarp and woody endocarp, and from Cascabela in the hypocrateritorm corolla, lack of scales in the corolla throat, lack of a floral disk and the woody endocarp (Lippold 1980).
The only previous taxonomic accounts of Cerbera in Australia have been by Bentham (1869) and Bailey (1900) who both recognised only a single species from the area, C. manghas (incorrectly as C. odollam Gaertner). C. floribunda Schumann and C. manghas L. were both recognised for New Guinea by Markegraf (1926, 1927), and the former for Australia by White (1933). Blake (1948) subsequently described a new species from north-east Queensland, firstly under the illegitimate C. dilatata S.T. Blake but later renaming it as C. inflata (Blake 1959). Up to the present there has been three described species recognised for the region although in many Australian herbaria there existed considerable undetermined material which has proved to contain collections of an undescribed species from Queensland.
As a precursor to my treatment of the genus in Volume 28 of ‘Flora of Australia’, the present revision 1s presented to enable description of the unnamed species from Australia, describe and provide a key for identification of all recognised species, and to typify the names of taxa where required. In addition, I have appended notes on local common names and ethnobotanical use, primarily of C. floribunda and C. manghas in Papuasia.
Materials and Methods
This revision is based on herbarium specimens in AD, BO, BRI, BSIP, CANB, CBG, DNA, MEL and QRS (herbarium acronyms as in Holmeren ef a/. 1990), and on plants observed in the field in north Queensland and Solomon Islands. The generic description is largely adapted from Smith (1988) who gave a good overview of the
GSES
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meme
570 Austrobaileya 3(4): 1992
important features. Species descriptions have been drawn up from dried material or material reconstituted by boiling in detergent and water; however, the descriptions of C. manghas and C. floribunda are supplemented by observations of fresh and spirit preserved material. With the distribution maps, those for C. manghas and C. floribunda include supplementary specimen data presented by Markgraf (1927).
Taxonomic Treatment
Cerbera L., Sp. ‘Pl. 208 (1753). Lectotype: C. manghas L. (fide Pichon, Notul. Syst. (Paris) 13: 221 (1948)).
Seem., Fl. Vit. 157 (1866); Pichon, Notul. Syst. (Paris) 13: 221 (1948); Markgraf, FI. Madagasc. fam. 169: 156 (1976): Lippold, Feddes Repert. 91: 51 (1980); Boit., Fl. Nouv.-Caled. et Dépend. 10: 212 (1981); A.C. Smith, Fl. Vit. Nov. 4: 89 (1988).
Perennial shrubs or trees, latex white; foliage glabrous, drying black. Leaves spirally arranged on stems, clustered at stem apices, petiolate; lamina coriaceous, entire or sinuate, secondary lateral nerves spaced, connected by a marginal nerve; lacking colleters at lamina base. Inflorescences terminal, pedunculate, laxly cymose, glabrous, bracteate. Flowers pedicellate. Calyx deeply divided, lobes + free, caducous, lacking colleters. Corolla hypocrateriform; tube glabrous externally, internally with or without simple indumentum, swollen about stamens, somewhat constricted above stamens; lobes ovate, obovate, obtuse or emarginate, sinistrorse in bud. Stamens inserted in upper part of tube; filaments short (1-2 mm long), slender, glabrous; anthers obliquely ascending, lanceolate, contiguous to style-head with filiform apical appendages; disk lacking. Ovary bicarpellate, each carpel usually with 4 biseriate ovules; style-head composed of 2 annular swellings surmounted by 2 apical, conical, obtuse, thick, appressed stigmatic appendages. Fruits apocarpous; mericarps drupaceous; exocarp fleshy: mesocarp reticulate-spongy with numerous fibers independent of the endocarp: endocarp lignifed, smooth or rugose- striate externally, with an apical wing; seeds | per locule, compressed, ellipsoid or obloid, not or narrowly winged, hilum large, lacking a coma.
Seven species distributed in Madagascar, Indian Ocean islands, Asia, Malesia, Melanesia and Australia. Four species in Papuasia and Australia.
Key to species of Cerbera in Australia and Papuasia
1. Leaf lamina with tertiary venation obscure below; flowers numerous in
cyme, generally more than 50... Rarer 2 Leaf lamina with tertiary venation reticulate and prominent below: flowers few in cyme, generally less than 30... .... cme ce seas ee ees iy alae 18, GA 3
2. Leaf lamina with 13-20 secondary veins below; corolla tube 8-12 mm
long .... 1, C. floribunda
Leaf lamina with 33-37 } secondary veins below: corolla 15-18 mm LONE? wou Vy ae . sige nt Voltn nes etn dy ES Inflate
3. Leaf lamina with 25-32 secondary veins below; corolla with red centre, tube 25-35 mm long... . 3..¢. manghas
Leaf lamina with 14-18 secondary veins below: corolla lacking red centre, tube 10-11 mm long ....... -...... 4, C, dumicola
1. Cerbera floribunda Schumann in Schumann & ‘Helleai. Fl. Kais. Wilh. Land 111 (1899). Type: Papua New Guinea. WeEsT SEPIK PROVINCE: Augusta fluss, Hollrung 849 (lecto (here designated): K (photo! BRI); isolecto: BO!, L (photo! Ban Papua New Guinea. 1887, A¢4. Hollrung 717 (lectopara: BO!, K. (photo! BRI
|Cerbera odollam auct. non Gaertner: S. Moore, J. Bot. 61, suppl. 32 (1923)).
Schumann, Notizbl. Bot. Gart. Berlin-Dahlem 2: 139 (1898): Boerl., Handl. FI. Ned. Ind. 2(2): 394 (1899); Schumann & Lauterb., Fl. Schutzgeb. Sudsee 505
Forster, Cerbera 57]
(1901); Markeraf, Nova Guinea 14: 284 (1926); Bot. Jahrb. Syst. 61: 198 (1927); C.T. White, Contr. Arnold Abor. 4: 92 (1933); van Royen et a/., Manual Forest Trees Papua & New Guinea, Part 9. Apocynaceae 22-23 (1964); D. Foreman, Checkl. Vasc. Pl. Bougainville. Bot. Bull. No: 5: 102-103 (1971).
Illustrations: van Royen et a/., Manual Forest Trees Papua & New Guinea, Part 9. PPOs CLOT 23 (1964); Foreman, Checkl. Vasc. Pl. Bougainville. Bot. Bull. No. 5: 103 (1971).
Tree to 30 m high, latex white; foliage and inflorescence glabrous. Bark brown to black erey; sap wood white, heart wood white. Leaf lamina lanceolate-elliptic, 9-24 cm long, 2-5 cm wide, discolorous, margins entire and not sinuate; upper surface dark glossy green, venation obscure; lower surface pale green, secondary veins 13-20 per side of midrib, tertiary venation obscure; tip acute to short acuminate; base cuneate; petiole 30-40 mm long, 1.8-—2.0 mm wide. Inflorescence a much branched cyme up to 17 cm long; peduncle 4-6 cm long; flowers numerous with generally over 50 present. Flowers 18-20 mm long, c. 5 mm diameter, sweetly scented; pedicels 2-3 cm long. Sepals lanceolate-ovate, 5.0~6.4 mm long, 3-5 mm wide. Corolla primarily white, but sometimes pinkish-white internally towards tips of lobes and yellowish at bottom of lobes; tube 8— 12 mm long, 2.7-3.0 mm diameter, white, constricted above anthers, glabrous internally; lobes lanceolate, 7-10 mm long, 1.8-—2.0 mm wide, light green, glabrous. Stamens inserted in upper 3 mm of tube; anthers 1.2-1.3 mm long, c. 0.7 mm wide. Fruit obloid-globose, 9-11 cm long, 5.0-5.5 cm wide, 4.5—4<8 cm thick, pomted at end away from pedicel, blue when ripe. Fig. 1D. as
%
Selected specimens: Irian Jaya. Sidei, c. 50 km W of Manokwari, Wwanggin BW5753 (CANB ex L); Japen Island, Soemberbaba, Jul 1961, Koster BWI1159 (BO,BRI-ex L); Bernhard Camp, Idenburg River, May 1939, Brass 14109 (BO,BRI ex A). Papua New Guinea. MANUS PROVINCE: Rambutyo Island, Peninsula Admin. Centre, 2°18’S 147°47’E, Mar 1981, Kerenga & Croft LAE77382 (BRI). EAST Sepik PROVINCE: Kankanaman, 4°05’S, 144°0S’E May 1969, Afillar NGF37518 (BRI). NEW BRiTAIN PROVINCE: Pirtloma Village, 6°06’S, 150°45’E, Mar 1965, Sayers NGF29192 (BRI), NEw IRELAND PRoviNncE: Inland from Lavongai, c. 26 miles [43.3 km] from Kavieng, 2°46’S, 151°02’E, Jan 1967, Coode et al. NGF29603 (BO,BRI). WESTERN HIGHLANDS PROVINCE: 6 miles [10 km] from Lake Kopiago on Paga Hill road, 5°22’S, 142°33’E, Nov 1968, Galore & Vandenburg NGF41029 (CANB). EASTERN HIGHLANDS PROVINCE: Okasa, 10 miles [16 km] SW of Okapa, May 1967, Womersley NGF24924 (BRI). MOROBE PROVINCE: Wagau, 6°50’S, 146°50’E, Jan 1965, Sayers NGF21566 (BRI,CANB). MADANG PROVINCE: Near Usino Village, Ramu Valley, Jul 1955, Hoogland 5016 (BRI,CANB). WESTERN PROVINCE: Dagwa, Oriomo River, Feb-Mar 1934, Brass 5991 (BO,BRI ex A). CENTRAL PROVINCE: Kagi Gap area, Kokoda trail, 9°09’S, 147°43’E, Sep 1973, Croft & Lelean NGF34727 (BRI). GULF PROVINCE: West bank, junction of Vailala & Lohiki Rivers, Jan 1966, Schodde 4311 & Craven (BRI,CANB). NORTHERN PROVINCE: Lower W slopes of Topographers, Aug 1954, Saunders 44 (BRI,CANB). MILNE BAY PROVINCE: Normanby Island, near Miadeba airstrip, 9°50’S, 1S0°55’E, Nov 1976, Croft et al. LAE68858 (BRI). Solomon Islands. Santa Ysabel, Bogotu Peninsula, near Koloajoa Village, Mar 1964, Whitmore BSIP4110 (BSIP). Australia. Queensland. Cook District: Daintree, Jul 1943, Blake 14992 (BRI,CANB); V.C.L. Noah, Oliver Creek, 16°10’S, 145°25’E, Nov 1978, Gray 1095 (QRS); Porn. 62, Alexandra, 16°10’S, 145°25’E, Dec 1972, Hyland 6606 (QRS).
3 3
Distribution and habitat: Widespread in New Guinea and Solomon Islands, but in Australia it is restricted to a small area near Cairns (Map 1). Plants grow along creeks or in marshy areas and are always found near permanent water.
Notes: C. floribunda is a distinctive species and the large blue fruits are often a feature of the forest floor where this species may be locally common. In Australia it is only likely to be confused with C. inflata, which is restricted to higher altitudes on the Atherton Tableland region. I have selected Hol/rung 849 as lectotype from the two syntypes, as it is more widely represented in herbaria and has a more precise collection locality than Hollrung 717.
Local names and ethnobotanical use: A listing of local names with dialects and region of origin 1s given in Table 1. The wood is used for mouldings and interior finishings in Bougainville (Foreman 1971) and other parts of Papua New Guinea, but is susceptible to blue stain fungal attack (van Royen et al. 1964). In Solomon Islands the wood is used for carving and medicine (Henderson & Santalau 491 (BSIP)). Medicinal application is for aches and sores with the leaves heated in a fire, after which they are rubbed on the sore area or the person lies down on the warm leaf.
Conservation status: Not rare or endangered. Conserved in Cape Tribulation National Park north of the Daintree River in north Queensland.
Ber ere Ene hh oes a ao ono eR REE ERT
372
Table 1. Local names for Cerbera floribunda.
Austrobaileya 3(4): 1992
Name Dialect & Region Voucher
Agar Onijob Hoogland 4603 (CANB) Aimalua Kwara’ae; S.I. Gaful et al. BSIP16430 (BSIP) Aitongatonga Kwara’ae; S.L Whitmore BSIP921 (BSIP) Babal Biak; Irian Jaya Koster BW11159 (BRI) Beembin Wagau™; Morobe Sayers NGF21566 (CANB) Biegbau Haltam; Irian Jaya Moll BW15725 (BO)
Brebong Kemtoek; Irian Jaya Jwanggin BW9121 (BO) Djokanabaiu Minufia; Northern Saunders 44 (BRI)
Ekwa Onjob; Northern Saunders 44 (BRI)
Grey Milkwood Australian; Cook Cause et al. (1989)
Iona Akuna; Madang Smith NGFI1088 (BRD)
Kartwi Pom; Irian Jaya Schram BW15046 (BO)
Milky Pine Atherton*; Cook personal obs.
Nibwafrim Amberbaken; Irian Jaya Koster BW6743 (BO) Ocro-Oego Kapaukoe; Irian Jaya Vink & Schram BW8713 (BO) Patega Usino; Madang Hoogland 5016 (BRI)
Pea,paa Maiama*™; Morobe Ridsdale NGF31684 (BRI) Rubbertree Australian; Cook Cause et al. (1989)
Sebokko Manikiang; Irian Jaya Iwanggin BW5753 (CANB) Sowoek Mooi; Irian Jaya Moll BW11727 (BO) Sungwiau Waskuk; Sepik Hoogland 10064 & Craven (BRI) Supuk Bembi; Madang Hoogland 5016 (BRI)
Tarau Subdidi*; New Britain Frodin NGF26589 (BRI)
Tero Rawa; Madang Hoogland 5016 (BRI) Totongwala Kwara’ae; S.I. Whitmore BSIP4110 (BSIP) Towl Jal; Madang Hoogland 5016 (BRI)
Vao Marovo; S.I. Chapman BSIP428 (BSIP)
* indicates locality name only, dialect not indicated. Abbreviation: S.I, = Solomon Islands.
2. Cerbera inflata S.T. Blake, Proc. Roy. Soc. Queensland 70: 33 (1959). Cerbera dilatata S.T. Blake, Proc. Roy. Soc. Queensland 59: 161 (1948), nom. illeg.; non Markgraf (1927). Type: Australia, Queensland. Cook DiIsTRicT: Near Goldsborough, Upper
Mulgrave River, 28 July 1943, S.7. Blake 15033 (holo: BRI!; iso: CANB)). _ [Cerbera manghas auct. non L.: C.T. White, Contr. Arnold Arbor. 4: 92 (1933)].
Tree to 30 m high, latex white; foliage and inflorescence glabrous. Bark grey, pustular with long fissures; sap wood white, heart wood white. Leaf lamina lanceolate-elliptic, discolorous, margins entire and not sinuate; upper surface dull green, venation obscure; lower surface pale green, secondary veins 33-37 per side of midrib, tertiary venation obscure; tip short acuminate; base cuneate; petiole 20-37 mm long, c. 2 mm diameter. Inflorescence a much branched cyme, up to 15 cm long; peduncles up to 5 cm long; flowers numerous, with generally more than 50 present. Flowers 18-27 mm long, 10- 13 mm diameter, sweetly scented; pedicels 11-15 mm long. Sepals lanceolate, 4-8 mm long, 1.5-3.0 mm wide. Corolla white to cream-pink; tube 15-18 mm long, 1-2 mm diameter, slightly constricted above anthers, glabrous internally; lobes obovate, c. 6 mm long, 3-4 mm wide, glabrous. Stamens inserted 2.2-2.5 mm from top of tube; anthers 2.5-2.8 mm long, 1.7—1.8 mm wide. Fruit ellipsoid, somewhat pointed at end away from pedicel, 6-7 cm long, 3.0-3.2 cm wide, 2.0-2.5 cm thick, purple when ripe. Fig. 1A.
Selected specimens. Australia. Queensland. Cook Disrrict: Kulara, 15 miles [25 km] WNW of Atherton, May 1921, Hamilton [AQ212353] (BRI); Atherton, Jan 1950, Webb 5118 (CANB); Lake Eacham, 17°17’S, 145°37’E,
Forster, Cerbera
Maps 1-2. Distribution in Australia and Papuasia of Cerbera spp.: 1. C. floribunda. 2. C. manghas.
573
SOYA A bra commana end ala Se
wes tal i le)
mene ye
km
574 Austrobaileya 3(4); 1992
Feb 1956, Dansie C36 (CANB); Juara Creek area, Danbulla, Aug 1948, Smith 3734 (BRD); Danbulla, near Atherton, Nov 1942, Blake {4749 (BRI,DNA); S.F.R. 185, Downfall L.A., 17°10’S, 145°35’E, Jul 1971, Sanderson 28 (QRS); S.F.R. 185, Danbulla, 17°10’S, 145°37E, Apr 1971, Stocker 673 (BRI,CANB,QRS); Gadgarra Reserve, Atherton, Jul 1929, Kajewski 1141 (BRI); S.F.R. 700, Gadgarra, Gillies L.A., 17°13’S, 145°42’E, Dec 1988, Hyland 13786 (QRS); S.F.R. 310, Caribou L.A., Dec 1981, Hyland 11414 (QRS); Yungaburra, Jan 1918, White [AQ212359] (BRI); near Malanda, Sep 1943, Blake 15249 (BRI,CANB); Mt Bartle Frere, north peak, Aug 1943, Blake 15244 (BRI); Mt Toressa, 1904, Bailey [AQ212354] (BRI); S.F.R. 310, Windin L.A., 17°21’S, 145°45’E, Aug 1979, Moriarty 2653 (QRS); S.F.R. 310, Bora L.A., 17°21’S, 145°46’E, Nov 1981, Gray 2267 (QRS); S.F.R. 755, Barong L.A., 17°32’S, 145°52’E, 1980, Gray 1727 (QRS); Jago, near Innisfail, Sep 1943, Blake 15270 (BRI); Babinda, Jul 1943, Blake 15024 (BRI,CANB); Deeral, near Babinda, Jul 1943, Blake 14970 (BRI).
Distribution and habitat: Restricted to north-east Queensland (Map 3). Plants grow in rainforest, generally away from permanent water.
Notes: As noted by Blake (1948), C. inflata is allied to C. floribunda but differs in the somewhat narrower leaves with more lateral nerves, the smaller flowers and the smaller ellipsoid fruit.
Local names: “Milky Pine” - Blake 14749 (BRD; “Joojooga” - Batley [AQ212354] (BRI); “Grey Milkwood” or “Rubbertree” — Cause et a/. (1989).
Conservation status: Widespread in north-east Queensland and not considered rare or endangered by the current author, despite the listing in Thomas and McDonald (1989).
20
Map 3. Distribution of Cerbera inflata @ and C. dumicola ©.
Forster, Cerbera S75
3. Cerbera manghas L., Sp. Pl. 208 (1753); Tanghinia manghas (L.) G. Don, Gen. hist. 4: 98 (1838). Type: description of Osbeck (Dagbok 6fwer en Ostindisk Res., ca 91 (1757)), cited in L., Sp. Pl. 208 (1753); fide Fosberg in Boiteau (1981).
J. Forster, Fl. Ins. Austr. Prodr. 19 (1786); Schumann, Notizbi. Bot. Gart. Berlin- Dahlem 1: 55 (1895); Merr. Interpret. Rumph. Herb. Amb. 432 (1917); Markeraf, Nova Guinea 14: 284 (1926); Domin, Biblioth. Bot. 89: 522 (1928); Markeraf, Bot. Jahrb. Syst. 61: 197 (1927); Guillaumin, J. Arnold Arb. 13: 18 (1932); Boit., Fl. Nouv.-Caléd. et Dépend. 10: 214 (1981); Fosb. & Sach. in D.J. Carr, Sydney Parkinson 186 (1983); A.C. Smith, Fl. Vit. Nov. 4: 89 (1988).
Cerbera odollam var. mugfordii Bailey, Queensland Agric. J. 3: 282 (1898) (as ‘mugforar), C. manghas var. mugfordii (Bailey) Domin, Biblioth. Bot. 89: 522 (1928). Type: [Mourilyan Harbour, W. Mugford| [?4Q332823] (holo: BRI).
[Cerbera odollam auct. non Gaertner: Benth., Fl. Austral. 4: 306 (1869): Bailey, Queensland fl. 3: 981 (1900)).
Illustrations: Sims, Bot. Mag. 43: t. 1845 (1816); K.A.W. Williams, Native PI. Queensland 1: 67 (1979), 3: 63 (1987); Fosb. & Sach. in D.J. Carr, Sydney Parkinson pl. 177 (1983); A.C. Smith, Fl. Vit. Nov. 4: 90, fig. 38 (1988).
Small tree to 12 m high; latex white; foliage and inflorescence glabrous. Bark light grey, scaly; sap wood white, heart wood white. Leaf lamina elliptic-obovate, 15-25 cm long, 4-7 cm wide, discolorous, margins entire and not sinuate; upper surface dark glossy green, venation + obscure; below pale green, secondary veins 25~32 per side of midrib, tertiary venation reticulate and prominent; tip acute, short acuminate or long acuminate; base cuneate; petiole 15-62 mm long, c. 1 mm wide. Inflorescence a little-branched cyme to 20 cm long; peduncles up to 8 cm long; flowers few, with generally less than 30 present. Flowers 30-40 mm long, 20-35 mm diameter, sweetly scented; pedicels 10-20 mm long. Sepals lanceolate to lanceolate-ovate, 12-25 mm long, 4-9 mm wide. Corolla white with red ‘eye’ at top of tube; tube 22—35 mm long, 2.0-2.5 mm diameter, constricted above anthers near top, with sparse to dense indumentum internally; lobes ovate to obovate, 18-26 mm long, 6-10 mm wide. Stamens inserted c. 2 mm below top of tube; anthers 1.8-2.0 mm long, 1.0-1.2 mm wide. Fruit ovoid-globose, 6-7 cm long, 3.5-4.0 cm wide; 3.5-4.0 cm thick, red when ripe. Fig. 1B, E-G, J-L.
Selected specimens. Irian Jaya. Batanta Island, W of Sorong, Marchesa Bay, Amdoei Village, Apr 1954, var Royen 3545 (BO,BRI ex L); Adi Island, Feb 1961, Afo// BW9827 (BRI, CANB). Papua New Guinea. MANUS PROVINCE: Pelikawa, 2°07’S, 146°44’E, Jun 1971, Stone & Streiinann LAE53788 (BRI). NEw BRITAIN PROVINCE: Torlu River, 6°00’S, {51°02’E, Mar 1965, Sayers NGF24208 (BRI). NEw IRELAND PROVINCE: Kavieng, Oct 1964, Millar NGF23816 (BRI). BOUGAINVILLE PROVINCE: Karngu, Buin, Oct 1930, Kajewski 2239 (BRI ex A). EAST SEPIK PROVINCE: Cape Wom International Park, c. 8 km NW of Wewak town, 3°35’S, 143°35’E, Nov 1976, Wiakabu & Yefle LAE70329 (BRI). MOROBE PROVINCE: Markham River mouth, W of Lae, 6°45’S, 147°00’E, Aug 1964, van Royen NGF20068 (BRI). MADANG PROVINCE: Tukai, 50 miles [83.3 km] N of Madang, Mar 1968, Coode & Katik NGF32749 (BRI). WESTERN PROVINCE: Daru Island, Mar 1936, Brass 6272 (BO,BRI ex A). GULF PROVINCE: Keuru, Mar 1926, Brass 1191 (BRI). MILNE BAY Province: Near Tapio Village, Cape Vogel Peninsula, Jul 1954, Heogland 4332 (BRI,CANB). Solomon Islands. Guadalcanal, Mbokokimbo River, logging road, c. 2 km SE of main road, 9°28’S, 160°22’E, Jun 1991, Forster 8652 & Lidd/e (BISH,BRI,K,L,MEL); Santa Cruz Islands, Tomotu Noi Island, 10°47’S, 166°04’E, Apr 1972, Powell et al. BSIP19891 (BSIP,CANB). Australia. Northern Territory. Wessel Islands, 11°19’S, 136°36’E, Oct 1972, Latz 3500 (BRI,DNA). Queensland. CooKx District: Mutee Head, Cape York, 10°55’S, 142°1S’E, Mar 1990, Forster 6430 (BRI,CANB,DNA,K,L,MEL,PERTH,QRS); Kennedy Hill Gorge, 12°28’S, 143°16’E, Jun 1989, Forster 5404 & Kenning (BRI,L,MEL,QRS); Claudie River (tidal reaches), 12°50’S, 143°20’E, Jul 1972, Hyland 6207 (BRI,QRS); Ella Bay, near Innisfail, Jul 1943, Blake mane (BRI,CANB). WipE BAY DISTRICT: Freshwater Creek, S of Double Island Point, Aug 1964, Everist 7639 RI).
Distribution and habitat: Widespread in Malesia and Melanesia. Common in New Guinea and Solomon Islands; in Australia relatively common in north Queensland with one locality in Northern Territory (Map 2). Plants at the Freshwater Creek locality in south- east Queensland have been collected numerous times since the 1920s and comprise 2 or 3 trees which must have resulted from a chance establishment long ago. This colony does not seem to have expanded over a period of 70 years and is not considered native to south-east Queensland.
Plants are generally found in the littoral zone behind the mangroves, often in monsoon vinethicket along the shore or may extend inland in lowland rainforest communities.
SET REARS
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576 | Austrobaileya 3(4): 1992
Notes: Smith (1988) discussed the typification of this species’ name and supported Fosberg’s (in Boiteau 1981) lectotypification directly from the Osbeck description based on a Javan specimen, that was cited by Linnaeus.
Boiteau (1981) recognised several varieties and forms under C. manghas; however, apart from C. manghas f. manghas, most of these are probably referable to C. odollam Gaertner (Smith 1988).
In a red type folder at BRI is a specimen dating from Bailey’s time that is probably the type of C. odollam var. mugfordii. Despite the lack of label data that agrees with the protologue, there are no other specimens at BRI that could be the Mugford collection and it seems reasonable to assume that this is the type.
Local names and ethnobotanical use: A listing of local names, the dialects of origin, and the relevant vouchers are given in Table 2. In Solomon Islands, the crushed leaves are used on boils or polio with accompanying prayers, and the wood for carving paddles and bowls (Powell BSIP19891 (BSIP), Henderson & Oimae 175 (BSIP)). In Torres Strait, the fruit is used by children as a ball (Lawrie [AQ004029] (BRI)).
Conservation status: Not rare or endangered. Conserved in Cape Wom International Park, East Sepik in Papua New Guinea, but not known with certainty to occur in any Conservation Reserves in Australia.
Table 2. Local names for Cerbera manghas.
Name Dialect & Region Voucher
Aikikira Kwara’ae; S.I. Kere BSIP5S094 (BSIP) Aimalua Kwara’ae; S.1. Boraule et al. BSIP9318 (BSIP) Aitu Laukana; Morobe White NGF11155 (BRI)
Babai Biak; Irian Jaya Moll BW9827 (CANB)
Bulo Tasia®; S.L. Brass 3284 (BRI)
Dalovi Manukiki; S.I. Griffith 4/24 (BRI)
Gingum Oomsis*; Morobe White NGF11170 (BRI)
Grey Milkwood Australian; Cook Cause et al. (1989)
Kaiohto Gabobora; Milne Bay Hoogiand 4332 (BRD
La Toto Rapurt; New Britain Floyd NGF6451 (BRI) Ngambako Nanegu; S.I. Powell et al, BSIP19891 (BSIP) Papoea Aniaeri; Irian Jaya Aet & Idjan 614 (BO) Rubbertree. Australian; Cook Cause ef al. (1989)
Sas Biak; Inan Jaya van Royen 3545(BRD)
Soto Manukiki; S.J. Griffith 4/24 (BRI)
To Santa Ana™; S.I. Yen BSIP18113 (BSIP)
To’o Are’are; S.I. de Coppet 146 (BSIP)
Toto Kulumo; New Britain Barker & Vinas LAE66541 (BRI) Totongwala Kwara’ae; S.I. Gafui ef al. BSIP17448 (BSIP) Totora Karngu*: Bougainville Kajewski 2239 (BRI)
Vao New Georgia; S.I. Waterhouse 303 (BRI)
Wal Dauan Is; Cook Lawrie [AQ004029] (BRI)
* indicates locality name only, dialect not indicated. Abbreviations: S.J. = Solomon Islands.
Forster, Cerbera S77
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Fig. 1. A-D. lower leaf surfaces showing venation, X 5. A. Cerbera inflata. B. C. manghas. C. C. dumicola. D. C. floribunda. E-G. C. manghas: E. inflorescence X 0.5. F. face view of flower x 1. G. side view of flower X 1. H-l. C. dismicola: H. side view’of flower x 1. I. face view of flower < 1. J-L. C. manghas: J. longitudinal section of flower showing disposition of anthers in tube X |. K. intact fruit xX 0.5. L. half fruit showing interior x 0.5. A, Smith 3734; B,E~G,J-L, Forster 6430; C,H-I, Batianoff 11705b, D, Blake 14992. Del. W. Smith.
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$78 Austrobaileya 3(4): 1992
4. Cerbera dumicola P. Forster sp. nov. ad Cer beram manghas L. affinis, a qua habitu fruticoso, venis lateralibus laminae folii paucioribus (14-18), tubo corollae 10- 11 mm longo, et lobis corollae 8-9 mm longis, differt. Typus: Queensland. PORT CuRTIS Districr: Howard Point, Middle Percy Island, 55 km NE of Arthur Point, Shoalwater Bay, 21°40’S, 150°16’E, 31 October 1989, G.N. Batianoff 11705B, I, Champion, P. Thompson & H.A. Dillewaard (holo: BRI; 1s0: MEL!, QRS!).
Shrub or small tree to 4 m high, latex white; foliage and inflorescence glabrous. Bark light grey, fissured longitudinally, somewhat scaly on taller plants; sap wood white, heart wood white. Leaf lamina elliptic-lanceolate to elliptic-oblong, 5-17 cm long, 1.5-—6.0 cm wide, discolorous, margins often variously lobed and sinuate; upper surface glossy light green, venation obscure; lower surface pale green, secondary veins 14-18 per side of midrib, tertiary reticulate venation prominent; tip obtuse, acute or short acuminate; base cuneate; petiole 5-12 mm long, 0.7-0.8 mm wide. Inflorescence a little branched cyme up to 8 cm long; peduncle up to 5 cm long; flowers few, with generally less than 30. Flowers 15-18 mm long, 14-20 mm diameter, sweetly scented; pedicels 31-45 mm long. Sepals lanceolate-ovate, 8-9 mm long, 3.5-5.0 mm wide. Corolla white; tube 10-11 mm long, 1.8-2.2 mm diameter, slightly constricted above anthers, with dense indumentum internally; lobes ovate to obovate, 8-9 mm long, 4.5-5.0 mm wide. Stamens inserted c. 2mm from top of tube; anthers 1.0-1.2 mm long, 0.6-0.7 mm wide. Fruit globose- ovoid, c. 5.5 cm long, 4 cm wide, 4 cm thick, colour unknown. Fig. 1C, H, I.
Specimens examined. Australia, Queensland. NORTH KENNEDY DISTRICT: Barrabas Scrub, 20°10’S, 146°45’E, May 1972, Hyland 6097 (QRS). Port Curtis District: West Bay, Middle Percy Island, 55 km NE of Arthur Point, Shoalwater Bay, 21°40’S, 150°16’E, Nov 1989, Batianoff 11627 et al. (BRI. LEICHHARDT DISTRICT: [400- 500 miles inland from Rockham pton], 1886, Govt. Surveyor [MEL 1515485] (MEL); range between the Dawson & Mackenzie, [MEL 1515808] (MEL); c. 80 km NE of Emerald, 10 km SSE of “Booroondara”’, 22°54’S, 148°31’E, Oct 1978, Jones 6 (CBG); Blackwater, Jun 1942, Flowers [AQ212427] (BRI); 5 km NW of Wowan, Banana Shire, 23°50'S, 150°10'E, Feb 1990, Gillespie 4837 (BRI); 23°55’S, 148°48’E, Dec 1985, Thompson [AQ399095] (BRI): Portion 130, Gogango Shire, Parish of Fleetwood, Dec 1983. Heppell [A 398635] (BRI); Below Blackdown Tableland, Nov 1975, Williams [AQ113486] (BRD); Duaringa, Oct 1943, Blake 15360 elt ditto, Oct 1943, Murray [AQ212426] (BRI): ditto, Nov 1943, White 12460 (BRI); Baralaba, Feb 1943, Maclean [AQ212430) (BRI); ‘Coolum’, Baralaba on eastern slope of Dawson Range, Nov 1943, Maclean [AQ212429 ] (BRD; near Stanwell, Apr 1876, O’Shanesy 1829 (MEL). MITCHELL DISTRICT: Romulus Tableland, 57 km SE of Blackall on the top of Enniskillen Range, 24°44’S, 145°52’E, Jul 1975, Beeston 1430c (BRI); Ravensbourne-Mount Edinburgh area, Blackall/Tambo Shire, Jul 1990, Roche [AQ472683] (BRI).
Distribution and habitat: Central coastal and subcoastal Queensland (Map 3). Plants occur primarily in lancewood (Acacia shirleyi Maiden) thickets away from the coast but may also be found in semi-evergreen vine thickets near the coast.
Notes: C. dumicola was discovered in 1876 by R. O’Shanesy who sent material (a sterile leafy stem) to von Mueller in MEL. Although collected sporadically since (collections in MEL), its distinctiveness was only recognised by the late S.T. Blake who made or organised a number of collections of the plant in the 1940s. Despite having fertile material (scrappy flowers only), Blake did not describe the plant and sporadic and nearly always sterile collections were made up until 1990, when good flowering material procured by G. Batianoff and associates allowed for its description. The fruit description is based on a single old sectioned fruit [MEL 1515485] (MEL) and further collections are still required for assessment of variation in the taxon.
C. dumicola is a distinctive species by virtue of the shrubby to small tree habit and the foliage often having sinuate margins. It is most closely allied to C. manghas but differs from that species in the shrubby habit, fewer lateral veins in the leaf lamina (14-18 per side of midrib); corolla lacking a red centre, the corolla tube 10-11 mm long and the corolla lobes 8-9 mm long.
ony The specific epithet alludes to the common occurrence of this species in thickets.
Conservation status: Not known to occur in any conservation reserves. The various populations in central Queensland must be considered under threat from agriculture and mining activities. Urgent survey work is required to determine the status of this taxon. A relevant conservation coding 1s 3RC (cf. Briggs & Leigh 1988).
Forster, Cerbera $79
Acknowledgements
Assistance with field work in north Queensland and Solomon Islands was given by G. Kenning, D. Liddle and M.C. Tucker. G. Batianoff (BRI) collected fertile material of C. dumicola. The illustrations were prepared by W. Smith (BRI) with funding from the Australian Biological Resources Study (ABRS). P.R. Sharpe translated various German texts. A.S. George (ABRS) translated the diagnosis into latin. The herbaria AD, BO, BRI, BSIP, CANB, CBG, DNA, K, L, MEL and QRS allowed access to collections either at their institutions or on loan. Loans staff at these institutions and BRI expedited rapid processing. G. Leach (DNA) while Australian Botanical Liaison Officer at Kew, U.K., located and photographed type material. The author was funded for this project by ABRS during 1991-92. This assistance is gratefully acknowledged.
References BAILEY, F.M. (1900). Apocynaceae. Queensiand Flora 3: 974-995. Brisbane: Government Printer. BENTHAM, G. (1869). Apocynaceae. Flora Australiensis 4: 301-324. London: L. Reeve.
BLAKE, S.T. (1948). Studies in Australian Apocynaceae and Asclepiadaceae, |. Proceedings of the Royal Society of Queensland 59: 161-168.
BLAKE, S.T. (1959). New or noteworthy plants, chiefly from Queensland, |. Proceedings of the Royal Society of Queensland 70: 33-46.
BOITEAY, P. (1981). Flore de la Nouvelle-Calédonie et Dependances, 10. Apocynacées. Paris: Muséum National d’Histoire Naturelle.
BRIGGS, J.D. & LEIGH, J.H. (1988). Rare or Threatened Australian Plants. 1988 Revised Edition. Australian pahone Parks and Wildlife Service Special Publication No. 14. Canberra: Australian National Parks and Wildlife Service.
CAUSE, M.L., RUDDER, EJ. & KYNASTON, W.T. (1989). Queensland Timbers, their nomenclature, density and lyctid susceptibility. Technical Pamphlet No. 2. Brisbane: Queensland Department of Forestry.
FOREMAN, D.B. (1971). A Check List of the Vascular Plants of Bougainville, with descriptions of some common forest trees. Botany Bulletin No. 5. Lae: Department of Forests, Division of Botany.
FOSBERG, F.R., BOTTEAU, P. & SACHET, M.-H. (1977). Nomenclature of the Ochrosiinae (Apocynaceae): 2. Synonymy of Ocirosia Juss. and Netsosperma Raf. Adansonia, ser. 2, 17; 23-33.
HOLMGREN, P.K., HOLMGREN, N.H. & BARNETT, L.C. (eds) (1990). Index Herbariorum. Part I: The Herbaria of the World. New York: New York Botanical Garden.
LINNAEUS, C. (1753). Species Plantarum. Stockholm: Laurenti, Salvii.
LIPPOLD, H. (1980). Die gattungen Thevetia L., Cerbera L. und Cascabela Rafin. (Apocynaceae). Feddes Repertorium 91: 45-55.
MARKGRAF, F. (1926). Apocynaceae. Nova Guinea 14(2): 278-291.
MARKGRAF, F. (1927). Die Apocynaceen von Neu-Guinea. Botanische Jahrbticher ftir Systematik, Pllanzenges- chichte und Pflanzengeographie 61: 164-222.
MARKGRAF, F. (1972). Florae Malesianae Praecursores LIII. Apocynaceae IH. 6. Urnularia, 7. Willughbeia, 8. Kopsia. Blumea 20: 407-425.
SMITH, A.C. (1988). Flora Vitiensis Nova. Vol. 4. Lawai: Pacific Tropical Botanical Garden.
THOMAS, M.B. & MCDONALD, W.J.F. (1989). Rare and Threatened Plants of Queensland. 2nd Edition. - Brisbane: Queensland Government.
VAN ROYEN, P., WOMERSLEY, J.S., WHITE, K.J., COLWELL, S.J. & PEARCE, D. (1964). Manual of the Forest Trees of Papua and New Guinea. Part 9. Apocynaceae. Port Moresby: Department of Forests.
WHITE, C.T. (1933). Ligneous plants collected for the Arnold Arboretum in north Queensland by S.F. Kajewski in 1929. Contributions from the Arnold Arboretum of Harvard University 4. 1-113.
Accepted for publication 7 February 1992
Austrobaileya 3(4): 581-583 (1992) 581
A NEW SPECIES OF LIPARIS RICH. (ORCHIDACEAE) FROM NORTH QUEENSLAND
B. Gray
Australian National Herbarium, CSIRO, Division of Plant Industry, PO Box 780 Atherton, Qid 4883, Australia
Summary
Liparis collinsit B. Gray, a new species of Liparis Rich. section Liparis, is described and illustrated. In addition notes on its taxonomic affinities, habitat, distribution and conservation status are given.
Taxonomy
Liparis collinsii B. Gray, sp. nov. Differt a L. simmondsio Bailey pseudobulbis brevioribus validioribus et foliis angustioribus lanceolatis epetiolatis basi vaginantibus. Typus: Cult. Atherton ex Queensland, Timber Reserve 14, Parish of Kesteven, 13°46’S, 143°21/E. 740 m, 13 September 1990, B. Gray 5269 (holo: ORS: iso: BRI, CANB, NSW).
Erect terrestrial herb, deciduous, glabrous, 25-50 cm tall. Pseudobulb fleshy, + elliptic in cross section, broadest near the base, tapering to the apex, 6-10 cm long, 2-3 cm diameter; bracts 2- 4, ovate, ee strongly ribbed, apex acute; new shoot arising from the base of the old. Leaves (3-4)-6; lamina lanceolate, 10-13 cm long, 4-5 cm wide, somewhat plicate, with 4 or § reddish purple ribs below, margin undulate, apex acute. Petiole broad, sheathing, 5-7 cm long. Inflorescence terminal, erect, a {0-20- flowered raceme, 25-35 cm long; peduncle 4-angled, slightly winged, about as long as the rhachis; bracts 1 or 2, linear-triangular, 6-9 mm long, 2.0-2.5 mm wide; floral bracts linear, 2-6 mm long, c. i mm wide, acute; pedicel with ovary 10-16 mm long. Flowers spreading widely, 2.0-2.5 cm diameter. Dorsal sepal linear to narrowly ovate, 12-14 mm long, 3.0-3.5 mm wide, pale purplish green, margin revolute. Lateral sepals elliptic, somewhat falcate, 9-10 mm long, 3.0-3.5 mm wide, pale purplish cream, revolute. Petals filiform, 11-14 mm long, 1.4-1.5 mm wide, purple, strongly revolute (rolled into a very narrow tube). Labellum obovate, c. 13 mm long, 8-10 mm wide, greenish to greenish cream at the base, suffused with maroon towards the apex, ventral surface with a narrow maroon central line from the base to apex; base cuneate; apex truncate to + emarginate; margin denticulate near the apex; semierect in the basal third, and channelled then decurved through 90° and spreading, with 2 calli on the ventral surface; calli c. 2 mm long, 0.5 mm diameter. Column arcuate, c. 7 mm long, 2 mm diameter, sreenish cream. ries: es cap 1.3-1.4 X 1.3 mm. Pollinia 4, in 2 pairs.
Distribution and habitat: This species is at present known only from the Mcllwraith Range on Cape York Peninsula in Queensland. It grows among granite boulders in relatively open areas in rain forest above 700 m altitude. Plants occur in loose colonies and are leafless throughout the dry season. (June to November or December).
Flowering period: Time of flowering in nature is not known but probably occurs soon after the onset of the wet season. In cultivation flowering occurred from November to January, while the plants were in full growth. Individual flowers last 7-10 days.
Notes: L. collinsii is most closely related to L. simmondsii Bailey but can be readily distinguished from that by its shorter, stouter pseudobulbs and the narrower, lanceolate non-petiolate leaves which are sheathing at the base. The two species have distinct geographic ranges. L. simmondsii has not been recorded north of the Mossman River, which is 350 km south of the MclIlwraith Range.
The widespread L. habenarina (F. Muell.) Benth. which occurs in open forest and grassland situations of north and eastern Australia differs from ZL. collinsii in having narrower leaves and pseudobulbs which are produced below ground level.
Of the several species eo New Guinea, L. oligantha Schltr. and L. finnisterrae Schltr. appear to be related to L. collinsii but are quite distinct.
582 Austrobaileya 3(4): 1992
Fig. 1. Liparis collinsii: A. habit. B. flower from side. C. flower from front. D. column from side. E. column from front. F. section through column and labellum. G. anther from front. H. anther from side. I. pollinia. J. labellum flattened. K. section through base of labellum. L. dorsal sepal flattened. M. lateral sepal flattened. N. sepal flattened. All from Gray 5269.
Gray, Liparis collinsti 583 Conservation status: This species is poorly known and its distribution, even within the Mcliwraith Range area is uncertain.
Etymology: It gives me great pleasure to name this orchid after the late Rev. R.D. (Ron) Collins of Atherton. Ron had a great interest in orchids, was a good friend and accompanied me on many field trips to Cape York Peninsula.
References
DOCKRILL, A.W. (1969). Australian Indigenous Orchids. Volume 1. Sydney: The Society for Growing Australian
Plants.
SCHLECTER, R. (1982). The Orchids of German New Guinea. Translated by R.S. Rogers, H.J. Katz & J.T. Simmons, edited by D.F. Blaxell. Melbourne: Australian Orchid Foundation.
Accepted for publication 18 March 1992
Syste oven yc eanone nt mm tee
Austrobaileya 3(4): 585-607 (1992) 585
STUDIES IN AUSTRALIAN GRASSES: 5* NEW SPECIES OF AND NEW COMBINATIONS FOR
QUEENSLAND PANICOID GRASSES
Bryan K. Simon
Queensland Herbarium, Meiers Road, Indooroopilly, Qld 4068, Australia
Summary
New species, Arthragrostis clarksoniana, Cyrtococcum capitis-york, Panicum bombycinum, Panicum chillagoanum, Panicum robustum, Paspalum batianoffii, Paspalum multinodum and Yakirra websteri, are described. New combinations made are Brachiaria atrisola, based on Urechloa atrisola and Brachiaria holosericea subsp. velutina, based on Urochloa holosericea subsp. velutina. An overview of the current situation regarding the taxonomy of Brachiaria and allied genera is given. The occurrence of Brachiaria distachya (L.) Stapf in Australia, New Caledonia and Fiji is established.
Introduction
There are a number of new Queensiand grasses, first drawn attention to in ‘A Key to Australian Grasses’ (Simon 1990), that require formal description. Of these, the panicoid species are treated in this paper, with nine new species described and two new combinations made in the genus Brachiaria for taxa already described under Urochloa by Webster (1987). For each new species there is a Latin diagnosis, a full botanical description, a listing of specimens examined, a section on etymology, a section on conservation status, following the system used by Briggs and Leigh (1988), and some additional notes.
Arthragrostis Lazarides
A panicoid grass with distinctive tubercle-based hairs on the upper glume and lower lemma was collected from York Downs, near Weipa on Cape York Peninsula, in 1981 by Anne Morton (now Gunness). More recently, two more collections of this grass have been made, one from Meripah in 1987 and the other from Batavia Downs in 1990. In my Australian grass key (Simon 1990) I indicated this species by Panicum sp. Morton 191, although the collector’s number should have been cited as 1191. The distinctive character of the absolute disarticulation of the panicle at the base of all the branches as well se beneath the spikelets, necessitates the species be placed in the genus Arthragrostis Lazarides.
Arthragrostis clarksoniana B. Simon, sp. noy. A. deschampsioidi (Domin) Lazarides affinis, sed gluma supera et gluma inferno in longitudine aequali plus minusve et pilis tuberculatis differt. Typus: Queensland. Cook District: 16 km from 7h pas homestead on road to the south, 13°49’S 142°22’E, 11 May 1987, JR. Clarkson 7149 & B.K. Simon (holo: BRI(AQ 452649); iso: MBA,NSW). Fig. 1.
Weak annual. Culms erect or basally decumbent, weakly tufted, 40-60 cm tall, sparingly branched at base, 2-4-noded, terminated by a solitary, large and open inflorescence about half the length of the culm. Internodes longer than the associated leaf sheaths which are rounded on the back. Ligule a fringed membrane c. 0.5 mm long. Leaf blades lanceolate, flat, 3-10 cm X 20 mm, pilose, margins smooth with scattered tubercle-based hairs. Inflorescence a panicle with main axis 23-30 cm long and smooth. Primary branches spreading, not whoried, 10-21 cm long, scabrous on the margins. Pedicels 2- 12 mm long, distinctly angled, straight, scabrous although with glabrous apices. Callus not differentiated. Spikelets abaxial, 15-25 on a typical lowermost primary branch, dorsiventrally compressed, ovate or elliptic 1n outline, 3.5-4.0 X c. 1.5 mm. Glumes + equal, 3.5~4.0 mm long: lower glume, ovate or elliptic in outline, acuminate, 5-7-nerved, membranous, hairy with tubercle-based cilia scattered over the back on the lower two-
*continued from Austrobaileya 3(1): 79-99 (1989)
586 Austrobaileya 3(4): 1992
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Fig. 1. Arthragrostis clarksoniana: A. plant X 0.5. B. base of primary branch, showing disarticulation. C. spikelet D. lower glume, back view. E. upper glume, back view. F. lower floret with minute palea within lemma. G. upper floret, back view. H. upper floret, front view. (B-H x 8). L caryopsis, view of side with embryo. J. caryopsis, view of side with hilum. (I,J < 16). From type specimen.
Simon, Australian Grasses 5 587
thirds; upper glume narrowly elliptic in outline, 7-9-nerved, membranous, hairy with tubercle-based cilia scattered over the back on the lower two-thirds. Rachilla conspicuous between the glumes. Lower lemma elliptic in outline, acute, membranous, glabrous, 3.0- 3.5 mm long. Palea of lower floret vestigial, apically truncate. Upper floret ‘perfect, elliptic in outline, shorter than the lower floret. Upper lemma c. 1.5 mm long, yellow, chartaceous, smooth and shiny, rounded on the back, glabrous. Palea of upper floret chartaceous, smooth. Caryopsis c. 1.5 mm long with hilum about half as long as caryopsis.
Specimens examined: Queensland. Cook District: York Downs on Myall Creek, 12°45’S, 142°18E, mg Pl 4QE,
tetrodonta — E. confertiflora grassy open forest, May 1981, Aforton 1191 (BRI); Batavia Downs, 12°40’ S. 142 fenced areas around homestead and yards, Apr 1990, Clarkson 8590 & Neldner (BRI,MBA).
Conservation status: 3K (Briggs & Leigh 1988).
Etymology: The species is named for John Clarkson, Queensland Herbarium Senior Botanist at Mareeba. He has enthusiastically collected much interesting botanical material from north Queensland and who has provided me hospitality, facilities and access to remote areas of Cape York on a number of occasions.
Notes: Although the presence of a lower palea for this genus was originally recorded by Domin (1915, under Panicum deschampsioides), it has not been mentioned in more recent papers (Lazarides 1984; Simon 1986). However it 1s present as a very small vestigial membrane in specimens of both Arthragrostis clarksoniana and A. deschamp- sioides. The glumes in A. clarksoniana differ from those in the other species in being of more or less equal length, although previously reported as having the lower glume longer than the upper (Simon 1990). In some spikelets the lower glume may be slightly longer than the upper glume, but in others the lower glume may be slightly shorter than the upper.
Brachiaria (Trin.) Griseb.
A state of instability exists regarding the taxonomic status of the genera Urochloa P. Beauv. and Brachiaria (Trin.) Griseb. The classical difference between these genera is whether the spikelet orientation is adaxial (with the lower glume adjacent to the inflorescence axis), the general situation in Brachiaria and some panicoid genera, or abaxial (with the lower glume remote from the inflorescence axis) in Urochioa and most other panicoid genera. To a lesser extent, the possession of a racemose inflorescence distinguishes Brachiaria from other panicoid genera. Both these characters cannot be universally applied, however, and the position regarding the taxonomic value of these characters is summarised in the relevant literature (Chase 1911, 1920; Hughes 1923; Gardner & Hubbard 1938; Blake 1958; Hsu 1965; Parodi 1969; Sendulsky 1978; Zuloaga & Soderstrom 1985; Clayton & Renvoize 1986).
Blake (1958) presented a good summary of the taxonomic history of this group of genera.
“Bentham (1878, 1883) treated under Panicum several groups that had been proposed as distinct genera by earlier authors, including all Brachiaria and Urochloa. Later Chase (1906-11) investigated the diagnostic value of the texture of the upper lemma and the nature of its margins as well as the orientation of the spikelets; she showed that these characters were so correlated with others that some of the groups previously proposed as genera could be sharply redefined as such. Most botanists soon accepted Chase’s principles for generic discrimination and new genera were described by Stapf (1917-1930) and others. Some of these genera (Paspalidium, Entolasia, and Ottochloa among others) were not accepted by Chase (1939, 1951 etc.), Hitchcock (1936a, 1936b etc.) or Reeder (1948), all of whom referred the species they dealt with to Panicum. These authors and Pilger (1940) treated under Panicum (Panicum sect. Fasciculata Hitch. & Chase or Panicum subgen. Urochloides Pilger) a group of species referred partly to Brachiaria and partly to Urochloa by Stapf (1917-1930) and Hughes (1923), and to Brachiaria by Gardner and Hubbard (1938).”
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588 Austrobaileya 3(4): 1992
In a manuscript of the Paniceae for A. Engler’s Pflanzenreich, recently distributed from Berlin (B), Mez placed Brachiaria, Urochloa and Eriochloa in Panicum subgenus Brachiaria (Griseb.) Benth. & Hook.
Brachiaria was originally described as a section of Panicum by Trinius (1826) and elevated to generic rank by Grisebach (1853), but neither author indicated the characters by which the taxon could be distinguished. In Trinius’s section were four species with reversed (adaxial) spikelets among others with normal (abaxial) spikelets. As Grisebach only mentions Brachiaria ucts in his treatment, the type of this name is the nomenclatural type of the generic name. He specifically describes the racemose primary branches but not the adaxial spikelets.
Further taxonomic history of the genus is documented by Webster (1987).
“In 1903 Nash, working on the grasses of the southeastern United States, recognised the adaxial orientation of the spikelets and transferred some Panicum species to Brachiaria. Stapf (1919) in ‘Flora of Tropical Africa’ used the presence of racemose primary branches as the distinguishing feature of Brachiaria and transferred numerous species from Panicum. The American authors, Chase and Hitchcock made additional combinations in Brachiaria, whereas [Gardner and| Hubbard (1938) and Hughes (1923) named many of the Australian species in Brachiaria. These various authors used the racemose primary branches and spikelet orientation to distinguish Brachiaria from Panicum. Chase (1920) states that Brachiaria spikelets are adaxial whereas Panicum is abaxial and this character appears in the various keys produced by Hitchcock and Chase.”
Blake (1969) transferred four American species of Panicum sect. Fasciculata to Brachiaria; two of these, however, had also been transferred to Brachiaria by Parodi (1969) and the combination published only days before Blake’s account (Blake 1973). Butzin (1970), in a paper proposing a new subtribal classification of the tribe Paniceae, placed Brachiaria in the subtribe Brachiariinae, together with nine other genera, on the basis of the abaxial lower glume, whereas Urochloa was placed in the subtribe Paspalinae with 13 other genera on the basis of the adaxial lower glume. Shaw and Siemens (1980) point out the strong affinity between Brachiaria, Eriochioa and Urochioa on the basis of their all possessing the PEP-ck subtype of C, photosynthesis (Gutierrez, Gracen & Edwards 1974), and that Urochloa differs from the other two by the abaxial orientation of the lower glume and a basic chromosome number of x = 10 as opposed to x = 9 in Eriochloa and Brachiaria. However they also add the rider that the latter two characters may not be as significant in the classification of the Paniceae as previously thought.
There appears some dispute whether the lower glume in Brachiaria is universally adaxial. Establishing whether the lower glume is adaxial or abaxial in species with long pedicels is difficult, but Gardner and Hubbard (1938) explained a method to overcome this which was summarised by Zuloaga and Soderstrom (1985).
“They suggested that the spikelet just below the terminal one on the axis or on a branch of the panicle be examined to determine the position of the lower glume, because the spikelet 1s generally short-pedicelled and appressed to the rachis. They commented that the orientation is still occasionally unclear due to suppres- sion or rudimentary development of the lower spikelet of the pair, in which case the terminal spikelet appears to be solitary and in an abaxial position.”
Webster (1987), however, implied that this anomaly of spikelet orientation in some of the spikelets of Brachiaria reptans is sufficient for the character to lose its significance. It is a reason, considered with other characters, for his placing all Australian species of Brachiaria, excluding B. eruciformis, in Urochloa. However Clayton and Renvoize (1986), while accepting the taxonomic proximity of Brachiaria, Urochloa, Eriochloa and Panicum, did not emphasise spikelet reversal in their comparisons but mentioned arbitrary characters such as ‘spikelet plumpness’ in Brachiaria and ‘plano- convex shape, cuspidate tip and mucronate upper lemma’ in Urochloa. However, under Urochloa, they stated that “when the spikelets are paired their orientation becomes ambiguous and diagnosis then rests upon their facies. Unfortunately orientation and facies are not wholly correlated, some intermediates being noted under Brachiaria, and it is a moot point whether generic rank is justified.”
Simon, Australian Grasses 5 589
However, the recent transfer of most species of Brachiaria and Panicum maximum to Urochloa by Webster (1987, 1989) and Webster ef a/. (1988, 1989) on the basis of morphological (the possession of rugose rather than smooth fertile lemmas, the spikelet disarticulating at the spikelet base as opposed to above the glumes, and the apex of the upper floret being mucronate to awned rather without mucros or awns) and anatomical (leaf anatomy associated with the PEP-ck biochemical variant of C, photosynthesis) characters, is probably premature as Ellis (1988) pointed out “the correlation between PEP-ck type anatomy with centrifugal chloroplasts and panicoid grasses with rugose lemmas 1s not perfect and that some species with smooth shiny lemmas also belong to this complex.” Furthermore the placing of Eriochloa, on the basis of its similar leaf anatomy and the lemmas being finely pitted or ridged (Chippindall 1955), in the same eroup as Brachiaria, Urochloa and Panicum maximum is not followed by Webster. The transfer of most Australian species of Brachiaria to Urochloa has been accepted by some botanists (Kenneally 1989; Wheeler, Jacobs & Norton 1990; Hnatiuk 1990; Wheeler in press) and the transfer of Panicum maximum to Urochioa has been accepted by Hnatiuk (1990) and Wheeler (in press) but not by Wheeler, Jacobs and Norton (1990). Watson and Dallwitz (1988), although appearing to agree with the transfer, do not actually accept it. Under Brachiaria they made the comment “all but the type species arguably best referred to Urochloa,” and under Urochloa “unsatisfactorily delimited from other close allies of Panicum, in particular Brachiaria.’ However, in terms of their ‘nearest neigh- bours’ analysis, Urochioa is listed as only the fourth closest relative to Brachiaria under the treatment of Brachiaria, although Brachiaria is listed as the closest to Urochloa under the treatment of Urochloa. Thompson and co-workers (Thompson & Estes 1986: Thompson 1988; Thompson, Tyrl & Estes 1990) have refrained from making generic transfers but indicate that within the Brachiaria group of Brown (Gutierrez, Edwards & Brown 1976; Brown 1977), Brachiaria (ncluding Panicum sect. Fasciculata), Eriochloa and Urochloa appear to constitute a monophyletic group. However they did not mention Panicum maximum as a member of the Brachiaria group, although included there by Brown and regarded by others as a candidate for generic transfer to the Brachiaria group on the basis of its rugose lemmas and PEP-ck anatomy (Ellis 1977; Hattersley 1987; Zuloaga 1987). Until a complete cladistic analysis is undertaken on all member species of the group it 1s probably better to leave the classification as it is.
Prior to Webster’s work a few species of Brachiaria had been placed in or transferred to Urochloa. Names provided for them in Urochloa are U. reptans (L.) Stapf (Stapf 1920), U. gilesii (Benth.) Hughes and U. praetervisa (Domin) Hughes (Hughes 1923), and U. mutica (Forssk.) Nguyen, U. kurzii (J.D. Hook.) Nguyen and U. ramosa (L.) Nguyen (Nguyen 1966). The latter two combinations were superfluously made again by Webster. Thompson (1988) retained in Brachiaria the Australian species transferred by Webster to Urochloa except Urochiloa gilesit and U. reptans, but I am retaining both se ce in Brachiaria as the lower glume in both of these species is for the most part adaxial.
Iam recognising Brachiaria in the classical sense pending results of cladistic work on the whole complex. Names are thus required in Brachiaria for taxa described by Webster for the first time in Urochloa.
Brachiaria atrisola (R. Webster) B. Simon, comb. nov.
Urochloa atrisola R. Webster, The Australian Paniceae (Poaceae) 232 (1987). Typus: Northern Territory. Barkly Tableland, 17°20’S, 135°45’E, undulating Astrebla grassland, grey pedocalcic soil, 14 May 1947, S.7. Blake 17776 (holo: BRI(BRI 186629)(AQ 256724); iso: BRICBRI 065747), CANB,DNA,K,MO),. Fig. 2.
Additional specimens examined: Northern Territory. BARKLY TABLELANDS: SW of Brunette Downs, dry bed of Lake Sylvester, May 1947 Blake 17830 (BRI,DNA). Queensland. BURKE District: 30 miles [48 km] S of Julia Creek, Mitchell grass downs, Mar 1959, Sillar 8 (BRI); Galway Downs, Hughenden, Mar 1933, Kirby s.n. (BRI,K,L,NSW).
Conservation status: Although the species does not appear threatened, I am aware of only four collections of it and therefore 1t should be placed in the category 3K (Briggs & Leigh 1988).
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590 Austrobaileya 3(4): 1992
a A at
att rattae 4 t + |
Fig. 2. Brachiaria atrisola: A. habit X 0.7. B. spikelet showing adaxial position of lower glume X 8. C. lower glume, back view. D. upper glume, front view. E. lower lemma, back view. F, upper floret, front view. G. upper floret, back view. H. caryopsis, view of side with hilum. I. caryopsis, view of side with embryo. J. lodicule. (C- J, X {2). K. anther X 25. From type specimen.
Simon, Australian Grasses 5 591
Etymology: Named for the black soils from which this species has been collected.
Notes: This species 1s represented only by four collections from the black-soil region of the Barkly Tableland of the Northern Territory and the adjacent parts of Queensland. According to Webster (1987) “it 1s morphologically similar to the glabrous form of Urochloa (Brachiaria) piligera but has a rounded apex on the second glume and lower lemma, turgid spikelets, relatively larger spikelets and lacks the distinct callus at the base of the spikelet.”
Brachiaria holosericea subsp. velutina (R. Webster) B. Simon, comb. nov.
Urochloa holosericea Webster subsp. velutina R. Webster, The Australian Paniceae (Poaceae) 240 (1987). Typus: Northern Territory. CENTRAL NORTH DISTRICT: Yeundumu Aboriginal Reserve, 22°10’S, 131°48’E, P.K. Latz 1979 (holo: CANB, n.v., fide Webster foc. cit.)
Specimens examined (All BRI; other herbaria cited where there are duplicates there): Northern Territory. DARWIN AND GULF: Near Balbirini Creek, May 1947, Blake 17740; Katherine, Feb 1947, Miles sn. CENTRAL NoRTH: Mt Doreen Station, Jan 1972, Latz 2046 (DNA). BARKLY TABLELAND: 35 miles [56 km] N of Newcastle Waters, Feb 1969, Afust 406 (DNA). Queensland. Cook District: 22 km W of Georgetown, Apr 1973, Henderson H1747; The Lynd, Feb 1968, Horsup 6; near Lynd Station, Feb 1954, Lazarides 4163. BURKE DISTRICT: Paroo Range N of Mt Isa, Jan 1990, Harris 434; Chudleigh Park, Feb 1931, Hubbard 7608 & Winders (K); ditto, Hubbard 7674 & Winders (K); Setthkement Creek, Mar 1923, Brass 312; Poison Creek N of Hughenden, Apr 1935, Blake 8609; Adel’s Grove, Camooweal, Mar 1947, de Lestang 302, 355; 14 miles [22.4 km] N of Mt Sturgeon, Jun 1953, Lazarides 3666 (CANB),; 88 miles [141 km] E of Camooweal, Feb 1937, Everist 189 & Smith; Normanton, May 1935, Blake 9044. NORTH KENNEDY DISTRICT: Warrigal Creek, Jun 1980, Rebgetz 422, near Burdekin and Bogie Rivers, Oct 1950, Blake 18690; Heathfield, Aug 1942, Smith T78; Milray, Oct 1935, Blake 9964: Pentland, Apr 1935, Blake 8375; Ayr Beach, Michael s.n. MITCHELL DISTRICT: Prairie, Feb 1931, Hubbard 7034 & Winders (K); Aramac-Torrens Creek Road, Jun 1977, AdcDonald 2622; Geera, Nov 1935, Blake 10330; E of Jericho, Feb 1931, Hubbard 7834 (K).
Etymology: Named for the velutinous leaves.
Notes: Webster (1987) stated that this subspecies “occurs in the arid and semiarid areas of north central Australia, whereas subspecies Holosericea occurs north of this area in tropical and subtropical subhumid woodlands. Morphologically, it differs from the typical subspecies in possessing velutinous leaves, fewer and shorter hairs on the inflorescence branches and pedicels, and generally longer awned lower lemmas.” Examination of the Queensland Herbarium material revealed that about half the material previously placed unsorted under Brachiaria holosericea actually belongs in B. holosericea subsp. velutina.
Brachiaria distachya (L.) Stapf
For some time confusion has existed concerning the correct application of the names Brachiaria distachya (L.) Stapf, B. miltiformis (Presl) Chase and B. subquadripara (Trin.) Hitche. Although these names have been regarded as applying to one species in the Malaya Peninsula (Gilliland 1971), the entity with smaller spikelets and a shorter inflorescence axis with fewer racemes, has been generally recognised as B. distachya (Jansen 1953; Bor 1960; Morat 1981; Clayton & Renvoize 1982). Regarding the other two names some authorities (Henrard 1950; Jansen 1953; Morat 1981) have synonymised B. miliiformis with B. subquadripara while others (Bor 1960; Blake 1948; Vickery 1961) have recognised both as applying to separate species. The case presented by Jansen (1953) for uniting these species is convincing to me in that a comparison of the drawing of a spikelet of B. subquadripara in Henrard (1950) with that of a spikelet on the type of Panicum miliiforme (the basionym of B. miliiformis) in Lamson-Scribner (1899) shows them to be almost identical. The drawing of a spikelet on the type of Panicum subquadriparum (the basionym of B. subquadripara) in Trinius (1829) also seems very similar to that of Panicum miltiforme in Lamson-Scribner (1899). As described and illustrated the types of both these names are annuals. Both types also show the presence ofa eee palea, whereas it appears to be absent in many other specimens of this species examined.
In tropical Australia and Asia there is a decumbent species, referred to in Simon (1990) as B. sp. Everist 5112, which has for some time been confused with B. subquadripara due to its very similar, but smaller spikelets. Because of its habit and the possibility that it could have some economic potential as a sward grass, I had been of the opinion it should be formerly recognised. All the specimens of this species have the lower palea
592 Austrobaileya 3(4): 1992
present and I was of the opinion this could be used as a supplementary morphological character to differentiate it from B. subquadripara till my recent discovery of the presence of a lower palea in some specimens of the latter, including the types of both Panicum miliiforme and P. subquadriparum. An examination of the microfiche of the type of Panicum distachyon 10 the Linnean herbarium (LINN) and specimens of Brachiaria distachya from India matched by C.E. Hubbard with the type (Gamble 17629 and Drummond 21156, both K with cibachrome photographs in BRI), has brought me to the conclusion that the decumbent sward-forming species referred to above, is in fact this species, although the inflorescence tends to be larger in some of the Australian material than the Indian specimens seen. Whereas the names Panicum distachyum
(Bentham 1878), or Brachiaria distachya (Hughes 1923) have been used in earlier times’
for an Australian grass species, they have been used for a broad concept of it that includes Brachiaria subguadripara. This 1s similar to what was done for the taxon on the Malaya Peninsula more recently (Gilliland 1971). C.E. Hubbard was of the opinion in 1933 that “true B. distachya does not occur in Australia” in a memorandum on Brachiaria subquadripara sent to the Queensland Herbarium and Webster (1987) did not include B. distachya in his treatment of the Australian Paniceae. However there are a fairly large number of specimens in BRI which can be referred to Brachiaria distachya. There are a few records of this species from outside Australia in BRI, including some from New Caledonia and Fiji, where it has previously been thought not to occur (Morat 1981; Toutain 1989; Parham 1979) but the specimens cited below belong to this species rather than B. subguadripara, where they were previously placed.
Specimens examined (Ali BRI; other herbaria cited where there are duplicates there): Sri Lanka. Chenkaladi, Dec 1974, Davidse 8997 & Sumithraarachchi (MO); Paranthan, Dec 1974, Davidse 9129 & Sumithraarachchi (MO). New Guinea. PoRT MORESBY PROVINCE: Nebiri Quarry, Apr 1970, Gebo UPNG 96. WESTERN PROVINCE: Mabaduan, Jun 1973, Henty NGF 49601. BOUGAINVILLE PROVINCE: Kieta, Oct 1969, Henty NGF 42711. Australia. Northern Territory. DARWIN AND GULF: Vanderlin Island, Aug 1988, Latz 11063 (CANB,DNA); Adelaide River, Aug 1946, Blake 16704. Queensland. Cook Disrrict: Thursday Island, Apr 1931, Hockings [AQ 255899]; Cairns, Jun 1930, Hifi [AQ 255903]: Low Isiand, Great Barrier Reef, Jun 1969, Done [AQ 8175]; Mareeba, Apr 1983, Clarkson 4593 (QRS,DNA,PERTH), Jun 1963, Veurman [AQ 255900]; Bamaga, May i981, Af/corn 8144, 8145; Yorkeys Knob, Apr 1962, McKee 9024, Gordonvale, Oct 1935, Blake 9853; Babinda, Mar 1973, Henderson H532; Cooktown, Apr 1973, Henderson H1602; South Johnstone, Apr 1938, Langdon 23; ditto, Jun 1963, Bailey 1; Daradgee, Jun 1936, Goodman s.n.; Kamarunga, May 1952, Everist 5112; Mornington Island, Sep 1981, Fosberg 62164; Freshwater, Mar 1938, Blake 13349; Bizant, Lakefield National Park, May 1987, Clarkson 6953 & Simon; Clifton Beach, Feb 1985, Lear s.n. [AQ 396685]; Corduroy Creek, Feb 1983, Steel 355. NoRTH KENNEDY DISTRICT: Dingo Beach, Apr 1978, Simon 3386; Trebonne, Hinchinbrook, May 1986, Stee/ [AQ 440235], Clark River Telegraph Station, Jul 1954, Blake 19436; Euramo, Mar 1961, Saint-Smith s.n.; Townsville, Jun 1975, Burmeister s.n.; Little Crystal Nationa! Park, May 1975, Simon 2621 & Andrews, Pettrides Bridge, May 1975, Simion 2644 & Andrews, Saunders Beach, Apr 1975, AfcDonald 1444 & Batianoff. SOUTH KENNEDY DISTRICT: Mackay, Apr 1975, AdcDonald 1301 & Batianoff. PoRT Curtis DISTRIcT: Emu Park, May 1975, Simon 2559 & Andrews. Vanuatu. Erromanga, May 1928, Kajewski 278; Tanna, Jun 1978, Morat 6011, 6022. New Caledonia. La Roche percee, Apr 1967, Schmid 1995 (NOU); Lifou, May 1969, Schinid 2829 (NOU); LF.O. Anse Vata, Jun 1963, Blanchon 179 (NOU). Fiji. Naitonitoni, Serua, Aug 1954, Shradha Nand 8652.
Cyrtecoccum Stapt
Cyrtococcum is a genus of 12 species native to the Old World tropics and its diagnostic features include the lateral compression of the spikelets, a crested apex on the upper lemma and a gibbous second glume and upper lemma. Until now only one species, C. oxyphyllum (Steudel) Stapf, has been recorded from Australia and that from the rainforests of north Queensland with a range extending to the tropics of Malesia, Melanesia and Asia. However John Clarkson and John Neldner, Queensland Herbarium Senior Botanists stationed at Mareeba, recently conducted a survey of the Batavia Downs region of Cape York Peninsula and among their collections was a delicate new species of Cyrtococcum.
Cyrtococcum capitis-york B. Simon, sp. nov. C. deccanensi Bor affinis sed spiculis parvioribus, C. patenti (L.) A. Camus affinis, sed spiculis grandioribus et pau- cioribus differt. Typus: Queensland. Cook District: 10.8 km S of Batavia Downs on Peninsula Development Road, 21 April 1990, J.R. Clarkson 8477 & VU. Neldner (holo: BRI(AQ 463916); iso: B,BRILDNA,NSW). Fig. 3.
Culms stoloniferous, basally decumbent, not tufted, 15-30 cm tall, 7—9-noded, sparingly branched, terminated by a solitary inflorescence. Internodes longer than the associated leaf sheaths. Sheaths slightly compressed and markedly ribbed. Ligule a membrane, c. 0.4 mm long. Leaf blades flat, lanceolate, 2.5-7.0 cm X 3-5 mm, glabrous, sparsely
Simon, Australian Grasses 5 593
Fig. 3. Cyrtococcum capitis-york: A. habit X 0.6. B. spikelet, lateral view x 16. C. lower glume, back view. D. upper glume, front view. E. lower lemma, back view. F. upper lemma, lateral view. G. upper palea, lateral view. (D-G, X 25). From type specimen.
594 Austrobaileya 3(4): 1992
pubescent adaxially, with scabrous margins. Inflorescence a panicle with main axis 2-8 cm long, smooth or rough. Primary branches spreading, not whorled, c. 4 cm long, smooth on the margins. Pedicels 2-5 mm long, distinctly angled, smooth, straight or twisted. Disarticulation at the base of the spikelet. Spikelets abaxial, c. 10 on a typical lowermost primary branch, laterally compressed, obliquely obovate in outline, 1.5-1.7 x 0.8-1.0 mm. Lower glume triangular or ovate, 0.6-0.8 mm long, 3-nerved, membranous, scabrous, setose with hairs tubercle-based. Upper glume obovate, c. 1. 5 mm long, 3- nerved, rounded on the back, membranous, hairy with tubercle-based hairs. Rachilla not conspicuous between the glumes. Lower lemma obovate, membranous, hairy, over- topping the spikelet, acute or rounded apically. Palea of lower floret absent. Upper floret perfect, shorter than the lower floret. Upper lemma white, softly cartilaginous smooth, gibbous, navicular, rounded on the back, glabrous, apically rounded. Palea of upper floret softly indurate, smooth. Anthers c. | mm long. Caryopsis not seen.
Conservation status: 1K (Briggs & Leigh 1988). Etymology: Named for Cape York.
Notes: Cyrfococcum capitis-york is very close to C. deccanense Bor, itself a distinctive species with local distributions in India and Sri.Lanka (Bor 1960), but has slightly smaller spikelets (1.5-1.7 mm long compared to 1.8-2.0 mm long in C. deccanense) in which its hairs are tubercle-based. Both these species have a sparse inflorescence of relatively few spikelets compared to that of C. patens (L.) A. Camus with a dense inflorescence of many spikelets, a species relatively widespread in southeast Asia with which C. ica ioigid was compared when it was described. These three species all have hairy spikelets
Panicum L..
Panicum is the largest genus in the grass family with recent estimates figures ranging from + 470 species (Clayton & Renvoize 1986) to + 600 species (Zuloaga 1987). As circumscribed last century the genus was even larger, but it was divided into smaller genera on the basis of well-defined morphological characters in the early part of this century by Chase (1906-1911) for the New World species and by Stapf (1917-1930) for the African species. Hughes (1923), working on the Australian species and following the example set by Stapf for Africa, divided Bentham’s genus Panicum in his Flora Aus- traliensis (1878) into 14 genera, leaving 22 species in Panicum. However Panicum is still a very variable genus and, together with its allied genera, is in need of critical revision at world level to make sense of this variability (Brown 1977). Nevertheless three Australian species of Panicum, two of them recognised in Simon (1990) as Panicum species and one as a species of Yakirra, require formal description.
Panicum bombycinum B. Simon, sp. noy. P. decomposito R. Br. affinis, sed glumis inferis multis longioribus et foliis bombycinis, P. gueenslandico Domin affinis, sed spiculis brevioribus et foliis bombycinis differt. Typus: Queensland. SouUTH KENNEDY DISTRICT, 27 km W of Mirtna HS, 21°18’46”S, 145°57’47”E, open savanna woodland, recently burnt, dominated by Eucalyptus whitei, with E. papuana and FE. brachycarpa on sandy soil, 6 April 1992, E./. Thompson BUC 1418 & B.K. Simon (holo: BRI{AQ560012); 1so: AD,B,BRI,CANB,DNA,K,L,MEL,MO, NSW,PERTH, US).
Plants perennial. Culms erect, tufted, to 35 cm tall, 2-4-noded, unbranched, terminated by a solitary inflorescence. Internodes shorter than the associated leaf sheaths. Sheaths rounded on the back. Ligule a fringe of hairs c. 1.5 mm long. Leaf blades linear, flat or involute, 9-21 cm * 2-4 mm, hairy, sericeous with scaberulous margins. Inflorescence a panicle with main axis to 12 cm long, scabrous. Primary branches spreading, to 6 cm long, scabrous on the margins. Pedicels 5-20 mm long, distinctly angled, scabrous, straight. Disarticulation at the base of the primary branches. Spikelets abaxial, 3-5 on a typical lowermost primary branch, dorsiventral compressed, ovate in outline, 3.0—3.5 x 1.0-1.5 mm. Lower glume triangular, acute, 1.7-2.0 mm long, 5- sub 7-nerved,
membranous, smooth, glabrous. Upper glume ovate, 3.0-3.5 mm long, 9- sub 1 1-nerved, rounded on the back, membranous, glabrous. Rachilla not conspicuous between the
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596 Austrobaileya 3(4); 1992
glumes. Lower lemma ovate, 3.0-3.5 mm long, membranous, glabrous, acuminate. Palea of lower floret vestigial, ovate, acute. Upper floret perfect, shorter than the lower floret. Upper lemma elliptic, 2.0-2.3 mm long, apically rounded, brown, coriaceous, smooth, rounded on the back, glabrous. Palea of upper floret coriaceous, smooth. Anthers c. 1.5 mm long. Caryopsis not seen. Fig. 4.
Additional specimens examined. Queensland. BURKE DISTRICT: near head of Poison Creek, about 90 mi [144 km] N of Hughenden, on compact sand in Eucalyptus forest, Apr 1945, Blake 8540 (Fig. 4); SourH KENNEDY District: 14.5 km SW of Mirtna on shot-line 35 km NW of Mirtna-Yarromere road, Apr 1992, Thompson BUC 146 & Simon (BRI,MBA).
Conservation status: 3K (Briggs & Leigh 1988). Etymology: Named for the silky, velvety hairs covering the leaf sheaths and blades.
Notes: Panicum bombycinum 1s distinctive of the Australian species of Panicum in that its leaf sheaths and blades have a dense covering of silky, velvety hairs. It differs also from P. decompositum by its longer lower glume (1.7-2.0 mm long compared to 0.5- 1.0 mm long in P. decompositum) long and differs from P. queenslandicum by its shorter spikelets (3.0-3.5 mm long compared to 3.5-5.0 mm long in P. queenslandicum). The spikelets thus appear plumper as they have the same width as the spikelets of P. queenslandicum (1.0-1.5 mm wide).
Because this species was originally thought to have an appendage at the base of the fertile, upper floret it was placed with Yakirra (as Yakirra sp. Blake 8570) in Simon (1990), but closer examination (Fig. 4) revealed it to have no such appendage. Furthermore the rachilla between the glumes is not conspicuous as in species of Yakirra, so the placement of this species in Panicum seems logical on the present evidence.
Panicum chillagoanum B. Simon, sp. nov. P. seminudo Domin affinis, sed spiculis brevioribus, P. mitchellio Benth. affinis, sed habitu annuo, habitans terram calcaream differt. Typus: Queensland. CooK DISTRICT: Chillagoe-Almaden road 8 km from Chillagoe, limestone outcrop, 8 March 1980, B.K. Simon 3556 & JR. Clarkson (holo: BRI(AQ 381541); iso: BRI,CANB,DNA,K,L,MBA,NSW). Fig. 5.
Plants annual. Culms erect, tufted, 15-60 cm tall, 2-3-noded, unbranched, terminated by a solitary inflorescence. Internodes shorter than the associated leaf sheaths. Sheaths slightly compressed. Ligule a fringed membrane c. 1 mm long, with marginal tubercle- based hairs at the sheath blade junction. Leaf blades flat, linear or triangular, 2-12 cm x 2-5 mm, hairy with a few tubercle-based hairs along nerves, especially the midrib, with scaberulous margins. Inflorescence a panicle with main axis 4-25 cm long and smooth. Primary branches spreading, 3-14 cm long, scabrous on the margins. Pedicels 3-12 mm long, distinctly angled, scabrous, straight. Disarticulation at the base of the spikelet. Spikelets abaxial, 15-20 on a typical lowermost primary branch, dorsiventral compressed although laterally compressed at apex, lanceolate in outline, 2.1-2.7 mm X 0.8-1.0 mm. Lower glume triangular, 0.9-1.2 mm long, 5-nerved, acute, membranous, smooth, glabrous. Upper glume ovate, 2.1-2.7 mm long, 5-nerved, apically acuminate, rounded on the back, membranous, glabrous. Lower lemma ovate, slightly shorter than upper glume, membranous, glabrous, apically acuminate. Rachilla conspicuous between the glumes. Palea of lower floret vestigial, apically truncate. Upper floret perfect, shorter than the lower floret. Upper lemma oblong, c. 1.5 mm long, brown, chartaceous, smooth, rounded on the back, glabrous. Palea of upper floret chartaceous, smooth. Anthers c. 1.5 mm long. Caryopsis not observed.
Conservation status: 1K (Briggs & Leigh 1988). Etymology: Named for the Chillagoe area, where it appears restricted.
Notes: Panicum chillagoanum differs from P. seminudum Domin by its smaller spikelets (2.1-2.7 mm long compared to 3.1-3.7 long in P. seminudum) and it differs from P. mitchellii Benth. by its annual habit and by its being restricted to limestone habitats of the Chillagoe area as opposed to P. mitchellii being perennial and occurring in non- calcareous wet sclerophyll forests and woodlands of north Queensland. This species has been collected only once, from a limestone outcrop in the Chillagoe area where it was
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598 Austrobaileya 3(4): 1992
locally fairly common. In its endemicity to this region of Queensland resembles other species such as Atalaya calcicola Reyn. (Sapindaceae) and Glossocarya calcicola Domin (Verbenaceae).
Panicum robustum B. Simon, sp. nov. P. trachyrhachidi Benth. affinis, sed spiculis longioribus, glumis inferis longioribus quam glumis superis differt. Typus: Queens- land. NORTH KENNEDY DISTRICT: Brandy Creek road, 12 km from its source, 13 April 1978, B.K. Simon 3370, J.R. Clarkson & N.B. Byrnes (holo: BRI(AQ 344285); iso: BRI,CANB,DNA,K,L,MO,NSW). Fig. 6.
Plants robust annuals. Culms erect or sometimes basally decumbent, weakly tufted, 80- 180 cm tall, 3-5-noded, sparingly branched or rarely unbranched. Internodes longer than the associated leaf sheaths. Sheaths rounded on the back. Ligule a fringed membrane c. 2 mm long. Leaf blades flat, linear, 10-35 cm X 5-10 mm, with a distinctive white midrib, hairy, hispid with tubercle-based hairs, and scaberulous on margins. Inflorescence a panicle with main axis 25-40 cm long, very lightly scabrous. Primary branches spreading, 10-18 cm long, scabrous on the margins. Pedicels 2-4 mm long, distinctly angled, scabrous, straight. Disarticulation at the base of the spikelet. Spikelets abaxial, dorsiventral compressed, lanceolate in outline, 4.5-5.0 X c. | mm. Lower glume lanceolate, 4.5-5.0 mm long, 5-7-nerved, membranous, smooth, glabrous, acuminate. Upper glume lanceo- late, 3.5~4.0 mm long, 5-7-nerved, rounded on the back, membranous, glabrous. Lower lemma lanceolate, 3.5-4.0 mm long, membranous, glabrous, apically acute. Rachilla conspicuous between the glumes. Palea of lower floret vestigial, cleft at the apex. Upper floret perfect, shorter than the lower floret. Upper lemma elliptic, c. 2 mm long, pale yellow, chartaceous, smooth, rounded on the back, glabrous, apically rounded. Palea of pone floret chartaceous, uniformly striate. Anthers c. 1.5 mm long. Caryopsis c. 2 mm ong.
Specimens examined: Queensland. Cook District: Laura River, Peninsular Development road, Apr 1983, Clarkson 4719 (BRI,CANB,K,QRS); Mareeba, Mar 1938, Blake 13479 (AD, BRI,CANB,DNA,K,L,MO,NSW,PERTH,PRE); Log Creek, 22 km W of Georgetown, Apr 1973, Henderson H1734 (BRI); Gilbert River crossing, 84 km WNW of Georgetown, Apr 1973, Henderson H1757 (BRI); Welcome Creek plateau, 13 km SSW of Battle Camp, via Cooktown, Jul 1990, Bean 1920 (BRI), BURKE DISTRICT: Poison Creek, near Mt. Sturgeon Station, Feb 1931, Hubbard 7726 & Winders (BRI,K). NoRTH KENNEDY DISTRICT: Burdekin River, 30 km NW of Charters Towers, Apr £978, Simon 3453 (BRI).
Conservation status: Not threatened (Briggs & Leigh 1988). Etymology: Named for the large culm and spikelet dimensions.
Notes: Panicum robustum is a robust annual species of rainforest clearings and margins and wet sclerophyll forests and woodlands has been collected a number of times from north Queensland. Like P. trachyrhachis it has large spikelets at least 4 mm long, differing in this respect from P. mindanaense with its spikelets up to 3 mm long. It is distinguished from P. trachyrhachis by its spikelets being more than 4.5 mm long and by its lower slume being longer than the upper one.
Paspalum L.
Paspalum is a tropical to subtropical genus of + 330 species (Clayton & Renvoize 1986), occurring mainly in the New World where they form an important component of the native grasslands. Of the 18 species recorded for Australia there are four fairly — common native species (although the two hydrophytic species P. distichurm and P. vaginatum are considered by Webster (1987) to be introduced) and two native species known from very limited material and which are described here.
Paspalum batianoffii B. Simon, sp. nov. P. notato Fluegge affinis, sed stolonibus, spiculis angustioribus, P. multinodo B. Simon affinis, sed stolonibus, spiculis longioribus, P. vaginato Sw. affinis, sed racemis et spiculis longioribus, rachidibus latioribus, differt. Typus: Queensland. PORT CuRTIS District: Statue Bay Beach, 6.5 km SE of Yeppoon, very narrow foredune with open woodland of Casuarina, Hibiscus and Excoecaria spp. and groundcover of Panicum maximum, Ipomoea pes-caprae and Zoysia macrantha; performing sand binding function just above high water
Simon, Australian Grasses 5 599
Fig. 6. Panicum robustum: A. plant X 0.4. B. spikelet x 8. C. lower giume, front view. D. upper glume, front view. E. lower lemma, front view. F. lower palea x 12. G. upper floret, front view. H. caryopsis, view of side with embryo. I. caryopsis, view of side with hilum. (C-I, x 12). From type specimen.
600 Austrobaileya 3(4): 1992
mark, 8 September 1977, G.N. Batianoff 651 & T.J. McDonald (holo: BRI(AQ 294456; 3 sheets BRI 294250, BRI 245882 and BRI 245883)). Fig. 7.
Plants. perennial. Culms stoloniferous, basally decumbent, to 40 cm tall, 2—3-noded, sparingly branched, terminated by a solitary inflorescence. Internodes shorter than the associated leaf sheaths. Sheaths rounded on the back. Ligule a membrane c. | mm long, entire. Leaf blades linear, flat or convolute, 8-16 cm X 2-4 mm, glabrous, with smooth margins, with marginal tubercle-based hairs at blade-sheath junction. Inflorescence of 2 or 3 spicate branches, with the main axis 1.5-2.5 cm long and smooth. Primary branches spreading, 6-9 cm long, smooth on the margins, rachis flattened, + 2 mm broad. Pedicels extremely short, c. 0.5 mm long, distinctly angled, smooth. Disarticulation at the base of the spikelet. Spikelets dorsiventral compressed, elliptic in outline, c. 4.5 * 1.5 mm. Lower glume absent. Upper glume elliptic, c. 4.5 mm long, 2- nerved, rounded on the back, coriaceous, glabrous. Lower lemma elliptic, coriaceous, the surface glabrous, acute, c. 4.2 mm long. Palea of lower floret absent. Upper floret perfect, slightly shorter than the lower floret. Upper lemma c. 3.5 mm long, yellow, coriaceous, smooth, elliptic, rounded on the back, glabrous, acute. Palea of upper floret coriaceous, smooth. Anthers c. 2 mm long. Caryopsis not observed.
Conservation status: This species is known only from the type and has been known about for a number of years by its inclusion in Thomas and McDonald (1987) as Paspalum sp. ‘Statue Bay’ (G.N. Batianoff 651) and in Briggs and Leigh (1988) as Paspaium sp. | (Statue Bay). In both these publications the designation 1K was assigned to it. Two further attempts to locate and collect the species at the type locality have not been successful so there is a possibility it should be designated 1X.
Etymology: The species is named for George Batianoff, a member of the staff of the Queensland Herbarium, who has shown great enthusiasm 1n collecting plants from coastal Queensland and associated islands.
Notes: Paspalum batianoffii differs from P. notatum Fluegge by its stoloniferous culms, longer and narrower spikelets (c-4.5 X 1.5 mm compared to 2.8-3.7 * 2.0-2.8 mm in P, notatum). It differs from P. multinodum B. Simon by its stoloniferous culms, fewer nodes (2-3 nodes compared to 10-13 nodes in P. multinodum) and longer spikelets (c. 4.5 mm long compared to c. 3.5 mm long in P. multinodum). It differs from P. vaginatum Sw. by its longer inflorescence branches (6-9 cm long compared to 2-5 cm long in P. vaginatum) and its longer spikelets (c. 4.5 mm long compared to 2.5-3.7 mm long in P. yaginatum) and by its broader rachis (c. 2 mm broad compared to c. | mm broad in P. vaginatum).
Paspalum multinodum B. Simon, sp. noy. P. wotato Fluegge affinis, sed habitu caespitosis et spiculis angustioribus, P. batianoffio B. Simon affinis, sed habitu caespitosis, spiculis brevioribus, P. vaginato Sw. affinis, sed habitu caespitosis, rachidibus latioribus, P. scrobiculato L.. affinis, sed. culmis elatioribus, rachidibus latioribus differt. Typus: Queensland. Cook District: Aurukun, collector unknown s.n., (holo: BRI(AQ 540191); iso: CANB,K,L). Fig. 8.
Plants perennial. Culms erect, tufted, 1.5-2.0 m tall, 10-13-noded, unbranched, terminated by a solitary inflorescence. Internodes longer or shorter (distally) than the associated leaf sheaths. Sheaths keeled and compressed. Ligule a membrane 1.0-1.5 mm long. Leaf blades flat, linear, to 38 cm X 5 mm, glabrous, with smooth margins. Inflorescence spiciform with 2-6 racemes on main axis 0.5-1.0 cm long and smooth. Primary branches spreading, 3-8 cm long, smooth on the margins, rachis flattened, + 2 mm broad. Pedicels extremely short, c. 0.5 mm long, smooth, straight. Disarticulation at the base of the spikelet. Spikelets dorsiventral compressed, elliptic in outline, c. 3.5 X 1.5 mm. Lower slume absent. Upper glume elliptic, c. 3.5 mm long, 3-nerved, rounded on the back, membranous, glabrous. Lower lemma elliptic, membranous, glabrous, subacute. Palea of lower floret absent. Upper floret perfect, shorter than the lower floret. Upper lemma elliptic, c. 3 mm long, white or yellow, coriaceous, smooth, rounded on the back, glabrous, a er aa Palea of upper floret coriaceous, smooth. Anthers and caryopsis not observed.
Additional specimen examined: Queensland, CooK DISTRICT: Mapoon Plain S of Cullen Point, N of Weipa, 11°5- ’S, 141°5’E, seasonally cracking clay plain, Sep 1980, Godwin A52 (BRI).
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Fig. 7. Paspalum batianoffit: A. base of culm X 0.7. B. apex of culm with inflorescence X 0.5. C. portion of rachis of a spike X 4. D, spikelet X 8. E. upper glume, front view. F. lower lemma, front view. G. upper lemma, front view. H. upper palea, front view. J. anthers. (E-I, X 12). From type specimen.
602 Austrobaileya 3(4): 1992
Conservation status: 2K (Briggs & Leigh 1988). Etymology: The name 1s derived from the many-noded culms.
Notes: P. multinodum is similar to P. batianoffii B. Simon in having a broad rachis, but differs from that species by its smaller spikelets (c. 3.5 mm long compared to c. 4.5 mm long in P. batianoffii) and its tufted habit. It differs from P. notatum Fluegge by its narrower spikelets (c. 1.5 mm wide compared to 2.0-2.8 mm wide in P. notatum) and its tufted habit. It differs from P. vaginaturm Sw. by its broader rachis (c. 2 mm broad compared to c. | mm broad in P. vaginatum) and tufted habit. It differs from P. scrobiculatum L. by its taller culms (1.5 to 2 m tall compared to less than 1.5 m in P. scrobiculatum) and its broader rachis (c. 2 mm broad compared to c. | mm broad in P. scrobiculatum). Only two collections of this species are known. One is from a plant grown from seed collected at Aurukun by a collector not designated and with the name of ““Moonpoon Grass’, according to a note attached to the specimen by S.T. Blake. This has been selected as the type, despite the scanty field information associated with it, as it has a number of duplicates that will be distributed to other herbaria. The other is a unicate specimen from the Mapoon Plain south of Cullen, and presumably the source of the name ““Moonpoon” in “Moonpoon Grass”’.
It was originally thought that P. batianoffii and P. muitinodum may be naturalised New World species but material of them could not be matched from the American collections of Paspalum in the Kew herbarium (S.A. Renvoize pers. comm.), nor were fragments and photographs of them recognised by Dr Fernando Zuloaga, a recognised authority of the genera Paspalum from the herbarium of the Instituto de Botanica Darwinion, Buenos Aires (SJ). It is on the basis of these communications that I decided to name them as new species.
Yakirra Lazarides & R. Webster
Diagnostic features of Yakirra include a straight, swollen rachilla between the florets, a conspicuous rachilla between the glumes and a smooth surface on the upper floret. All species except Y. nulla Lazarides & R. Webster also have two appendages from the apex of the stipe of the fertile, upper floret There are two specimens of a new species of Yakirra from western Queensland which have these stipe appendages but they are short and hard compared to the appendages in all other species of this genus which have them. Superficially this entity resembles Panicum effusum var. effusum.
Yakirra websteri B. Simon, sp. nov. Y. mue//eri (Hughes) Lazarides et R. Webster, Y. majusculae (F. Muell. ex Benth. ) Lazarides et R. Webster, Y. australiensi (Domin) Lazarides et R. Webster et Y. pauciflorae (R. Br.) Lazarides et R. Webster affinis, sed stipitis flosculi fertili appendicibus brevibus et duris, flosculo supero brunneo vel luteo et habitu perenni differt. Typus: Queensland. MITCHELL DISTRICT: 93 k N of Langlo Crossing, 20 May 1975, G.R. Beeston 1361C, (holo: BRI(AQ 268164); 1s0: BRI,CANB,K,NSW) Fig. 9
Plants perennial. Culms erect, tufted, 25-80 cm tall, 2-3-noded, unbranched, terminated by a solitary inflorescence. Internodes shorter than the associated leaf sheaths. Sheaths rounded on the back. Ligule a fringe of hairs c. 0.5 mm long. Leaf blades linear, flat, 7-12 cm X 2-5 mm, pubescent with tubercle-based hairs; margins scaberulous. Inflo- rescence a panicle with main axis 12-20 cm long, hairy with tubercle-based hairs. Primary branches spreading, to 10 cm long (longest at base of rachis), scabrous on the margins. Pedicels 2-8 mm long, distinctly angled, scabrous, straight. Disarticulation at the base of the spikelet. Spikelets abaxial, dorsiventral compressed, ovate in outline, 2.5 * 1.5 mm. Lower glume triangular, c. 1.5 mm long, 3- sub 5-nerved, membranous, scabrous on nerves, glabrous, acute. Upper glume ovate, c. 2.5 mm long, 9-nerved, rounded on the back, membranous, glabrous. Rachilla conspicuous between the glumes. Lower lemma ovate, membranous, slabrous, acuminate. Palea of lower floret ovate, acute. Upper floret perfect, shorter than the lower floret. Basal stipe producing two small hard appendages. Upper lemma elliptic, c. 1.8 mm long, brown with white nerves or yellow, chartaceous, rounded on the back, glabrous, apically rounded. Palea of upper floret chartaceous,
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604 Austrobaileya 3(4): 1992
uniformly striate. Anthers not observed. Caryopsis c. 1.5 mm long with hilum less than half as long as caryopsis.
Additional specimen examined. Queensland. MARANOA DiSTRICT: 39 km from Roma on Injune road, May 1975, Simon 2859 & Clarkson (BRI).
Conservation status: 3K (Briggs & Leigh 1988).
Etymology: The species 1s named for Robert Webster, U.S.D.A. at Beltsville, who studied the Australian taxa of the tribe Paniceae at the Research School of Biological Sciences, A.N.U., using DELTA and is currently editor of the DELTA Newsletter.
Notes: Yakirra websteri differs from Y. muelleri (Hughes) Lazarides et R. Webster, Y. majuscula (F. Muell. ex Benth.) Lazarides et R. Webster, Y. australiensis (Domin) Lazarides et R. Webster and Y. pauciflora (R. Br.) Lazarides et R. Webster by the stipe of the upper, fertile floret having shorter and hardened appendages by the fertile floret being brown to yellow in colour and by the plant having a perennial habit.
Acknowledgements
I extend my gratitude to colleagues on the staff of the Queensland Herbarium for preparing the figures, Will Smith for the fine illustrations of Arthragrostis clarksoniana, Brachiaria atrisola, Cyrtococcum capitis-york, Panicum robustum, Paspalum batianoffii and Yakirra websteri, and Hans Dillewaard for the photographs of the types of Panicum bombycinum, Panicum chillagoanum and Paspalum multinodum. I thank Rod Henderson for a critical scrutiny of and improvement to the manuscript. I also thank Greg Leach, Australian Botanical Liaison Officer at Kew, 1990-1991, for providing me with a copy of Neuyen (1966) and Gerrit Davidse, Senior Curator of the herbarium at the Missouri Botanical Garden for working facilities during my visit to St. Louis in 1991.
References BENTHAM, G. (1878). Flora Australiensis 7: 463-491. London: L. Reeve & Co. BENTHAM, G. (1883). In G. Bentham & J.D. Hooker, Genera Plantarum 3: 1100-1103. London: L. Reeve & Co.
BEA S.T. (1948). Studies in Queensland grasses, II]. Proceedings of the Royal Society of Queensland 59: 153- 160. BLAKE, S.T. (1958). New criteria for distinguishing genera allied to Panicum (Gramineae), Proceedings of the
Royal Society of Queensland 70: 15-19.
BLAKE, S.T. (1969). Taxonomic and nomenclatural studies in the Gramineae, No, 2. Proceedings of the Royal Society of Queensland 81: 1-26.
BLAKE, S.T. (1973). Taxonomic and nomenclatural studies in the Gramineae, No. 3. Proceedings of the Royal Society o Queensland 84: 61-70.
BOR, N.L. (1960). The grasses of Burma, Ceylon, India and Pakistan. London: Pergamon Press.
BRIGGS, J.D. & LEIGH, J.H. (1988). Rare or Threatened Australian Plants. 1988 Revised Edition. Australian serra Parks and Wildlife Service Special Publication Ne. 14. Canberra: Australian National Parks and Wildlife Service.
BROWN, W.YV (1977). The Kranz syndrome and its NYO in grass systematics. Mfemoirs of the Torrey Botanical Club 23: 1- 97, 126-130.
BUTZIN, F. (1970). Die systematische Gliederung der Paniceae. Willdenowia 6: 179-192.
CHASE, A.S. (1906-1911). Notes on genera of Paniceae, J-lV. Proceedings of the Biological Seciety of Washington. (1906: I, 19: 184-192; 1908: II, 21: 1-10; IIL, 21: 175-188; 1911: TV, 24: 103-160).
CHASE, A.S. (1920). The North American species of Brachiaria. In A.S. Hitchcock & A. Chase, Revisions of North American grasses. Contributions of the United States National Herbarium ( Washington) 22(1): 33-43 (Uchnanthus, Lasiacis, Brachiaria, Cenchrus).
CHIPPINDALL, L.K.A. (1955). A guide to the identification of grasses in South Africa, pp. 5-527. In D. Meredith (ed.), The Grasses and Pastures of South Africa. Cape Town: C.N.A.
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Fig. 9. Yakirra websteri: A. plant X 0.33. B. spikelet, lateral view X 16. C. lower glume, front view. D. upper glume, front view. E. lower floret, front view, showing lemma enclosing palea. F. upper floret, front view, showing the two basal appendages. G. caryopsis, view of side with embryo. H. caryopsis, view of side with hilum. (C-H, x 25). From type specimen.
DYN ec NI a ec DAM ANA AC DANAE ERMA
gg one nimim men nen neg romney tN
606 Austrobaileya 3(4): 1992 CLAYTON, W.D. & RENVOIZE S.A. (1982). Flora of Tropical East Africa, Gramineae 3: 575-600 (Brachiaria). Rotterdam: A.A. Balkema.
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Accepted for publication 12 February 1992
SS RL an Cod coon ec Can Dad ent Ca oan ard ave ten Ta
Austrobaileya 3(4): 609-614 (1992) 609
PEROTIS AITON (GRAMINEAE) IN AUSTRALIA AND SOUTHEAST ASIA
J.F. Veldkamp and H. van Steenbergen Rijksherbarium, P.O. Box 9514, 2300 RA Leiden, The Netherlands
Summary
A survey of the 3 species of Perotis Aiton (Gramineae) in Australia and Southeast Asia is given. One species, P. clarksonii is described as new, while P. rara is recorded from Southeast Asia for the first time.
Introduction
Perotis Aiton (Gramineae) is a small genus of about 10 Old World tropical species. By some (Hubbard in Bor 1960) it has been regarded as belonging to a separate tribe, Perotideae, while others have included it in the Lappagineae (Pilger 1956), Zoysieae (Jacques-Felix 1962), or Cynodonteae-Zoysiinae (Clayton & Renvoize (1986).
Until about 10 years ago only one species, P. rara R. Br., was known to occur in Australia. Simon (1981) mentioned the presence of P. indica (L.) Kuntze in Queensland, and a third species, new and closely related to P. rara, also from Queensland, is described here.
Perotis
Perotis Aiton, Hort. Kew. 1: 85 (1789). Type: Perotis latifolia Aiton, nom. superfl. (based on Saccharum spicatum L. = Perotis indica (L.) Kuntze). Xystidium Trin., Fund. Agrost. 102, t. 2 (1820). Type: Xystidium maritimum Trin. (= Perotis rara R. Br,).
Annuals (in Australia). Ligules collar-shaped, membranous. Blades broadest at the rounded to amplexicaul base. Inflorescence a spike-like raceme. Spikelets 1-flowered, solitary, falling entire at maturity, pedicels persistent. Glumes enclosing the floret, dorsally rounded, l-nerved, long-awned. Rachilla process absent. Lemma membranous, 3-nerved, acute, muticous. Anthers 3. x = 10.
Distribution: About 10 species in the Old World tropics, 3 in Australia. Anatomy: See Clayton and Richardson, Kew Bull. 27: 44 (1973).
Key to the species
1. Racemes laxly spikeled; spikelets at maturity reflexed; lower glume grad- ually passing into awn, body not distinct from awn; callus up to 1.5
mm long, obconical, slightly flattened, laterally pubescent ee he tons 2 Racemes densely spikeled: spikelets at maturity patent; lower glume with the body distinct from awn; callus up to 0.3 mm nies terete, stipe-
like, puberulous all over. Caryopses FEveTC! vishal BS ar, we ee: 2. Perotis indica
2. Leafblades stiffly patent, distichous at short intervals; glumes smooth, glabrous to shortly pubescent; awns at base with c. 1 mm long, spirally twisted hairs along the infolded margins; caryopses flattened, sulcate .... 1. Perotis clarksonii Leafblades more or less erect, rather flaccid, not distinctly distichous: glumes more or less scaberulous to setulose: awns rarely setulose at base, bristles then at most 0.5 mm long, Vileaast caryopses terete Ae esse Se eR ear ee ......... 3 Perotis rara
1. Perotis clarksonii Veldk., sp. nov.
Planta annua, ad 40 cm alta; foliorum laminae patentes, |.7—2.5 cm longae, 1.7-2,.3 mm latae, planae ad involutae; racemi laxi, 12~27 cm longi, incomplete exserti; spiculae patentes sub anthesi, (35-47. 5-55 mm longae (callo 0.75-1.5 mm longo, obconico leniter complanato, saltem lateraliter pubescenti excluso), glumae gradatim in aristis transientes laeves glabrae ad tote breviter pubescentes, costa scaberula; aristae longe pilosae ad basin pilis lenibus spiraliter tortis ad | mm longis argenteis; caryopsides complanatae sulcatae.
610 Austrobaileya 3(4): 1992
Typus: Australia, Queensland. Cook DISTRICT: 0.8 km South of the Laura River crossing on the Peninsula Development road, 15°35’S, 144°27’E, 90 m altitude, 7 March 1987, Clarkson & McDonald 6802 (holo: L!; iso: BRI,CANB fragm.!, DNA,NSW!,PERTH,QRS).
Annual, up to 40 cm high. Leaf blades patent, 1.7-2.5 cm * 1.7-2.3 mm, flat to infolded, bristly at base and along the throat of the sheaths. Racemes lax, 12- 27 cm long, not completely exserted. Spikelets patent at anthesis, (35—)47.5-55.0 mm long (excluding the 0.75-1.50 mm long, obconical, slightly flattened, at least laterally pubescent callus). Glumes passing gradually into the awns, smooth, slabrous to shortly pubescent all over, midrib scaberulous, awns long-hairy at base, hairs soft, becoming spirally twisted, up to 1 mm long, silvery. Caryopsis flattened, sulcate. Fig, 1
Specimens examined: Australia, Queensland. Cook DISTRICT: 30-40 miles [48- 64 km] S of Coen, 14°15’S, 143° 10°E, A 4843 (ADW,AHUC, CANB!, K, L!); Koolburra Station, 15°18’S, 143°587E, Clarkson 3152 (BRI, CANBI, L!); Clarkson & McDonald 6802, the type (see above); 15 km W of Battle Camp Road, 29 km NE of Laura, 15° 32'S, 144°27’E, Forster 4017 (BRI, L fragm.!); Chillagoe-Wrotham Park Road, 16°45’S, 144° S’E, STHi0n & Clarkson 3584 (BRI, CANBI, Li); 3 miles E of Mareeba, Thorne 21162 (BI,L fragm.!).
Distribution and habitat: Far north-eastern Australia, grows in savannah woodland with a grassy understorey, at low altitude.
Collector’s notes: Small, tufted grass; nodes red; underside of leaves dark wine red, dark sreen above; scape and inflorescence pale green.
Note: The awns of the glumes in P. rara exceptionally have rather long bristles up to 0.75 mm long, which then also have a tendency to become spirally twisted. An example from the Cook District is /4cKee 9241 (BRI,CANB!) from 5 miles (8 km) S of Dimbulah (c. 17°14’S, 145°6’E), where not only are the awns long-hairy but the glumes and rachis are as well. I (JFV) am not too sure whether this should or should not be regarded as a hairy form of P. rara or of P. clarksonii. A similar specimen is Thorne 21162 (BRI,L fragm.!,RSA) from 3 miles (4.8 km) E of Mareeba (Fig. 2A & B). In their leafblades they resemble P. clarksonii.
From the Northern Territory I have seen a few specimens with the P. rara type of leaf blades, very rough glumes, and awns with such long hairs. J am inclined to retain these in P. rara: Latz 564, Frew River, 3 miles (5 km) ENE of Epemarra Homestead, c. 20°30’S, 135°30°E (CANBI,NT), a mixture of ‘typical’ P. rara, suggesting that the hairy plant is just a form of that; Perry 3258, 9 miles (14.4 km) S of Undoolya Station, 23°41’S, owe Winkworth 815, 15 miles (24 km) SW of Alcoota, 22°50’S, 134°27’E (CANB!).
Etymology: Named after Mr John Richard Clarkson, Mareeba, who collected three out of the six known representatives of this new taxon, in recognition of the many important collections he has made in northern Queensland.
2. Perotis indica (L.) Kuntze, Rev. Gen. Pl. 2 787 (1891); Anthoxanthum indicum L., Sp. Pl. 1: 28 (1753). Type: Hermann s.n. (holo: LINN).
Saccharum spicatum L., Sp. Pl. 1 54 (1753); Perotis latifolia Aiton, Hort. Kew. 1: 85 (1789), nom. superfl., Perotis spicata (L.) Dur. & Dur., Syll. Fl. congol. 628 (1909); Perotis latifolia var. typica Domin., Bibl. Bot. 85: 285 (1915), nom. inval. Lectotype: Hb. Linn. 77.5 (holo: LINN).
Alopecurus bengalensis Houtt., Nat. Hist. 2/13: 206, t. 90, f. 4. (1782) Type: not indicated (not found in Hb. Houttuyn in G or L).
Perotis hordeiformis Nees in Hook. & Arn., Bot. Beechey Voy. 248 (1838); Fl. Afr. Austr. 1: 139 (June 1841). Type: Royle 280 (holo: K; iso: LIY).
Perotis glabrata Steud., Syn. 1: 186 (1854). Type: Cuming 1399 (holo: P; iso: L).
Perotis birmanica Gand., Bull. Soc. Bot. France 66: 301 (1920). Type: Mokin 560 (holo: P).
Perotis chinensis Gand., Bull. Soc, Bot. France 66: 301 (1920). Type: Chanet 14 (syn: P), 137 (syn: P US) (Jisosyn: an unnumbered Chanet collection in L).
Perotis perrottetii Gand., Bull. Soc. Bot. France 66: 301 (1920). Type: Perrottet 1290 (‘1390’) (holo: P; iso: ‘L).
Veldkamp & van Steenbergen, Perotis 611
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Fig. 1. Perotis clarksonii: A. Habit < 0.67. B. Spikelet x 4. C. Area at base of awns X 16. D. Caryopsis, ventral view X 16. A-D, Clarkson 3152, iso at BRI.
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612 Austrobaileya 3(4): 1992
Annuals, up to 75 cm high. Leafblades 1-3 cm X 2-10 mm, base more or less cordate to amplexicaul, usually bristly along the basal margins. Spike 2-20 cm long. Spikelets patent at maturity, 6.5-30.0 mm long (excluding the up to 0.3 mm long, terete, stipe- like, puberulous callus). Glumes scaberulous all over, with midrib scabrous; lower glume 1.5-2.5 mm long, not gradually passing into the up to 16 mm long awn. Caryopsis terete. 2n = 20, 40
Distribution and habitat: India to E China (Hopeh), throughout Malesia to Australia (Queensland, Cook District, Walsh River); introduced elsewhere in tropical countries. It srows in sandy soil near the coast, under Casuarina sp., 1n coconut plantations, dry grasslands, usually at low altitudes, ‘rarely as high as 1065 m in Malesia.
Note: In most works P. indica and P. hordeiformis are regarded as distinct and are distinguished as follows:
Callus acute, 0.2-0.5 mm long; glumes not conspicuously ciliolate on the keel; lower glume obscurely lobed, evenly and closely hairy all over, with hairs very short, white, appressed to spreading Ter . P. indica
Callus truncate, up to 0.2 mm long; glumes conspicuously Siolate on. 1 the keel: lower glume abruptly acute to more or less obtuse, with haus in close lines, especially at base Ce ee ee Se P. hordeiformis
In fact the length and shape of the callus are variable, and the hairs may be both irregular and in rows on the same specimen. Thus both names are taken to refer to only one species here.
3. Perotis rara R. Br., Prod. 172 (1810); Perotis rara var. typica Domin., Bibl. Bot. 85: 285 (1915), nom. inval. Type: R. Brown (holo: BM; iso: NSW).
Xystidium ils Trin.,, Fund. Agrost. 102, t. 2 (1820); Mem. Acad. Sc. St. Petersb. VI, 6: 266 (1915); Perotis rara var. maritima (Trin.) Domin., Bibl. Bot. 85: 285 (1915). Type: Chamisso s.n., Eschscholtz s.n. (syn: LE).
Xypstidium barbatum Presi, Rel. Haenk. 1: 228 (1830). Type: Haenke s.n. (holo: PR; iso: MO).
Perotis longiflora Nees in Hook. & Arn., Bot. Beechey Voy. 247 (1838); Perotis latifolia var. ones (Nees) Domin.., Bibl. Bot. 85: 285 (1915). Lectotype: Vachell 38 (holo: K; iso: GCE).
Perotis patula Nees = Hook. & Arn., Bot. Beechey Voy. 248 (1838). Type: Meyen s.n. (holo: K).
Perotis rara var. euryphylla Domin., J. Linn. Soc. 41: 274 (1912). Type: Clement s.n. (holo: K).
Perotis macrantha Honda, Bot. Mag. Tokyo 41: 638 (1927). Type: Yamazaki s.n. A® 1923 (holo: TT).
Annuals, up to 40 cm high. Blades 1-5 cm X 1-4 mm, rounded at base, bristly along the margins at base and in the throat. Spike 8-30 cm long. Spikelets reflexed at maturity, (7-)13-35 mm long (excluding the up to 1.5 mm long, obconical, slightly flattened, laterally pubescent callus). Glumes scaberulous all over, with midrib scabrous, gradually passing into the awns. Caryopsis terete. x = ?. Fig. 2C-F.
Distribution and habitat: Asia (Vietnam, Taiwan, SE China) to Australia (excluding Victoria, Tasmania). Grows on beach; eucalypt savanna; sandy river banks; locally common, 0-275 m altitude.
Uses: Decorative in flower; cattle are said not to like it but it might supply some slight grazing for sheep (Vickery, Fl. N.S.W. 19/2: 305 (1975)).
Vernacular name: Comet grass (E).
Note: This species is here first reported for continental Asia based on the following specimens from Vietnam present in L: d’Alleizette 2033 (Hanoi), Balansa s.n. (27 Oct.
Veldkamp & van Steenbergen, Perotis 613
WASMITH
Fig. 2. Intermediate specimen between Perotis rara and Perotis clarksonti: A. Spikelet_X 4. B. Area at base of awns X 16, C-F. Two specimens of Perotis rara. C,E. Spikelets X 4. D,F. Area at base of awns < 16, A,B, Thorne 21162, BRI; C,D Clarkson 5453, BRI; E,F Clarkson 5707, BRI.
1876, Quinhon), Balansa 378 (Baie de Fi-tsi-long), and Robinson 1039 (Natrang). It is also recorded for Taiwan (as P. macrantha) and China (as P. longiflora and P. patula).
Acknowledgements
This research was based on material present in L, CANB, and NSW; the latter two institutes were visited during a period of study at the Research School of Biological Sciences, Australian National University (A.N.U.), on A.N.U. and the Nederlandse Organisatie voor Wetenschappelijk Onderzoek (N.W.O.) grants, which are gratefully acknowledged. Mr B.K. Simon’s valuable advice on the specimens present in BRI and. the status of the present taxa were much appreciated. The synonymy of P. indica and P. hordeiformis 1s based on research undertaken during a course in Angiosperm Taxonomy at the Rijksherbarium by HvS. The figures of Mr William A. Smith, illustrator at BRI, are acknowledged.
References
CLAYTON, W.D. & RENVOIZE, S.A. (1986). Genera graminum: 255. Kew Bulletin Additional Series XIII. London: Her Mayjesty’s Stationery Office.
HUBBARD, le in N.L. BOR. (1960). The grasses of Burma, Ceylon, India and Pakistan: 611. Oxford: Pergamon Press Ltd.
614 Austrobaileya 3(4): 1992
JACQUES-FELIX, H. (1962). Les graminees d’Afrique tropicale. I: 230. Paris: Institut de Recherches Agronomiques Tropicales et des cultures Vivrieres.
PILGER, R. (1956). Das System der Gramineae. Botanische Jahrbticher fiir Systematik, Pflanzengeschichte und Pflanzengeographie 76: 349.
SIMON, B.K. (1981). Annual Report Queensland Herbarium 1979/1980: 23. Brisbane: Queensland Department of Primary Industries.
Accepted for publication 6 February 1992
Austrobaileya 3(4): 615-625 (1992) 615
STUDIES IN EUPHORBIACEAE A.L. JUSS., SENS. LAT. 2*. A REVISION OF NEOROEPERA MUELL. ARG. & F. MUELL. (OLDFIELDIOIDEAE KOHLER & WEBSTER, CALETIEAE MUELL. ARG.)
Rodney J.F. Henderson
Queensland Herbarium, Meters Road, Indooroopilly, Qld 4068, Australia
Summary
Neoroepera is endemic in central-eastern and north-eastern Queensiand, Australia. It contains two species, namely N. banksti Benth. and N. buxifolia Muell. Arg. & F. Muell. A lectotype is chosen for N. duxtfolia. Placement of the genus in Caletieae Muell. Arg. rather than Phyllantheae Dumort. is justified on grounds of its spinulose pollen and attributes of flowers, fruit and seed. Comments on dates of publication of parts of Adansonia 6 (1865-6) are siven in Appendix I.
Introduction
The genus Neoroepera was established by Johannes (Jean) Mueller (Mueller Argoviensis) and Ferdinand Mueller in the former’s contributions on Euphorbiaceae to the de Candolles’ great ‘Prodromus’ (Mueller 1866). It was accepted as containing only one species, NV. buxifolia, which was based on two specimens collected in central-eastern Queensland by Edward McArthur Bowman (1826-1872) for the latter Mueller, in Melbourne. By 1863, these specimens had reached the de Candolle herbarium in Geneva, Switzerland (G-DC), in time for the former Mueller’s study.
Since publication of Neoroepera and N. buxifolia, only one other species belonging to the genus has been recognised; that was named N. banksti and dealt with by Bentham in his ‘Flora Australiensis’ (1873).
Origin and author of name Neoroepera
The two specimens on which N. buxifolia is based are both mounted on the one sheet in G-DC. Duplicates of these collections are in MEL where again they are mounted on the one sheet (MEL 697068). A note by Bowman on that sheet indicates, however, the most likely course of events in relation to his material is as follows. To Mueller, in Melbourne, he sent flowering twigs with both copious male and a few female flowers (but no fruit or seed) from plants he found in central eastern Queensland. Mueller thought these specimens were from a new species of F/ueggea Willd. so labelled them ‘Flueggea buxifolia F.M.’ Later, Bowman sent fruiting and seed material of these plants to Mueller which then convinced him they were from a new species of Roeperia Sprengel, or possibly of a new genus (he tentatively named ‘Flueggella’), so annotated them either mon buxifolia J. & F. Mueller’ (MEL) or ‘Roepera (oder Flueggella) buxifoliad (G- DC).
Bowman’s note at MEL reads “55./Princhester./I sent this in flower in a former collection.” My experience with plants of this species growing in the Marlborough area is that ones copiously flowering do not have many fruit with mature seed at the same time and vice versa. Thus the material on sheets at G-DC and MEL must have been collected on two different occasions as Bowman indicated. Mueller, in his usual fashion, distributed material to Geneva, Paris and to Kew in London, and probably other places too, to assist workers then writing up groups of the Euphorbiaceae. In publishing Neoroepera, based on the material in Geneva, Johannes accorded Ferdinand joint authorship, though there is no evidence the latter was involved in selecting the name or preparing the protologue. Indeed, Bentham (1873), working with Kew material, accorded Mueller Argoviensis sole authorship of both the generic and species names and
(r90n) number 1, a revision of Aniperea Adr. Juss., was published in Australian Systematic Botany 5: 1-27 (1992).
616 Austrobatleya 3(4): 1992
cited F. Mueller’s manuscript “Roepera buxifolia’ in the synonymy of N. buxifolia when dealing with the species. Hutchinson (1969) and Webster (1975) followed Bentham in accepting only Mueller Argoviensis as author of the generic name, but this seems contrary to the latter’s wishes. As Johannes Mueller appears responsible for the generic name (he treated Roeperia Sprengel as a synonym of Ricinocarpos Dest. when publishing Neoroe- pera) and Ferdinand Mueller for the specific epithet, their wishes for joint authorship should be respected.
History of Classification
When formally describing Neoroepera with Ferdinand Mueller, Mueller Argoviensis (1866)! grouped the genus with Hemicyclia Wight & Arn. and Cyclostemon Blume within his subtribe Cyclostemoninae? in tribe Phyllantheae Dumort. Though Cyclostemon and Hemicyclia are now considered congeneric with Drypetes Vahl and in the distinct tribe Drypeteae (Griseb.) Hurusawa, Mueller considered Drypetes belonged in his subtribe Securineginae (“‘Securinegeae’) within tribe Phyllantheae. To him, this tribe fundamentally contained euphorbs with the lobes of the male perianth imbricate, anthers erect in bud, ovaries with 2-ovulate locules, and cotyledons two or more times wider than the radicle. Subtribe Cyclostemoninae was attributed flowers without petals and commonly in clusters, and with stamens inserted round and from below a central disc. While this gave Mueller a practical, rather simplistic grouping of relevant genera, it failed to take into account several other attributes now considered critical for indicating relationships of the genus Neoroepera. Besides, the anthers in Neoroepera are transverse on the apex of the filaments, somewhat similar to those in Mic ‘antheum Desf. and Beyeria Miq. (though held vertical in bud), but not like the erect ones in Phyllanthus and its allies.
From Stafleu and Cowan (1976, p. 97), it could be inferred that Baillon (1866) was the first to provide a name for the taxon here treated as genus Neoroepera when he described it as section Neoroepera of genus Securinega Juss. and named its sole species S. muelleriana, Though it seems this publication dates from July 1866, the relevant pages of ‘Adansonia’ probably appeared in September of that year (Appendix 1), some weeks after Mueller’s publication. Thus, his Sauropus sect. Neoroepera is a new com- bination of the Muellers’ Neoroepera, and his S. muelleriana is an illegitimate name (Greuter et a/. 1988) provided for Bowman’s Princhester Creek specimens.
Bentham (1873) maintained Neoroepera, on the basis of flowers of both sexes with a perianth, embryo with broad cotyledons and a narrow radicle, and two ovules in each ovary locule, in tribe Phyllantheae but did not recognise subtribes within that. He later (Bentham 1883) maintained Neoroepera within Phyllantheae but there associated it with several genera such as Phyllanthus L. (in a very broad sense), Sauropus Blume and Securinega Juss., none of which is now considered closely related to it.
The association of Neoroepera with Phyllanthus and its allies in the Phyllantheae persisted apparently until Hutchinson (1969) segregated it into the Drypeteae (Pax) Hutchinson (correctly Drypeteae (Griseb.) Hurusawa according to Webster 1975). As Hurusawa (1954) did not name Neoroepera anywhere in his account of the Euphorbiaceae, it is not clear where he intended it to be placed though presumably it remained within tribe Phyllantheae and covered by the ‘etc.’ in his list of genera included in that tribe. Hutchinson’s segregation was prompted, no doubt, by Pax’s inclusion of Neoroepera in subtribe Drypetinae of Phyllantheae in 1890 (Pax & Hoffman 1931) based, seemingly, principally on the disc in male flowers in Neoroepera being central. This genus invariably has carunculate seeds and N. banksii is a dioecious species. Pax apparently did not know that pollen of Neoroepera is spiny, fundamentally different from the non-spiny pollen of Drypetes. Thus this genus’ association with Drypetes, persisting from 1866 when the Muellers first described it, was perpetuated by Hutchinson even though he dissociated it from Phyllanthus.
With his different approach to classification within Euphorbiaceae, Webster assessed more than the traditional attributes used to divide the family and produced a new
' Published in late August 1866 according to Stafleu & Cowan 1976, p. 447.
2 “Cyclostemoneae’ Muell. Arg. (1865), equivalent to “Cyclostémonées’ of Batllon (1858).
Henderson, Neoroepera
classification of it (Webster 1975) based primarily on ovule numbers per ovary locule, morphology of poilen and other data from such fields as wood anatomy, cytology and biochemistry. Thus, based on the paired ovules in each ovary cell, the spiny pollen and carunculate seed, Webster grouped Neoroepera with Micrantheum Desf., Pseudanthus Sprengel and Stachystemon Planchon in Caletieae within sub-family Oldfieldioideae, totally dissociating it from Phyllanthus and Drypetes which he retained in subfamily Phyllanthoideae. Webster later admitted (1987) that his classification is by no means definitive or wholly satisfactory, for much information to test the robustness of his cheme is lacking. However, in the case of Neoroepera it seems logical to associate it with at least Micrantheum wherever that is placed, because, besides sharing the attributes
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ig. 1. Scanning electron micrographs of Neoroepera and Micrantheum pollen and seed: A. seed of N. buxifolia. . seed of N. banksii. C, seed of M. ericoides. D. pollen grain of M. hexandrum. E. . pollen grain of N. buxifolia. A, Bowman 55, MEL; B, Staples 2213, BRI; C, SW slopes of Mt Coolum, Qld,
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618 Austrobaileya 3(4): 1992
of two ovules per locule, spiny pollen (Fig. 1), and carunculate seed (Fig. 1) with that genus, it has a petaloid, single-whorled perianth, bilocular anthers transverse on the filaments, a central, gland-like disc lobed between the stamen filaments in male flowers, styles that are entire (though somewhat dorsi-ventrally expanded rather than more or less terete), and seeds somewhat tumid proximally about the hilum (at least in N. banksii) as do Micrantheum species. Thus, Webster’s transfer of Neoroepera to Caletieae seems fully justified and is accepted here.
This close relationship of Neoroepera with Micrantheum was noticed by F. Mueller who annotated a Walter specimen of N. banksii from Lizard Island in 1871 (MEL 697066) as ‘Neoroepera micrantheoides’ and commented that the plant was “similis Micrantheum hexandrum’”’, though he (? later) incorrectly accepted that the plants represented N. buxifolia ‘J. M. & F. M.’.
Whether or not the Oldfieldioideae warrants family status (as Paivaeusaceae) as suggested by Meeuse (1990), will have to wait till a more detailed study of the Euphorbiaceae sens. /at. 1s undertaken.
Origin of central structure in male flowers
The finding of a flower with a pistilode in a specimen of male N. banksii (Gittins 1833, in BRI and NSW), an attribute that occasionally occurs in the family (Baillon 1858: Webster 1984) but not to my knowledge recorded for Neoroepera before, helps clarify the nature of the central structure in male flowers. In this particular flower, the ovary, complete with three typical styles and stigmas but with the ovules abortive, arises from the centre of a gland-like, lobed structure otherwise typical of male flowers of this genus, that is, internal to the stamens and lobed with the lobes protruding between the filaments. Thus, the central structure found in male flowers of Neoroepera is homologous with the disc that subtends the ovary in female flowers and as such confirms traditional thought that it is a disc internal to the stamens, not a vestigial gynoecium as suggested by Baillon (1866).
Taxonomy
Neoroepera Muell. Arg. & F. Muell. in DC., Prodromus 15(2): 488 (August 1866); Securinega section Neoroepera (Muell. Arg. & F. Muell.) Baillon, Adansonia 6: 333 (September 1866). Type: NV. buxifolia Muell. Arg. & F. Muell.
Derivation of name: Named from Greek neos, new, and ‘Roepera after the generic name Roeperia Sprengel (a later synonym of Ricinocarpos Desf. also in the Euphorbiaceae) which honours Johann August Christian Roeper, German botanist, who, in the 1860s, was a professor at Basel (then Rostock) and who published on the Euphorbiaceae of Germany and Hungary (Baines 1981).
Shrubby monoecious or dioecious perennials with stems erect or ascending, branching, the branches leafy throughout. Leaves alternate, stipulate, shortly petiolate, persistent or caducous. Stipules entire. Flowers pedicellate, solitary or paired or in clusters or short spikes (reduced branchlets) in leaf axils, subtended by minute bracts; perianth lacking a corolla, petaloid, of several imbricate lobes; lobes dimorphic, usually a small sepal-like one alternating with a larger, petal-like one, + free. Male flowers 6(rarely 4, 5, 7 or 8)- merous; perianth lobes entire, emarginate, ciliate or shortly erose; a whorl of few to many, discrete, finger-like glands present between tepals and stamens; stamens exserted; anthers of two, separate, obloid, parallel but contiguous locules each transverse on the apex of a reduced lobe of the shortly bifid filament, dehiscing by longitudinal slits; disc a central, squat, + sessile, lobed structure embracing base of filaments. Female flowers 6(~8)-merous; perianth persistent, of + distinct, imbricate lobes; lobes appressed to ovary but spreading in fruit, entire or somewhat ciliate or erose on upper margins; glands in two whorls, the outer of few, discrete finger-like lobes, the inner of discrete, flattened, irregular lobes on a continuous rim. Ovary 3(rarely 4)-celled with two pendant ovules from an enlarged placenta in each locule; styles 3(rarely 4), + free from the base, sulcate adaxially, horizontally spreading at first but becoming erect or ascending with age, entire, the distal portion expanded and dorsi-ventrally flattened into a large stigmatic zone.
Henderson, Neoroepera 619
io, 2, Neoroepera banksii: A. apical portion of stem from male plant showing axillary flower clusters xX 2. B. le flower x 6. C. apical aoean of ‘stent with long narrow leaves, from female piant x 2. D. female flower from side showing solitary pedicel, perianth lobes and bracts X 4. E. ovary from above showing styles and distally flattened, broad, stigmatic portions X 8. F. sub-mature fruit viewed from the side showing pedicel and persistent styles and perianth X 2. G. apex of stem with short broad leaves x 2. A,B, Gittins 1833; C-E, Ross [AQ473929]; F, Scarth-Johnson 1271A; G, Isbell [AQ204124]. All BRI.
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620 Austrobaileya 3(4): 1992
Fruit capsular, separating septicidally into three 2-valved cocci. Seeds somewhat curved, becoming dorsi-ventrally flattened at maturity, smooth or minutely pitted, carunculate, pale to dark brown when mature (and containing embryo); caruncle whitish to reddish, waxy-fleshy; endosperm copious; cotyledons several times broader than the radicle.
A genus of 2 species endemic in tropical eastern Australia. Key to species of Neoroepera
1. Plants dioecious; leaves broadest above the middle; perianth lobes of male flowers virtually smooth on margins; stamen filaments spreading hairy to above the middle; stigma limb much wider than long; far north Queensland ...... ae 1. N. banksii
Plants monoecious: leaves broadest at or ec below the niiddle perianth lobes of male flowers ciliate on margins; stamen filaments glabrous, smooth or sparsely papillose peta nee limb + longer than wide; central Queensland .... , oe ete te ue... .. 2, .N. buxifolia
1, Neoroepera banksii Benth., Flora Australiensis 6: 117 (1873). Type: Queensland, sandy ridges, north shore, Endeavour River, A.Cunningham (holo: ?K. n.v.; iso: MEL).
Dicecious shrubs (0.15~)0.5—2 m high. Stems smooth, rounded, shortly spreading-hairy when young, later glabrescent. Leaves evenly spaced along stems and branches, spreading; petiole 1.0-1.2 mm long, shortly curved hairy adaxially and abaxially; blade oblanceolate to very narrowly obovate or + spathulate, broadly obtuse or emarginate at tip and shortly attenuate to base, 6-16 mm long, 1.0-6.8 mm wide, smooth except for raised nervation, and glabrous except for short curved hairs proximally on midrib and around recurving ‘tip above, smooth and glabrous below; midrib produced as a short, recurving, usually reddish subula from the emarginate tip: margins entire, a little thickened and recurving. Stipules dark red at least distally, narrowly triangular with tip acute, 0.3-1.0 mm long and to c. 0.3 mm wide; margins glabrous to densely hairy. Flowers single or in few-flowered clusters, subtended by numerous bracts similar to but smaller than stipules. Male flowers solitary or in pairs or threes, shortly pedicellate, 6-8-merous; pedicels 6.0-9.5 mm long in flower to c. 18.0 mm long in fruit. Perianth spreading: lobes ovate to obovate with tips rounded and entire or erose, and margins entire or few toothed, sepal-like ones 0.8-1.3 mm long and 0.6-0.8 mm wide, somewhat concave, the petal-like ones 2.0-2.3 mm long and 1.7-1.9 mm wide, dished; glands of the outer whorl to c. 0.15 mm long. Stamens 6-8; filaments stout, 1.9-2.4 mm long, spreading long hairy in the lower three quarters; anthers 0.85-1.00 mm long; glandular disc c. 0.9 mm across and 0.3 mm high; pistilode rarely present. Female flowers solitary, pedicellate; pedicels 9-15(-25) mm long in flower, to c. 30 mm long in fruit. Perianth spreading, persistent and reflexed beneath fruit; lobes semi-elliptic to oblong to obovate, with margins entire, the sepal-like ones 1.4—2. 4 mm long and 0.8-1.3 mm across and rounded or acute at the apex, smooth or ciliate on margins, the petal-like ones 2.5-3.4 mm long and 1.2- 1.6 mm across, and cucullate acute at the apex, smooth or ciliate distally on margins: glands of the inner disc forming a continuous, flattened, 3-lobed ring at base of the ovary, the lobes + triangular and to c. 0.7 mm long. Ovary ovoid, c. 1.5 mm high and 1.75 mm across; styles stout, with column 0.5-0.7 mm long, and flattened limb + reniform in outline, c. 1.6-2.2 mm across. Fruit ovoid, to c. 6.5 mm long, at first conspicuously crowned with the 3 (or 4) long-persisting styles, reddish green when mature. Seed + obloid, a little tumid proximally (around hilum), 4. 0-4.7 X 2.2-2.7 X 2.2-2.7mm, longitudinally striate with lines of minute fovea, later smooth; caruncle irregularly shaped and + coralloid (dried state). Figs 1, 2.
Selected specimens (63 Peri Queensland. Cook DISTRICT: 5 km NE of Bamaga airstrip, Aug 1978, Paijnians 3020 (2) (BRI,CANB); 11°35’/S, 142°27’E, vicinity of McDonnell, Jul 1970, Isbell [AQ204125] (8) (BRD: ditto, Isbell i4 (sterile) (BRD, 11° 36'S, 142°46'E, between Heathlands homestead and Captain Billy beach, May 1980, Morton 638 (2) (BRI,MEL); 12°07'S, 143°0S’E, Olive River, near mouth, Sep 1974, Tracey 14494 @) (BRD): c. 34 km from ‘Bromley’ along road to Bolt Head, 11 km from Olive River crossing. turnoff, Jul 1990, Ross [AQ473929] (2) (BRI); 14°08’S, 143°21’E, c. 48 miles (77 km] N of Musgrave Telegraph Station, Gittins 1833 ($) (BRI) (6 & 2) (NSW); Lizard Island, in 1871, Waiter (MEL); ditto, May 1975, Byrnes 3146 a(@) & b(d) (BRI); ditto, Jul 1990, Batianoff 12186 (8) (BRD: Hopevale, Jul 1977, ‘Scarth-Johnson 545A (2) (BRI: 15°17’S, 145°19’E, 3 km SW of South Cape Bedford, Aug 1978, Kanis 1928 (2) (BRI,CANB); Cooktown, mouth of Endeavour River, Jun-Aug
Henderson, Neoroepera 621
Fig. 3. Neoroepera buxifolia: A. apical portion of a flowering branch X 0.33. B. portion of a twig showing clusters of male and female flowers x 2. C. partial cluster of flowers showing central female and surrounding male flowers — note 5- and 6-merous male flowers < 4. D. ovary from above showing styles with distal, flattened, stigmatic portions x8. = portion of a stem showing sub-mature fruit with persistent styles X 4. A-E, Batianoff MC9108001 & Robins, BRI.
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622 Austrobaileya 3(4): 1992
1770, Banks & Solander [AQ450766/MEL 515923} (@) gris ,MEL); ditto, May 1970, Blake 23311 (2 & &) (BRL MEL); 15°34’S, 147°34’E, approximately | km S Annan River mouth, Aug 1974, Tracey 14734 (8) (BRILMEL,QRS).
Distribution and habitat: Confined to far north-eastern Queensland north of about Cooktown (Map 1). Commonly occurs in sandy soils on or close to the coast, in dune communities or Eucalyptus forest on the landward side of coastal sand-dunes.
Notes: The variability in leaf shape suggests that at least two infraspecific taxa can be recognised so distinctive are the extremes. The most northerly material generally has remarkably small, narrow leaves, e.g. Isbell [AQ204125], whereas that from around Cooktown mostly has comparatively longer, quite broad leaves, e.g. Kanis 1928. However, since small-leaved forms can also occur near Cooktown, e.g. ‘Scarth-Johnson S45A, and broader leaved ones near Cape York, e.g. Isbe// 14 (Fig. 2G), and forms that cannot be grouped with either the above with certainty, e.g. Tracey 14494, occur throughout the species’ range, no attempt has been made to formally recognise this variability. The MEL isotype has leaves within the range of small/narrow to large/broad somewhat closer to the large than small end of the range. There seems no qualitative differences between leaves of the different forms.
| Notes accompanying Batianoff 12186 indicate this species has horticultural poten- tial as it is in cultivation on Lizard Island where it is described as a spectacular ornamental because of its striking red mature fruits.
Risk coding: This species is evenly distributed throughout its range and can be classed as common. It is conserved in at least the Lizard Island National Park and is not at risk,
2. Neoroepera buxifolia Muell. Arg. & F. Muell. in DC., Prodromus 15 (2): 489 (August 1866). Type: In New Holland at ‘Prenchestic’ [= Princhester] Creek [Queensland], Bowman (lecto chosen here: G-DC n.y. [BRI-microfiche IDC 800-74. 2508: I, 2], twig bearing male and (?)female flowers; 1solecto: MEL).
Securinega muelleriana Baillon, Adansonia 6: 333 (September 1866), nom. illeg. Based on Neoroepera buxifolia Muell. Arg. & F. Muell. |
Monoecious shrubs or small trees to c. 6 m high. Stems smooth, at first shortly antrorsely ferruginous pubescent later glabrescent, rounded, robust, many-branched with branches spreading or ascending. Leaves evenly spaced along stems; petiole 1.0-2.0 mm long; blade narrowly to broadly ovate to elliptic (or occasionally orbicular), with margins slightly thickened and a little recurving, tapered to broadly obtuse and a little emarginate tip and also to base, 10.0-40.0 mm long, 6.5-20.0 mm wide, smooth above and below; midrib produced as a spreading, microscopic subula from the emarginate tip; margins entire, smooth. Stipules subulate, to c. 0.3 mm long, glabrous, reddish, caducous. Flowers clustered along short axis, subtended by numerous microscopic, semi-circular to trian- gular, externally appressed-hairy bracts. Male flowers several in each cluster, pedicellate, (4, 5 or) 6 (or 7)-merous; pedicels 4.5-8.0 mm long. Perianth + crateriform: lobes ovate to obovate with tips rounded and entire, emarginate or erose, and margins regularly ciliate, sepal-like ones 0.75-1.8 mm long and 0.65-1.2 mm wide. somewhat cupular, the petal-like ones 2.2—3.2 mm long and 1.5-2.5 mm wide, dished; disc of many, spreading filiform, entire or bifid or secondarily lobed lobes up to 1.5 mm long. Stamens (4, 5 or) 6 (or 7); filaments straight, glabrous, smooth or sparsely papillate, 1.7-3.6 mm long, incipiently bifid distally; anthers ellipsoidal, 0.7-1.2 mm long. Female flowers solitary, apical on a short axis with several male flowers below it, pedicellate; pedicels to c. 8.5 mm long in flower, to c. 18 mm long in fruit, stouter than that of males, shortly antrorsely hairy throughout but denser distally, glabrescent, bracteolate near or below middle; bracteoles + ovate, to c. 0.5 mm long. Perianth + crateriform, persistent and somewhat reflexed beneath fruit; lobes narrowly ovate to oblong, rounded at the apex, and with margins shortly ciliate, the sepal-like ones 1.2-1.5 mm long, 0.75—1.1 mm across, the petal-like ones 2.5-2.8 mm long and 1.7-1.9 mm across. Ovary + ovoid, to c. 1.5mm long; styles stout, adnate only near base, ascending, with column c. 0.5 mm long, and flattened limb + narrowly ovate in outline, c. 1 mm long, its margins revolute. Fruit obloid to obovoid, 5-8 mm long, at first conspicuously crowned with the 3 long- persisting styles, olive green (?) at maturity. Seed + obloid to ovoid becoming dorsiven-
Henderson, Neoroepera 623
trally flattened with maturity, 4.2-5.1 x 2.1-3.3 x 1.7-2.3 mm, smooth but with contiguous minute fovea visible below surface of testa; caruncle reduced to a small, red-
coloured flap or cone-shaped outgrowth of tissue from testa that abutted the hook-like placenta. Figs 1, 3.
Specimens examined: Queensland. PoRT CURTIS DISTRICT: between Marlborough and Yaamba, Oct 1937, White 12095 (BRI); Livingstone Shire, about 11 km S of Marlborough homestead, at Marlborough Creek crossing, Nov 1981, Anderson 2361 (BRI); 22°58’S, 149°52’E, Marlborough Creek Crossing, Mar 1989, Reeves 630 (BRI); ditto, May 1991, Batianoff & Franks (BRI); 23°04’S, 150°15’E, gut W of Canoona, c. 45 km NW of Rockhampton, on road to Mona Vale, Nov 1990. Henderson H3490 & Robins (BRIL.K,MEL,NSW)
Distribution and habitat: Known only from a few creek-side localities north-west of
Rockhampton in the Port Curtis pastoral district, in areas of serpentinite soils (Map 1).
Notes: As explained above, the sheet of type material at G-DC, as well as that at MEL, contains material of Neoroepera buxifolia collected on two different occasions. Thus each of the two specimens on the G-DC sheet is a syntype while those at MEL are isosyntypes. To be sure of the application of the name, Bowman’s flowering specimen at G-DC, as opposed to the fruiting material there, is selected as its lectotype.
The duplicate material distributed to K was cited by Bentham (1873). That loaned to Baillon in Paris was obviously examined and reported on to Mueller prior to Baillon’s publication of 1866, for annotation on the MEL sheet reads ‘Securinega muellerii Baill’. That name was not published but Securinega muelleriana was. However, as it was proposed as a later alternative for Neoroepera buxifolia it is not a legitimate name.
This species appears restricted to the bed and banks of creeks in areas of outcropping ultramafic (serpentine) rock which itself is, in central Queensland, restricted largely to the Yaamba to Marlborough area, with a second, though somewhat smaller, occurrence between Rockhampton and Yeppoon. Thus the species could be expected to be found at other creek-side situations where serpentinite soils occur in this area.
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14 Na Na PA UNA mnt KG MEAN MNGIE OE Oa
624 : Austrobaileya 3(4): 1992
Risk coding: Though this plant is quite common in the riparian habitat it is confined to, It is restricted in occurrence at any one site. None of the known sites of occurrence is within any declared conservation reserve though the plant is known to occur within one State Forest. Most sites are subject to roadside clearing and/or grazing, and none of them is more than 50 km from the others. The species must, therefore, be considered vulnerable. A risk coding of 2V, as recorded by Briggs and Leigh (1988), is thus still appropriate.
Acknowledgements
The Directors of MEL and NSW are thanked for loan of their holdings of Neoroepera (including types from MEL) to BRI for study. I am grateful to Will Smith for producing the illustrations and maps, and to Andrew Franks and Hans Dillewaard for the SEM photomicrographs. George Batianoff, Paul Robins and Andrew Franks collected pickled material of N. buxifolia for me for illustration purposes. Support by grants from the Australian Biological Resources Study (ABRS) for my ‘Stenolobeae’ studies since 1988 is gratefully acknowledged.
References BAILLON, H.E. (1858). Etude générale du groupe des Euphorbiacées p. 561. Paris: Victor Masson. BAILLON, H.E. (1866). Species Guohorbineesram Euphorbiacées Australiennes. Adansonia 6: 282-345. BAINES, J.A. (1981). Australian Plant Genera. Society for Growing Australian Plants. BENTHAM, G. (1873). Neoroepera. Flora Australiensis 6: 116-117. London: L. Reeve.
BENTHAM, G. (1883). Euphorbiaceae. In G. Bentham & J.D. Hooker, Genera Plantarum 3: 239-340. London: L, Reeve.
BRIGGS, J.D. & LEIGH, J.H. (1988), Rare or Threatened Australian Plants. 1988 Revised Edition. Australian ae Parks and Wildlife Service Special Publication [14]. Canberra: Australian National Parks and Wildlife Service.
GREUTER, W. ET AL. (1988). International Code of Botanical Nomenclature. Articles 62.1, 63.1 & 63.2. Regnum Vegetabile 119: 63 & 64.
HURUSAWA, I. (1954). Eine nochmalige Durchsicht des herkommlichen Systems der Euphorbiaceen im weiteren Sinne. Journal of the Faculty of Science University of Tokyo Section HI Botany 6: 209-344.
HUTCHINSON, J. (1969). Tribalism in the family Euphorbiaceae. Atmerican Journal of Botany 56(7): 738-758. MEEUSE, A.D.J. (1990). The Euphorbiaceae auct. p/ur., an Unnatural Taxon. Delft: Eburon.
MUELLER, J. (1865). Euphorbiaceae. Vorlaufige Mittleilungen aus dem fiir De Candolle’s Prodromus best immten manuscript uber diese Familie. Linnaea 34: 64.
MUELLER, J. (1866). Euphorbiaceae. In A.L.P.P. de Candolle (ed.), Prodromus Systematis Naturalis Regni Vegetabilis 15(2): 189-1260. Paris: Masson.
PAX, F. . pee ec, K. (1931). Euphorbiaceae. In A. Engler & K. Prantl (eds), Die Natiirlichen Pflanzenfamilien Band i9c: 11-233.
STAFLEU, F.A. & COWAN, R.S. (1976). Taxonomic Literature edn 2, 1{A-G): 97, 447. Regnum Vegetable 94. WEBSTER, G.L. (1975). Conspectus of a new classification of the Euphorbiaceae. Taxon 24: 593-601. WEBSTER, G.L. (1984). A revision of Flueggea (Euphorbiaceae). Allertonia 3(4). 259-312.
WEBSTER, G.L. (1987). The saga of the spurges: a review of the classification and relationships in the Euphorbiales.
Botanical Journal of the Linnean Society 94: 3-46.
Appendix 1]
It is reasonable to assume from Stafleu and Cowan (1976) that Baillon’s publication dealing with Neoroepera in Adansonia appeared in July 1866, before that of Mueller Argoviensis. However, Dr Grady Webster (pers. comm.) has pointed out that since Baillon cited actual page numbers in his reference to de Candolle’s Prodromus under Securinega muelleriana, and considering the personal relationship between Mueller and himself, it is likely his publication appeared after Mueller’s (ate August 1866).
Henderson, Neoroepera 625
Despite what may be inferred from the title page and from Stafleu and Cowan, it seems clear that for Volume 6 of ‘Adansonia’, the publication schedule slipped beyond August 1866. Though parts comprised of 32 pages may have appeared monthly in early volumes, it is clear that the journal was produced in fascicles of 16 pages which were numbered consecutively in each volume in the bottom right-hand corner of the first page of that fascicle, and did not necessarily appear two per month. In Volume 6, though fascicles | to 11 are undated, fascicles 12 to 24 carry a date (presumably of publication) in the bottom left-hand corner as follows.
Fascicle Pages Date Fascicle Pages Date
12 177-192 07 Oct 1865 19 289-304 30 Aug 1866 13 193-208 11 Mar 1866 20 305-320 Sep 1866 14 209-224 li Mar 1866 21 321-336 sep 1866 15 229-240 18 May 1866 22 337-352 Sep 1866 16 241-256 12 Jun 1866 23 353-368 Oct 1866 17 297-272 12 Jul 1866 24 369-384 Oct 1866 18 273-288 30 Jul 1866
Thus fascicles 18 to 22 of this volume, covering Baillon’s paper on Australian euphorbs, apparently appeared over two months with the critical fascicle (number 21) appearing in (probably) mid to late September 1866, indeed later than Mueller’s. Incidentally, though Volume 7 fascicle 1 is undated, Volume 7 fascicle 2 is also dated September 1866.
Accepted for publication 17 March 1992
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Austrobaileya 3(4): 627-641 (1992) 627
TAXONOMIC STUDIES ON THE GENUS HOYA R (ASCLEPIADACEAE) IN PAPUASIA, 1-5
Paul I. Forster Queensland Herbarium, Meiers Road, Indooroopilly, Qld 4068, Australia and David J. Liddle P.O. Box 794, Mareeba, Qid 4880, Australia
Summary
i. The group of species of Hoya, known colloquially as ‘New Guinea Whites’, is revised. This group includes H. albiflora (Zipp. ex Blume) Boerl., H. magnifica P. Forster & Liddle sp. nov., H. naumanii Schitr., H. australis subsp. tenuipes (K, Hill) P. Forster & Liddle and H. calycina Schitr. Two subspecies are recognised for H. calycina with subsp. glabrifolia P. Forster & Liddle newly described. 2. H. pottsit Traill is lectotypified and H. nicholsoniae F.Muell., H. hellwigiana Warb. and H. sogerensis S. Moore are placed in synonymy. 3. H. anulata Schltr, is found to be the earliest name for the taxon described as H. schlechteriana S. Moore, H. poolei C.White & Francis, H. pseudolittoralis C., Norman or H. alata K. Hill. 4. H. reveluta Wight ex J.D. Heok. is found to be the earliest name for the taxon described as H. litoralis Schitr., H. inconspicua Hemsley or H. dodecatheiflora Fosb. 5. H. sussuela (Roxb.) Merr. is the earliest name for a widespread taxon to which the name A. lauterbachit Schumann oro abd ae misapplied. H. hAollrungii Warb. and H. coronaria var. papuana Bailey are placed in synonymy of Hi. sussuela,
Introduction
Approximately eighty published names have been applied to species of Hoya R. Br. in Papuasia (rian Jaya, Papua New Guinea, Solomon Islands). An account of the genus in German New Guinea (now part of Papua New Guinea) was provided by Schlechter (1913) who utilised mainly his own field collections. There is no overall account of the genus in Papuasia and many species have been subsequently described without reference to previously described taxa (e.g. Moore 1916).
Despite this early taxonomic activity, there is considerable confusion as to the identity of the taxa present in the region. A large number of unidentified collections have accumulated in herbaria, mainly as a result of the collecting activities of officers of the Department of Forests, Lae (NGF and LAE series) and the Archbold expeditions. The primary set of the NGF-LAE collections 1s at the National Herbartum of Papua New Guinea (LAE), with the second set at the Rijksherbarium, Leiden (L), the third set at the Queensland Herbarium (BRI), and minor sets at other herbaria such as the Australian National Herbarium (CANB). The collections of L.J. Brass (Archbold expe- ditions) are distributed in several herbaria with primary sets mainly at the Arnold Arboretum, Harvard (A), New York Botanic Gardens (NY) and L, with minor sets of some numbers at BM and BRI.
While there are many collections of Hoya available, few have been identified fully. This is undoubtedly a result of the lack of a comprehensive revision and key to the species. Also, it has been assumed that the types for many of the species described by Schlechter were destroyed in the firebombing of the Berlin Herbarium (B) in 1942. However, this is not the case and most are still extant in B (B. Leuenberger, pers. comm. 1988). Unfortunately, the same cannot be said for the species named by Engler, Schumann or Warburg. Very few duplicates of type collections relevant to taxa named by these workers have been located. Although the large number of published names does not give an accurate picture of the actual number of species of Hoya present in Papuasia, the region, nevertheless, does have a rich diversity of taxa comprising approximately 70 species.
From examination of Papuasian holdings at the herbaria A, BO, BM, BISH, BRI, CANB, L, LAE, MEL, NY and SING, and relevant type material at K, P and WRSL, it is evident that some of the taxa collected and named by Schlechter, often from single collections, have not been recollected in more recent times. Conversely there are
628 Austrobaileya 3(4): 1992
collections, particularly from Inan Jaya and southern Papua New Guinea that represent undescribed taxa not seen by him.
A comprehensive revision of the genus in this region will require extensive field work and cultivation of field-collected material which will take many years. In the interim we believe we have been able to match a significant proportion of existing herbarium collections with the types from various taxa. These identifications have represented significant changes to known distribution records of the taxa concerned in the Papuasian region and beyond. We believe it to be of some importance to progressively publish these findings, if only to stimulate curation in other herbaria, and to encourage further fieldwork and study. In the long-term we hope to produce a revision of the genus in the region.
At this preliminary stage of our investigations, some comment on nomenclatural priority is necessary. We have previously stated (Forster & Liddle 1990) that for certain widespread taxa, it is possible that earlier legitimate names exist than the ones we previously accepted. Such has proved to be the case with H. revoluta Wight ex J.D. Hook., AH. anulata Schlitr., H. sussuela (Roxb.) Merr. and H. pottsii Traill. which have been found to be earlier names for species previously known by more familiar names.
Notes on Terminology
Indumentum cover is described using the terminology of Hewson (1988), except that ‘scattered’ is used instead of ‘isolated’. In previous papers we have described the glandular structures at the base of the leaf lamina as ‘extrafloral nectaries’ and those at the sepal sinus bases as ‘glands’. However, following the excellent review of the mor- phology and evolution of the corona and related structures in Asclepiadaceae and allied families by Kunze (1990), we are now calling both these structures ‘colleters’.
1. Revision of the ‘New Guinea Whites’
In Hoya horticulture, there is a group of taxa originating from Papuasia that have been known colloquially as the ‘New Guinea Whites’. These taxa have large, showy, usually totally white flowers. Naming of cultivated material of these plants has been hopelessly confused and generally based on ill-informed conjecture of horticulturalists attempting to match original descriptions to live plants without recourse to the types. Much of this confusion has centred on the definition of H. australis R. Br. ex Traill, (see Forster & Liddle 1991), Hoya albiflora (Zipp. ex Blume) Boerl. and H. calycina Schltr. (e.g. Burton 1989). The existence of two undescribed taxa in the group has also added to the confusion.
The present study is based both on live and herbarium specimens. Key to taxa of ‘New Guinea Whites’ Hoya
1. Flower campanulate; corolla not red under coronal lobes; corolla lobes
triangular ...... ee 2 Flower akapaniilatesotate: cored ved ander doteiial Bbes! eaveia lobes IAHCCOTESOVATE: 0. One ley beet veins Sewn «Gye e toe eee oveg ter kh 3
2. Corolla 40-45 mm diameter; lobes horizontal or reflexed, 15-18 mm long ...... Br H. magnifica
Corolla 20-25 mm deartietor’ Idlies iriflexed: Aly mm viows ...... H. albiflora
3. Staminal column elongated with staminal corona raised well above corolla ....... a again H. naumanii
Staminal column not Siosinled: rae af veucettial ¢ corona Siaated more or less flush with corolla .. 2... 0... ee eae 4
Forster & Liddle, Hoya in Papuasia, 1-5 629
4. Corolla internally with dense puberulous indumentum of short white
hairs; staminal corona lobes 1.2-3.5 mm long ........ .. H. australis Corolla internally glabrous or with only scattered puberulous indumentum of short white hairs; staminal corona lobes 3.4-4.8 mm long .. H. calycina
Note: H. naumanii Schltr. 1s restricted to Bougainville and Solomon Islands and will be dealt with in detail in a later publication when recently collected material (June 1991) has been grown on and flowered. H. australis subsp. tenuipes (K. Hill) P. Forster & Liddle, the only subspecies of H. australis present in Papuasia, has been covered previously (Forster & Liddle 1991).
1. Hoya magnifica P. Forster & Liddle, sp. nov. a H. albiflora (Zipp. ex Blume) Boerl. corolla 40-45 mm diametro eius lobis 15-18 mm longis reflexis vel horizontaliter portatis differt. Typus: Papua New Guinea. MOROBE PROVINCE: Palenkwa, Golden Pines, 7°06’S, 146°35’E, 24 April 1969, H. Strelmann & A. Kaitro NGF39381 (holo: CANBI!; iso: A!, BO!, BRI, L!, LAE!; K 7...)
Liane to several metres long, latex white. Stems cylindrical, with dense indumentum when young, becoming sparse with age; internodes up to 270 mm long and 5 mm diameter. Leaves petiolate, held erect or at angle of 10-20° from vertical; lamina elliptic- ovate to narrow-ovate, up to 190 mm long and 100 mm wide; upper surface with sparse indumentum, venation obscure; lower surface with dense indumentum, venation obscure; tip acute, shortly acuminate to somewhat apiculate; base cordate to rounded; petiole 14- 43 mm long, 2.3-2.5 mm diameter, with dense indumentum; colleters 4 at lamina base. Cymes racemiform, up to 90 mm long, held horizontally to pendulous; peduncles 9~10 mm long, 5-6 mm diameter, with dense indumentum; bracts triangular, 1.4-1.5 mm long, 1.2-1.3 mm wide, with sparse to dense indumentum. Flowers 10-15 mm long, 40-45 mm diameter; pedicels 20-50 mm long, 1.1-2.6 mm diameter, with dense indumentum, Sepals ‘lanceolate- ovate, 13-16 mm long, 7-10 mm wide, with dense indumentum externally; colleters 5 large and 5 small (minute) at base of each SINUS, each large colleter subulate, 2-3 mm long, 0.7-0.8 mm diameter, with dense indumentum. Corolla campanulate, cream to white; tube 10-14 mm long, 22-25 mm diameter, glabrous; lobes triangular, held horizontally or reflexed, 15-18 mm long, 15-18 mm wide, glabrous. Staminal corona cream to white, 4-5 mm long, 10-16 mm diameter; each lobe 4.8-7.0 mm long, 4.0-4.5 mm high, 2. 84. Q mm wide at base and c. 0.5 mm wide at upper tip, outer edge rounded and upturned, inner tip lanceolate, not extending beyond style-head. Staminal column 4-7 mm long, 4.6-5.0 mm diameter: anther appendages lanceolate, 1.7-2.0 mm long, 1.3-1.7 mm wide; alar fissure 2.0-2.5 mm long. Style-head depressed- globose, 6.5-7.0 mm long, 4.5-5. QO mm diameter. Ovaries c. 4 mm long and 2 mm diameter, with sparse indumentum. Pollinartum 1.3-1.4 mm long, 1.2-1.3 mm wide; pollinia oblong, 1.12-1.15 mm long, 0.35-0.42 mm wide, with pellucid germination mouth on outer edge; corpusculum ovate, 0.8-0.9 mm long, 0,55-0.58 mm wide; caudicles 0.30-0.35 mm long, 0.13-0.14 mm wide, not winged. Fruit fusiform, c. 190 mm long and 12 mm diameter, glabrous. Seed not seen. Fig. 1.
Specimens examined. Papua New Guinea, MOROBE PROVINCE: Moikisung — Manga track, c. 2 km above Moikisung, Huon Peninsula, 6°37’S, 147°38’E, Sep 1976, Clunie et al. LAE63319 (BRI,L,LAE); Mapos, Snake River, 6°55’S, 146°45’E, Mar 1971, Streimann & Kairo NGF25879 (BRLCANB,L,LAE); Bulolo - Watut Divide, Rd 46, 9 km SW of Bulolo, 7: 15S, 146°35’E, Dec 1980, Kairo 691 (A,L; K a.y.); Wantoat, Feb 1940, Clemens 11126 (A, BISH,BRI L): Patep, Mumeng, Mar 1979, Rau 482 (LAE). Cultivated. Lae Botanic Gardens (origin not stated), 1957, Millar NGF9213 (A, BRI); Arawa Plantation, Bougainville (origin not stated), Apr 1970, Millar & Vandenberg NGF48503 (BRI,LAE); Emerald Creek, Mareeba (ex Lukins Rubber Estate near Sogeri, Central Province, USDA354244), Apr 1990, Liddle IML76 (BRI).
Distribution and habitat: Known with certainty only from the Morobe and Central Districts of Papua New Guinea (Map 1). Plants grow as hanes in lowland rainforests at altitudes of 650 -— 2000 m.
Notes: This species has been confused with H. albiflora (Zipp. ex Blume) Boerl. in cultivation. It is probably closely related to H. albiflora but is immediately distinguishable by the much larger flowers of which the corolla lobes are held horizontally or reflexed at anthesis, as opposed to the smaller flowers with corolla lobes inflexed at anthesis in the latter species.
Etymology: The specific epithet alludes to the magnificent white flowers of this species.
. atta ade ae MA BEM CC 8 tm a aa A GOAN A tat tach kn teetat cama DEAT Ott dd ttt td fet tatoo atte ede ais ie gy eee a nt an sti geen . . i RE i ea RE ty Ne eee eee eRe eee Ry En Ee PLB DE CTA oP ae TG TL LE FE de acd BO aa PgR eI nS Sate ARE en A PUN eee RENE «
630 : Austrobaileya 3(4); 1992
Fig. 1. Hoya magnifica: A. habit of flowering stem X 0.5. B. side view of flower X 0.5. C. face view of flower X 0.5. D. side view of staminal column and corona X 2.5. E. face view of staminal column and corona X 2.5. F.,
face view of calyx X 2.5. G. verticai cross-section of flower X 2.5. H. pollinarium (inverted) X 16. All from Liddle IML76. Del. D.J. Liddle.
Forster & Liddle, Hoya in Papuasia, 1-5 631
2. Hoya albiflora (Zipp. ex Blume) Boerl., Handi. Fl. Ned.-Ind. 2(2): 440 (1899); Pterostelma albiflora Zipp. ex Blume, Rumphia 4: 33, t. 188 (1849). Type: Nov. | ile [New Guinea], Zipp. [Zippelius| (holo: L! [898168-96, 898168-95, 898168-
— 94)).
Blume, Fl. Ned. Bat. 1: 513 (1856).
Liane to several metres long, latex white. Stems cylindrical, up to 6 mm diameter, with Sparse indumentum when young, becoming sparse to glabrous with age; internodes up to 180 mm long. leaves petiolate; lamina elliptic, up to 180 mm long and 90 mm wide: upper surface glabrous, venation obscure; lower surface glabrous, secondary venation of 6-7 veins per side of midrib nearly obscure, tertiary venation obscure; tip acute to shortly acuminate; base cuneate, truncate or cordate; petiole 18-30 mm long, 1.4—-3.0 mm wide, glabrous or with scattered indumentum; colleters 4 at lamina base. Cymes racemiform, up to 70 mm long, held pendulously; peduncles 4-6 mm long, 2-3 mm diameter, with scattered to dense indumentum; bracts triangular to lanceolate-ovate, 0.8— 1.6 mm long, 0.6-1.0 mm wide, with scattered to dense indumentum. Flowers 15-18 mm long, 20-25 mm diameter; pedicels 25-50 mm long, 1.0-1.6 mm diameter, with scattered to dense indumentum. Sepals lanceolate-ovate, 13-15 mm long, 6-7 mm wide, with sparse to dense indumentum externally; colleters 5 at base of each sinus, each gland -subulate, 3-7 mm long, c. 1 mm diameter, with sparse indumentum. Corolla campanulate, cream to white; tube 8-10 mm long, 15-17 mm diameter, glabrous; lobes triangular, inflexed, 7-!1 mm long, 5.5-12.0 mm wide, glabrous. Staminal corona cream to white, 4-5 mm long, 7-8 mm diameter; lobes 3-4 mm long, 4.5-5.0 mm high, 2.7-2.8 mm wide at outer edge, c. 0.5 mm at inner tip, outer edge upturned and sharply rounded, inner tip subulate and reflexed from style-head. Staminal column c. 4 mm long, 5.0-5.4 mm diameter; anther appendages lanceolate, obscuring style-head, 1.5-1.8 mm long, 0.9- 1.4 mm wide; alar fissure 1.5-2.5 mm long. Style-head depressed-globose, 1.8-2.0 mm diameter. Ovaries 2.6-3.0 mm long, 1.9-2.0 mm diameter, glabrous. Pollinarium 1.15- 1.20 mm long, 1.0-1.1 mm wide; pollinia oblong, 1.15-1.20 mm long, 0.44-0.45 mm wide, with pellucid germination mouth on outer edge; corpusculum ovate, 0.80-0.85 mm long, 0.32-0.50 mm wide; caudicles unwinged, 0.35-0.37 mm long, 0.12-0.13 mm wide. Fruits and seeds not seen. Fig. 2.
Specimens examined, Papua New Guinea. MOROBE PROVINCE: Patep III, 6°20’S, 146°45’E, Aug 1969, Adillar Eons fa (BRLL,CAE; K n.yv.). Cultivated. Emerald Creek, Mareeba (from unknown origin), Apr 1990, Liddle IML299 (BRI). |
Distribution and habitat: H. a/biflora is known only from the Morobe Province in Papua New Guinea (Map 2) where it grows as a liane in rainforest.
Notes: There are few collections of H. albiflora, either in herbaria or in cultivation. While there are several differently numbered clones of H. albiflora in cultivation, none have collection data, and all are probably from the same original plant.
3. Hoya calycina Schltr., Bot. Jahrb. Syst. 50: 125 (1913). Type: Papua New Guinea. MT BRI PROVINCE: Kani Geb., Mar 1908, R. Schlechter 17510 (holo: B (photo at BRI)).
[Hova albiflora auct., non (Zipp. ex Blume) Boerl.: Burton, Hoyan 11(2): 57-58 (1989)].
Liane to several metres long, latex white. Stems cylindrical, up to 6 mm diameter, glabrous or with scattered to dense indumentum when young, becoming corky with age; internodes up to 160 mm long. Leaves petiolate; lamina elliptic to elliptic-ovate, up to 170 mm long and 90 mm wide; upper surface glabrous or with scattered indumentum, venation obscure; lower surface glabrous or with scattered to dense velutinous indu- mentum, venation obscure; tip acute, acuminate, or apiculate: base rounded, cuneate, or slightly cordate; petiole 12-30 mm long, 1.5-—3.5 mm diameter, with scattered to dense indumentum; colleters 4 at lamina base. Cymes racemiform, up to 70 mm long; peduncles 6-25 mm long, 3~7 mm diameter, glabrous or with scattered to dense indumentum; bracts triangular to lanceolate, 0.8-1.0 mm long, 0.5-1.0 mm wide. Flowers 9-10 mm long, 18-28 mm diameter; pedicels 24-44 mm long, 1.6-2.0 mm diameter, glabrous or with scattered to dense indumentum. Sepals lanceolate to lanceolate-ovate, 2.8-5.0 mm
632 Austrobaileya 3(4): 1992
Fig, 2. Hoya albiflora: A. habit of flowering stem X 0.5. B. face view of flower X 0.5. C. side view of staminal column and corona X 3. D. face view of staminal column and corona x 3. E. face view of calyx X 0.75. F. vertical cross-section of flower X 3. G. pollinarlum (inverted) X 8. All from Liddle IML299. Del. D.J. Liddle.
Forster & Liddle, Hova in Papuasia, 1-5 633
long, 1.2-3.0 mm wide, glabrous or with scattered to dense indumentum; base of each sinus with an irregular glandular ridge. Corolla campanulate-rotate, cream to white, red to purple under coronal lobes and around gynostegium; tube 3.0-5.4 mm long, 8-14 mm diameter, glabrous; lobes lanceolate-ovate, 7-13 mm long, 4-9 mm wide, glabrous or with scattered to sparse indumentum on the edges and externally. Staminal corona 4.0-4.3 mm long, 7-13 mm diameter; lobes 3.4-4.8 mm long, 4.0-4.3 mm high, 1.8- 2.5 mm wide. Staminal column 3.6-4.0 mm long, 2.5—-3.5 mm diameter; anther appen- dages lanceolate, 1.6-—2.0 mm long, 1.7-1.8 mm wide; alar fissure 1.3—2.0 mm long. Style- head depressed-globose, 1.5-1.9 mm diameter. Ovaries 2.3-2.4 mm long, 1.8-1.9 mm wide, glabrous. Pollinarium c. 1.1 mm long, 0.85-0.90 mm wide; pollinia oblong, 1.05- 1.10 mm long, 0.35-0.40 mm wide, with pellucid germination mouth on outer edge; corpusculum ovate, 0.60-0.65 mm long, 0.35-0.40 mm wide; caudicles unwinged, 0.25- 0.30 mm long, 0.06—0.07 mm wide. Fruit and seed not seen. Fig. 3.
Notes: H. calycina is a distinctive species that has been confused with H. australis and H. albiflora, mainly because of its superficially similar flowers. It differs most markedly ‘from H. australis in the more-or-less internally glabrous corolla and the much larger staminal corona and from H. albiflora in the rotate corolla. Like H. australis, H. calycina * variable in terms of indumentum and two subspecies can be recognised on this character,
Key to subspecies of Hoya calycina
1. Lamina with dense velutinous indumentum below, calyx and corolla with
sparse to dense indumentum externally ................ subsp. calycina Lamina glabrous or with scattered indumentum below, calyx and corolla glabrous or with scattered indumentum externally ...... subsp. glabrifolia
3a. Hoya calycina subsp. calycina
Lamina with dense velutinous indumentum below. Calyx and corolla with sparse to dense indumentum externally.
Specimens examined. Papua New Guinea, NEw IRELAND: Wanup near Lossuk, 2°45’S, 151°04’E, Jan 1967, Coode et al. NGF29625 (L). MOROBE PROVINCE: S of Boana, 6°30/S, 146°50’E, Feb 1977, Conn et al. 86 (K,L); Patep Creek, 6°35’S, 146°25’E, May 1959, Millar NGF9967 (A,BRI,CANB,LAE), Zenag, 7°00’S, 146°35’E, Jul 1968, Millar NGF12115 (A,BRIL.CANB,L,LAE); Kwatmengu, Aseki Patrol Area, Apr 1966, Craven & Schodde 1459 (A,L,LAE). SOUTHERN HIGHLANDS PROVINCE: Mt Bosavi, northern side, N of the Mission Station, 6°26/S, 142°50'E, Oct 1973, Jacobs 9496 (L,LAE). NORTHERN PROVINCE: c. 2 km W of Popondetta along road near airstrip, Jui 1953, Hoogland 3379 (A,BM,BRI,CANB,K,L,LAE). MILNE BAY PROVINCE: SW of Nowata airstrip, 9°59’S, 149°44’E, Jul 1969, Kanis 1104 (CANB,LAE). Cultivated. Emerald Creek, Mareeba (ex plant collected MOoOROBE PROVINCE: Patep II Village, [USDA354236]), Apr 1990, Liddle IML201 (BRD.
Distribution and habitat: Widespread in Papua New Guinea (Map 3). Plants grow in rainforest at altitudes of 20-1400 m.
3b. Hoya calycina subsp. glabrifolia P. Forster & Liddle subsp. noy., a H. calycina Schltr. subsp. calycina lamina foliorum glabra vel infra pilis dispersis praedita, calyce corollaque glabra vel extus pilis non nisi sejunctis praedita differt. Typus: Irian Jaya. 4 km SW of Bernhard Camp, Idenburg River, Mar 1939, L.J. Brass 13465 (holo: BRI}; iso: A!, BO!, L!).
Lamina glabrous or with scattered indumentum below; calyx and corolla glabrous or with scattered indumentum externally.
Specimens examined. Papua New Guinea. WESTERN HIGHLANDS PROVINCE: Jimmi Valley, near Karap, Jun 1955, Womersley & Millar NGF7648 (BRI,LAE). EASTERN HIGHLANDS PROVINCE: Kassam, Nov 1959, Brass 32470 (LAE). MOROBE PROVINCE: Bupu Village, Wampit, 6°50/S, 146°55/E, Jul 1967, Millar NGF22936 (BRI,L,LAB). CENTRAL PROVINCE: Isuarava, 9°00’S, 147°44’E, Mar 1936, Carr 16107 (BM,CANB,L,NY,SING; EK #.1.).
Distribution and habitat: Widespread but rarely collected in Papua New Guinea (Map 4). Plants grow as lianes in rainforest at altitudes of 1500-1900 m.
634 Austrobaileya 3(4): 1992
Fig. 3. Hoya calycina subsp. calycina: A. habit of flowering stem x 0.5. B. face view of flower X 12.5. C. side view of stamina! column and corona X 3. D. face view of calyx X 3. E. vertical cross-section of flower X 3. F. pollinarium (inverted) X 28. All from Liddle IML201. Del. D.J. Liddle.
Forster & Liddle, Hoya in Papuasia, 1-5 635
2. Typification and synonymy of Hoya pottsti Traill
Hoya pottsii Traill, Trans. Hort. Soc. 7: 25 (1827). Type: based on plant in cultivation. (lecto (here designated): Traill, Trans. Hort. Soc. 7: 25. fig 1 (1827)).
Hoya nicholsoniae F. Muell., Fragm. 5: 159 (1866). [October, not specified to day], synon. noy. Type: ‘In arboribus ad sinum litoreum Rockingham’s Bay, Dallachy (holo: MEL 72.¥.).
Hoya hellwigiana Warb. in Fedde, Repert. Spec. Nov. Reg. Veg. 3: 342 (1907). synon. nov. Type: ‘Kaiser Wilhelms-Land: Bussum bei Finschhafen’, O. Warburg 21313 (holo: B n.v., destroyed).
Hoya sogerensis S. Moore, J. Bot. 52: 293 (1911). synon. nov. Type: Papua New Guinea. CENTRAL PROVINCE: River side Sogere, 1885-6, H.O. Forbes 691 (holo: BM!}).
Additional selected specimens, Celebes, Sulawesi Selatan, Soroako, S. shore of Lake Matano, de Vogel 5793 (BRI). Irian Jaya. Sorong, Roefei River N of the town, Mar 1954, van Royen 3007 (L); Mamberamo, Oct 1914, Feurlletau de Bruyn 130 (BO,L); Rouffaer River, Aug 1926, Docters vy. Leeuwen 10122 (BO,L,SING; K u.¥.); Waigeo Is, Lupintol Village on SW coast of Majalibit Bay, Feb 1955, van Royen 5483 (L); Mairipi, near Andai, SW of Manokwari, Nov 1961, Vink BW12104 (L). Papua New Guinea. EAsT SEPIK PROVINCE: Ramu fluB, Tappenbeck 37 (WRSL).
Distribution and habitat: Widely distributed in Celebes, New Guinea and Australia (Forster & Liddle 1990).
Notes: Since the publication of our account of H. nicholsoniae F. Muell. (Forster & Liddle 1990), we have been able to examine a much greater range of material, particularly from L, that is referrable to this taxon. From this it is clear that H. nicholsoniae F. Muell., AH. hellwigiana Warb. and H. sogerensis 8. Moore are all conspecific, based on the examination of type collections, original descriptions and collections from the areas where these taxa originated. However, it 1s evident that H. pottsii Traill 1s also conspecific with these taxa as suggested by Burton (1983) and hence, due to priority, its name is the correct one to be used for the aggregate taxon.
H. pottsii has been newly named in most geographic regions where it has been collected, and although there is wide variation in flower colour and to some extent leaf size and shape, which are both dependent on environment (Forster & Liddle 1990), there are no valid reasons for upholding any of the later names.
The status of H. samoensis Seem. described from Samoa, H. neocaledonica Schltr. described from New Caledonia, H. neoebudica Guill. described from Vanuatu and H. cominsii Hemsley, described from Solomon Islands, with respect to H. pottsii is unclear at this stage. We have examined a wide range of collections from these areas (holdings at BSIP, P and NOU) and recently collected (June 1991) a range of material from Solomon Islands of A. cominsii, however, further comparisons with H. pottsii from Australia and New Guinea must wait until this recent material has been grown and flowered under similar conditions.
Typification of H. pottsii is critical to the application of the name and this species was named somewhat informally by Traill (1827). There appears to be no specimen at K that could be unequivocally considered as a type for H. pottsii. However, there is a flowering specimen at K labelled ‘Hoya Pottsii Traill. Hort Glasg. bot. Mag. t. 3425’ that may possibly represent the cultivated material illustrated both by Traill and by Hooker (1835). This plant may well have persisted 1n cultivation at K for a considerable time as there is a further flowering collection of 2 sheets labelled [in part] ‘Hoya pottsi Trail native in China... EN464-63 Sir George Taylor H2855/86 .. .’. While the geographic origin of both these cultivated collections 1s problematic, it is unlikely that they came from China [Macao], the reputed origin for H. pottsii (Traill 1827; Hooker 1835), and both are conspecific with the Australian and Papuasian material. While of dubious status as types of H. pottsii, both specimens lend credence to our application of the name H. potisii to the taxa treated as conspecific in this paper. In the absence of an unequivocal type specimen for H. potisii, we have selected as lectotype, the plate published by Traill (1827) which agrees perfectly with the taxon we apphed the name to.
shone SMO SOM EM ae dh a cata
636 Austrobaileya 3(4): 1992
150°E
O 130 E
150°E
Maps 1-2. Distribution of Hoya spp. i. H. magnifica. 2. H. albiflora.
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Forster & Liddle, Hoya in Papuasia, 1-5
150°E
150°E
f
Maps 3-4. Distribution of Hoya calycina. 3. H. calycina subsp. calycina. 4. H. calycina subsp. glabrifolia.
637
FN eee RN EN Sets aR RCE
Sg seonmm costae renee vee MS ANE SOS NR ee na oe nee a it Wad tao cta gs sertt
638 Austrobaileya 3(4): 1992
No type material of H. hellwigiana Warb. has been located; however, we have found the collection Tappenbeck 37 that was cited by Schlechter (1913) in his account of this species. Given that Schlechter would have surely seen the type of H. hellwigiana, it seems reasonable to allocate this name to synonymy based on this collection and the original description.
3. Typification and synonymy of Hoya anulata Schltr.
Hoya anulata Schitr. in Schumann & Lauterb., Nachtrage Fl. Schutzgeb. Stidsee 362 (1905). Type: Papua New Guinea. MADANG PROVINCE: Auf Baéumen am oberen Nuru, auf dem Wege vom Ramu zur Kiste, 4 February 1902, R. Schlechter 14185 (holo: BI).
Schlitr., Bot. Jahrb. Syst. 50: 118 (1913).
Hoya schlechteriana 8. Moore, Trans. Linn. Soc. 9: 112 (1916), synon. nov. Type: Irian Jaya. Camps III-VI, 1912-13, Utakwa River to Mt Carstensz, C_B. Kloss (holo: BM!).
Hoya poole: C. White & Francis, Proc. Roy. Soc. Queensl. 39: 69, fig. 13 (1928), synon. noy. Type: Papua New Guinea. MADANG PROVINCE: Joangey, South eastern end of Finnisterre Range, December 1923, CLE. Lane-Poole 566 (holo: BRII!; iso: K!).
Hoya pseudolittoralis Norman, Brittonia 2: 328 (1937), synon. nov. Type: Papua New Guinea. WESTERN PROVINCE: Dagwa, Oriomo River, Feb-March 1934, L./. Brass 5990 (holo: NY n.yv.; iso: A!, BM (photo at BRI), BO!, BRI).
Hoya alata K. Hill, Telopea 3: 249 (1988), synon. noy. Type: Australia, Queensland. Cook District: Pascoe River rockpile, B. Wallace 83250 (holo: NSW, 2.y7.).
Hoya sp., Jones & Gray, Austral. Climbing PI. Fig. 126 (1977). [Hoya gracilipes aact. non Schltr.: Jones & Gray, Climbing Pl. Austral. 242 (1988)]. Forster & Liddle, Austrobaileya 3: 228-230 (1990).
Additional specimens examined. Irian Jaya. Orob River, Feb 1913, Pulle 1202 (BO,L). Papua New Guinea. WESTERN HIGHLANDS PROVINCE: Baiyer River, Nov 1954, Womiersiey & Floyd NGF6809 (LAE). MOROBE PROVINCE: 4 miles [6.7 km] NE of Boana, near Guambot, 6°30’S, 146°50’E, Mar 1977, Conn et al. 70 (CANB,K,LAE); Boana, Jul 1938, Clemens 8480 (A,B,L); ditto, May-Nov 1940, Clemens 41725 (A,BRI); Bupu Village above Wampit River, Mar 1964, Afi/lar NGF23296 (A, BRI,;CANB,L,LAE); Ekwap, Wantoat, Aug 1968, Avi/lar NGF12130 (L,LAE); Wantoat, Jun 1957, White NGF9523 (LAE); Patep, Mar 1959, Millar NGF9904 (BRI), Wareo, Jan 1936, Clemens {$21 (A); Finnisterre Range, Ekwap, near Wantoat, Aug 1968, Philipson 3305 (A,L); Zenag, 7°00’S, 146°35’E, May 1969, Avi/iar NGF40862 (LAE). GuLF PROVINCE: near Iori Village, Ihu, Apr 1971, Stone 10143 (L,LAE). Australia. Queensland, Cook District: Garraway Creek rockpiles, Apr 1988, Forster 4234 & Liddle (BRI); South Pap, Tozers Gap, Jul 1991, Forster 9079 (BRI).
Notes: Examination of the type specimen of H. anulata Schltr. has found it to be conspecific with H. pseudolittoralis Norman (Forster & Liddle 1990). H. anulata is widely distributed in New Guinea, but has only been found in the Tozer Range area of Queensland.
Of importance with respect to the synonymy for this taxon, is H. poolei C.White & Francis which was applied by Liddle (1986) to the Australian material. Both the BRI holotype and the K isotype of H. poolei lack flowers; however, the foliage is a good match for those of H. anulata (based on the range of specimens examined) and the flowers figured by White and Francis (1928) agree with those of this taxon.
Isotype specimens of Hf. alata K. Hill have yet to be examined, despite repeated requests to NSW for their distribution.
4, Typification and synonymy of Hoya reveluta Wight ex J.D. Hook.
Hoya revoluta Wight ex J.D. Hook., Fl. Brit. India 4: 55 (1883). Type: Malaysia. Malacca, LO eects 1867, A.C. Maingay 1127 (ecto: K!, fide Rintz, Malay. Nat. J. 30: 486 (1978)).
Hoya inconspicua Hemsley, Bull. Misc. Inform. Kew 1894: 213 (1894), synon. nov. Type: Solomon Islands, Officers of H.M.S. Penguin (holo: K!).
Forster & Liddle, Hoya in Papuasia, 1—5 639
Hoya litoralis Schltr. in Schumann & Lauterb., Nachtrage Fl. Schutzgeb. Stidsee 363 (1905), synon. nov. Type: Papua New Guinea. MADANG PROVINCE: Auf Baumen am Strande von Potsdam Hafen, 16 October 1901, R. Schlechter 13675 (holo: B (photo at BRI')).
Forster & Liddle, Austrobaileya 3: 228-229 (1990).
Hoya dodecatheiflora Fosb., Lioydia 3: 118 (1940), synon. nov. Type: Solomon Islands. TEMOTU PROVINCE: Santa Cruz Islands, Vanikoro, Tevia Bay, 6 May 1933, Stewart (holo: BISH n.y.).
Additional selected specimens, Malaysia. JOHORE: between G. Blumut & G. Bechua, May 1923, Holttum 10844 (K); Bukit Paloh Estate, Apr 1958, Shah & Kadim 390 (K,L)}; Kg. Hubang Development Area, 100 m. s. Endau Road, Jul 1959, Burkill 1904 (L). SELANGOR: Sg, Buloh, R.R.LE.E. Forest Reserve, Nov 1956, Burkill & Shah 1066 (K,L). Indonesia. Sumatera. Vicinity of Ack Mocute (Aer Moette} Asahan, NE of Tomoean Dolok & W of Salabat, Jul 1936, Boeea 9335 (L); Mt Sago near Pajakumbuh, Jun 1956, Meijer 5099 (L). Borneo. Central Kutei, Belajan R., near Kembang Dyangut, May 1955, Kostermans 10692 (L); West Kutei, Mt Palimasan near Tabang on Belajan River, Sep 1956, Kostermans 12751 (L). Irian Jaya. East bank of Merauke River, 8 of Senajo, Aug 1954, van Royen 4668 (A,CANB,L),; Merauke River, West bank between Djedjoerah & Eramboe, Aug 1954, van Royen 4812 (CANB,L). Papua New Guinea. WEST NEw BRITAIN: Nantambu, Feb 1971, Lelean & Stevens LAES1180 (L,LAE). WESTERN PROVINCE: c. 8 miles a km] S of Morehead Patrol Post, Trans-Fly area, Aug 1967, Paijmans 291 (CANB); Weam, Jul 1967, Ridsdale NGF33505 (BO,CANB,L,LAE; K 7.¥.). Solomon Islands. GUADALCANAL PROVINCE: Mt Austen area, Jun 1991, Forster 8608, 8612 & Liddle (BRI); NW of Tinomeat Village, Goldridge area, Jun 1991, Forster 8722 & Liddle (BRI,K); NW Guadalcanal, Mataniko’o River, Nov 1967, Nakisi et al. BSIP8246 (BSIP,K,L). MALAITA PROVINCE: Malaita, Are-are dist., Moka village, Nov 1965, Hunt 3080 (K), MAKIRA PROVINCE: San Cristobal, Kira Kira, Aug 1932, Brass 2761 (A,L). TEMOTU PROVINCE Santa Cruz Group, Vanikoro Island, Dec 1928, Kajewski (K); Luendambu area, Tomotu Noi, Apr 1972, Powell BSIP19500 (BISH,BSIP,CANB,L).
Notes: Like H. pottsii, H. revoluta has a very wide geographic distribution from Malaysia (Rintz 1978), through Malesia to New Guinea, Solomon Islands and northern Queensland. Once again, this species has been formally renamed in various geographic regions of its occurrence, e.g. H. inconspicua Hemsley (Hemsley 1894) and H. dodecatheiflora Fosb. (Fosberg 1940) from Solomon Islands and H. /itoralis Schltr. from New Guinea (Schlechter 1905). The type of H. inconspicua is notable for its very long leaves and the type of H. dodecatheiflora, as illustrated by Fosberg (1940), for a more raised style-head in relation to the anthers than is typical in H. revo/uta from Malesia. Examination of many flowering plants in Solomon Islands shows that there are plants with both long and short leaves. Hence this character 1s unreliable for species distinction. Some plants have raised style- heads whereas others did not. However, there are numerous intermediates and this character is also unreliable for distinguishing species. Hence there is no justification for recognition of these taxa at specific rank distinct from H. revoluta s. lat. However, further fieldwork and cultivation under uniform conditions of plants of known origin from throughout the range of H. revoluta, may well produce data that justifies an infraspecific taxonomy based on vegetative characters, similar to that proposed for H. australis (Forster & Liddle 1991).
_ H. revoluta appears to be a commonly collected species that occurs in lowland riverine and coastal situations throughout its geographic range.
5. Typification and synonymy of Hoya sussuela (Roxb.) Merr.
Hoya sussuela (Roxb.) Merr., Interpret. Rumph. Herb. Amboinense 438 (1917); Asclepias “Her (19503 Fl. Ind. ed. 2, 2: 31 (1832). Type: Rumph., Herb. Amboinense 5: t. 172 (1750).
Hoya corona ariadnes Blume, Rumphia 4: 31 (1849). Type: Rumphia 4: t. 182, 185 (lecto: fide Merrill, Interpret. Rumph. Herb. Amboinense 439 (1917)).
Hoya speciosa Decne. in DC., Prodr. 8: 634 (1844). Type: Amboina, La Billardiere (holo: ?P n.v., fide Merrill, Interpret. Rumph. Herb. Amboinense 439 (1917)).
Hoya ariadna Decne. in DC., Prodr. 8: 635 (1844). Type: Rumph., Herb. Amb. 5: t. 172 (1750) (lecto: fide Merrill, Interpret. Rumph. Herb. Amboinense 439 (1917)).
Hoya coronaria var. papuana Bailey, Queensl. Agric. J. 3: 156 (1898), synon. nov. Type: Papua New Guinea: Foot of Mt Trafalgar, F.A4. Bailey (holo: BRI [AQ360787!)).
Sa
se te aN a a ee encom fa AEM atc tft tg ME A EM eh an ne a
640 Austrobaileya 3(4): 1992
Hoya hollrungii Warb., Feddes Repert. Spec. Nov. Regni. Veg. 3: 342 (1907), synon. noy. Type: Papua New Guinea. WEST SEPIK PROVINCE: Augusta Station, 1887, M. Hollrung (holo: Bf; iso: MEL 1520112!; K n.y.).
Hoya sp., Liddle, Hoya in Australia 26, 28-33 (1986); Jones & Gray, Climbing PI. Austral. 237, 252 (1988).
[Hoya rubida auct. non Schitr.: Jones & Gray, Austral. Climbing Pl. 126-127 (1977)].
[Hoya an auct. non Schumann: P. Forster & Liddle, Austrobaileya 3: 220 (1990)]
Additional specimens examined. Malesia, Amboina, Jul-Nov 1913, Robinson 90 (L; K 7.¥.); South Celebes, Danau Towuti, Timampu, Nuha, Luwu, Apr 1984, Ramilanto 168 (L; K n.y.).
Notes: H. sussuela has had a chequered taxonomic history; however, the analysis of Merrill (1917) is relatively unambiguous and his synonymy is followed here. Paramount to the identification of the common taxon that occurs in far northern Queensland and New Guinea now recognised as H. sussuela, 1s the interpretation of plates, 182 and 185 of Blume (1848) and 172 of Rumphius (1750), all of which are based on plants from the Moluccas to the west of Irian Jaya. Table 172 of Rumphius is not particularly helpful, apart from depicting a plant of section Eriostemma Schltr. Blume’s Table 182, by comparison, is well executed and the plant depicted is conspecific with this taxon common in New Guinea and Australia. Blume’s Table 185 depicts only fruit and seed. C.B. Robinson in 1913, attempted to recollect those taxa depicted by Rumphius (Merrill 1917), and his collection No. 90 from Amboina is conspecific both with the plants illustrated in Blume’s plates cited and the taxon common in New Guinea and Australia.
The original type citation for A. hollrungii requires the collection Hollrung 661. The K sheet, which we have not seen, has this number, and although the MEL sheet does not have this number, in all other respects the label data is in agreement with the original citation, and the specimen agrees with the original description. Also we have located the specimen Schlechter 14297 (WRSL) which was cited by Schlechter (1905) in his account of HW. hollrungii and this is conspecific with the taxon delimited here.
In retrospect, we believe that the name AH. lauterbachii Schumann has been misapplied both to H. sussuela in Australia and New Guinea and A. gigas Schltr. from New Guinea (Forster & Liddle 1990). The plate of H. /auterbachii in Schumann (1905) depicts a plant with more flattened campanulate corollas, long thin pedicels and a sharper angled staminal corona than the plants under consideration here. H. gigas has markedly larger flowers (50-70 mm diameter) with more widely spaced coronal lobes. As yet we have not seen either herbarium or live material that quite matches Schumann’s plant and its identity remains uncertain.
We still have not found any authentic type material of H. neoguineensis Engler, which as mentioned previously (Forster & Liddle 1990), may also be referable here. In the absence of such material this name should still be regarded as of uncertain application. As outlined previously, H. sussuela is very variable in flower size and colour (Forster & Liddle 1990), and in the herbaria cited, collections of this taxon account for nearly one third of all the holdings of Hoya material.
Acknowledgements
B. Leuenberger (B) provided a listing of extant Hoya types at B and answered various queries on the existence of specimens. The Directors/Managers of the herbaria A, B, BO, BM, BRI, BSIP, CANB, K, L, LAE, MEL, MICH, NOU, NY, P, US and WRSL allowed access to collections either at their institutions or on loan. The latin diagnoses were translated by L. Pedley. G. Leach (DNA) while Australian Botanical Liaison Officer at Kew, U.K., located and photographed various specimens in K and BM. G. Dennis, Honiara, gave freely of his field knowledge of Hoya in Solomon Islands. Some spirited discussions on various aspects of this work were held with Hoya devotees Ted Green of Hawaii and Chris Burton of Atlanta, Georgia, U.S.A. An anonymous referee kindly drew our attention to duplicates of various collections held at K. The Australian Biological Resources Study provided funding to P.I. Forster during 1988- 1990. We gratefully acknowledge this assistance.
Forster & Liddle, Hova in Papuasia, 1-5 641
References BLUME, C.L. (1849). Rumphia. 4: 31. Amsterdam: J.G. Sulpke. BURTON, C.M. (1983). Hova pottsitt recap. Hovan 5; 2. BURTON, C.M. (1989). New determination. Hoyan 11: 57-58.
FORSTER, P.I. & LIDDLE, D.J. (1990). Hoya R. Br. (Asclepiadaceae) in Australia - an alternative classification. Austrobaileya 3: 217-234,
FORSTER, P.L. & LIDDLE, D.J. (1991). Variation in Hoya australis R.Br. ex Traill (Asclepiadaceae). Austrobaileva 3: 503-521.
FOSBERG, F.R. (1940). Melanesian vascular plants. Liovdia 3: 109-124 (1940). HEMSLEY, W.B. (1894). Flora of the Solomon Islands. Bulletin of Miscellaneous Information 1894: 211-215.
HEWSON, H.J. (1988). Plant Indumentum. A Handbook of Terminology. Australian Flora & Fauna Series No. 9. Canberra: Australian Government Publishing Service.
HOOKER, W.J. (1835). Hoya pottsii. Mr. Potts’s Hoya. Curtis’ Botanical Afagazine 62: 1. 3425.
KUNZE, H. (1990). Morphology and evolution of the corona in Asclepiadaceae and related families. Tropisciie und subtropische Pflanzenwelt 76: 1-51.
LIDDLE, D.J. (1986). Preliminary observations on Hoya and Dischidia (Asclepiadaceae) in Australia. In P.I. Forster (ed.), Hova in Australia, pp. 2-37. Brisbane: Queensland Succulent Society.
MERRILL, E.D. (1917). An interpretation of Rumphius’s Herbarium Amboinense. Manila: Bureau of Printing.
MOORE, S. (1916). Asclepiadeae. In H.N, Ridley, Report on the botany of the Wollaston Expedition to Dutch New Guinea, 1912-1913. Transactions of the Linnean Society 9: 112-116.
RINTZ, ete (1978). The peninsular Malaysian species of Hoya (Asclepiadaceae). Afalaysian Nature Journal 30: 4 2.
RUMPHIUS, G.E. (1750). Herbarium Amboinense. Vol. 5. Amsterdam: F. Changuton, J. Catuffle, H. Uytwerrf.
SCHLECHTER, R. (1905). Asclepiadaceae. In K. Schumann & K. Lauterbach, Nachtrage zur Fiora der Deutschen Schutzgebiete in der Stidsee. Leipzig: Gebruder Borntrager.
SCHLECHTER, R. (1913). Die Asclepiadaceen von Deutsch-Neu-Guinea. Botanische Jahrbticher fiir Systematik, Pflanzengeschichte und Pflanzengeographie 50: 81-164.
srabenlicea hy K. (1896). Hova lauterbachii K. Sch., eime neue Wachsblume. Afonatsschrift fiir Kakteenkunde 6(1). 7
TRAILL, J. (1827). Accounts and descriptions of the several plants belonging to the genus Hoya, which are cultivated in the garden of the Horticultural Society at Chiswick. Transactions of the Horticultural Society 7: 16-30.
WHITE, C.T. & FRANCIS, W.D. (1928). Plants collected in the Mandated Territory of New Guinea. Proceedings of the Royal Society of Queensland 39: 61-70 (1928).
Accepted for publication 2 December 199]
to to ms oe AA Wht MN A ul A ta tinier a YS AA NSE AI Ak NG Ad St PO i St
Austrobaileya 3(4): 643-659 (1992) 643
THE GENUS LEPTOSPERMUM (MYRTACEAE) IN NORTHERN AU
A.R. Bean
Queensland National Parks and Wildlife Service, Box 5391, Townsville, Qld 4810, Australia
Summary
An annotated list of the 31 Leptospermum species accepted for northern Australia and Malesia is provided. L. madidum A. Bean replaces the iHegitimate name L. longifolium (C. White & Francis) S.T. Blake. Two new species, L. pallidum and L. venustum, and one new subspecies, LZ. madidum subsp. sativum, are described. L. amboinense Blume is reinstated at species level, and L. peftersoniit subsp. lanceolatum J. Thompson is synonymised with it. A multi-access key and a dichotomous key to the 31 species are provided. Significant distributional data additional to Thompson’s revision are given.
Introduction
| Leptospermum is characterised within the family Myrtaceae by its flowers with a single whorl of free stamens which are shorter than the petals, its versatile anthers, capsular fruits and alternate leaves.
A revision of the genus Leptospermum was recently published by Joy Thompson (Thompson 1989), which elucidated many matters of concern to taxonomists and provided a basis for further research. Readers are referred to that publication for descriptions and full synonymy of species not treated in this paper.
The purpose of this paper is twofold: (a) to describe two new species from Queensland, and one new subspecies from Northern Territory and Western Australia, and (b) to deal with taxonomic problems which remain in two northern Australian and Malesian species groups, 1.e. the informal L. brachyandrum and L. petersonii subgroups of Thompson (1989), in particular the circumscriptions of L. parviflorum Valeton and L. javanicum Blume.
L. parviflorum sensu Thompson and L. javanicum sensu Thompson are hetero- geneous. Thompson’s concept of L. parviflorum includes all rheophytic Leptospermum species from New Guinea and far northern Australia. However, typical L. parviflorum from New Guinea is quite different from the Australian taxon. L. parviflorum s. str. has leaves which are dark green above, densely pubescent below, with strongly recurved margins on dried material; the floral hypanthia are densely pubescent, and stem flanges are present and conspicuous. The Australian material has leaves which are pale green, concolourous, glabrous to sparsely pubescent, margins not recurved, floral hypanthia sparsely pubescent, and stem flanges absent. This taxon has been known as L. longifolium (C. White & Francis) §.T. Blake but that name is illegitimate and is replaced here by L. madidum A. Bean. Populations of L. madidum from Northern Territory and Western Australia have much narrower leaves and smaller fruits than those on other populations and have been separated as a subspecies, L. madidum subsp. sativum A. Bean.
Thompson has applied the name L. javanicum to all woody-fruited Malesian Leptospermum species except the Mt Kinabalu population of ZL. recurvum Hook. The type of L. javanicum has broad obovate discolourous leaves, pubescent floral hypanthia and fruits measuring 4-5 X 6-7 mm. While many specimens do match this description, it is clear that many presently identified as L. javanicum do not, and that they represent a distinct taxon, differing clearly from L. javanicum by its narrower lanceolate leaves (concolourous or nearly so), mostly glabrous floral hypanthia and fruits measuring 3-4 X 4.0-5.5 mm. The type of L. amboinense Blume belongs in this taxon so that name is reinstated here for it. :
L. amboinense is widespread in Malesia, even more so than L. javanicum, and also extends to north Queensland. While these two species are clearly in the same
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644 Austrobaileya 3(4): 1992
subgroup, L. amboinense is most closely related to L. petersonii Bailey and is identical with L. petersonii subsp. /anceolatum J. Thompson. In view of the altered circumscription of L. javanicum and L. parviflorum accepted here, these species are described fully in the text, as are the reinstated species L. amboinense and L. madidum.
In the annotated species list provided, species are arranged in natural order as perceived by the present author. This is similar to a subset of that used by Thompson (1989), particularly in the placement of major groups. However I have changed the position of some species. For example, L. /amellatum is placed next to L. trinervium on the basis of its brown, papery bark and seeds with small lateral wings. L. /iversidgei is removed from the Z. petersonil subgroup because it appears to have little in common with that group; for example, it lacks stem flanges, its sepals possess very few hairs, and the bark is grey and scaly. In contrast, the species in the L. petersonii subgroup have prominent stem flanges, their sepals possess hairs on the margins and apex, and the bark is brown and fibrous.
For the purposes of this paper, northern Australia is defined as including Queens- land, Northern Territory and that part of Western Australia which hes north of the 26th parallel. Malesia 1s the area defined for the Flora Malesiana project (van Steenis-Kruseman 1950), and includes the Malay peninsula, Sumatra, Borneo, Java, Philippines, Sulawesi, Moluccas and New Guinea. The genus Leptosper mum does extend slightly beyond Malesia into southern Burma (Corner 1940).
Diagnostic Attributes
Two characters useful in the identification of Leptospermum species, the presence/ absence of stem flanges and seed morphology, are worthy of brief review.
Stem flanges
One of the most diagnostic of vegetative characters in Leptospermum 1s the presence or absence of stem flanges. This term was introduced by Thompson (1989) and refers to the raised ridges of tissue on the branchlets of many Leptospermum species, extending from the base of each leaf (Fig 1A.). Stem flanges had been noted previously, as ‘angular’ or ‘triangular’ twigs for L. javanicum (Corner 1940; Backer & Bakhuizen Van Den Brink 1963; van Steenis 1972), and for L. wooroonooran (Bailey 1900). Stem flanges are consistently either present or absent for a given species, and when present they are readily visible with a hand lens. In L. sericatum, the flanges are present but poorly developed. Stem flanges are not confined to the genus Leptospermum. Of the genera in the Lepfospermum suballiance of Briggs and Johnson (1979), stem flanges are present in Homalospermum and Pericalymma (both monotypic genera), they are con- sistently absent in Neofabricia and Asteromyrtus, while in Agonis and Kunzea stem flanges are present in some species but not others e.g. they are prominent in Kunzea graniticola Byrnes but absent in K. opposita F. Muell.
Seed morphology
The seeds of most of the northern Leptospermum species have been examined in the present study. In most cases, seeds were collected from plants in their natural habitat, rather than from herbarium specimens. Thus, seed maturity could be properly assessed. Leptospermum species can be placed into two broad groups according to the characteristic of their seeds. 1. Fertile and infertile seeds are identical in appearance, being linear and striate. Germination tests have revealed that each seedlot comprises a mixture of fertile seeds and unfertilised ‘chaff. In this group are all the “woody-fruited’ species e.g. L. polygalifolium, and also L. pallidum. 2. Seeds and chaff are usually readily distinguishable, the seeds being obovoid to cuneate with a reticulate surface, and the chaff narrower and paler. This group comprises the ‘soft-fruited’ species, and the seed characters are heterogeneous. Seeds of L. venustum and L. semibaccatum are dark brown, conspicuously reticulate, and only twice as long as wide. L. microcarpum seeds are quite black, while seeds of L. trinervium have rows of extended cells forming small lateral wings.
Bean, Leptospermum 645
The Species
1. Leptospermum pallidum A. Bean sp. nov. affinis L. madido A. Bean a quo cortice aspera, pedicellis longioribus, fructibus majoribus 5-valvibus, seminibus linearibus, habitatione in summis collium differt. Typus: Marble Creek mesa, SE of Greenvale, 19°07’S, 145°04E, 20 April 1991, A.R. Bean 2949 (holo: BRI, iso: CANB,K, MEL,NSW).
Spreading shrub to 3 metres high; bark rough, grey, longitudinally fissured, closely adhering, persistent throughout; branchlets virtually glabrous, stem flanges absent. Leaves alternate, sessile or with petioles up to 2 mm long, concolourous, pale yellowish-green, narrow-lanceolate, 35-52 X 5-9 mm, more or less triplinerved, oil glands numerous, conspicuous; leaf base cuneate, apex acute to acuminate; young leaves with sparse long hairs on underside, glabrescent. Inflorescence comprising bracteolate monads on con- densed axillary shoots, 2-3 flowers on each shoot, often appearing to arise directly from leaf axils; floral bracts and bracteoles shed in early stages of bud development. Flowers 10-15 mm in diameter; pedicels 9-12 mm long, with bracteole scar approximately midway along pedicel; hypanthium slabrous, 1.5—2.0 mm long; sepals obtuse, glabrous, oil dots conspicuous; petals white, orbicular, 3-5 mm long; stamens shorter than petals, 30-40 in a single whorl; anthers versatile, cells parallel, opening by narrow slits; gland conspicuous, globular; style inset, stigma broad, capitate; roof of ovary glabrous, ovary 5-locular. Fruits dry, conspicuously domed above hemispherical base, 5-locular, 4.5-5.0 x 5-6 mm, sepals persistent. Seeds and chaff identical, pale brown, linear, striate, 2.0— 2.5 mm long, 0.2 mm wide. Fig 1.
Specimens examined: Queensland. BURKE District: Porcupine Gorge, Apr 1988, Fe// DF796 (BRI). NorTH KENNEDY DISTRICT: 20 km east of Greenvale, May 1989, Fe/i DF1809, DF181i0 (BRI): beside Charters Towers- Greenvale road, 32.3 km from Greenvale, Jun 1989, Bean 1068 (BRD): Marble Creek mesa, SE of Greenvale, Apr 1991, Bean 2942 (BRI,DNA,MEL NSW).
Distribution and habitat: L. pallidum has a restricted distribution in northern Queensland, mostly in the Greenvale area, but also at Porcupine Gorge, north of Hughenden. It occurs on lateritic ridges, often on cliff edges with skeletal soil. It sometimes grows near vine-thicket communities on rocky slopes. Associated species include Eucalyptus exserta F. Muell., 2. similis Maiden, E. persistens L. Johnson & K. Hill, F. figmen yila Brooker & A. Bean, Myrtella mic ‘ophylla (Benth.) A.J. Scott and Triodia Sp.
Flowering period: March - June.
Affinities: Leptospermum pallidum shows some resemblance to species in the genus Neofabricia J. Thompson by virtue of its rough grey bark, large leaves and domed fruits. However, L. pallidum lacks all of the most diagnostic characters of Neofabricia, which are the very numerous stamens in several irregular whorls, the dorsifixed anthers and the winged seeds (Thompson 1983). Hence it clearly belongs in Leptospermum but it has no very close relatives there. It is probably closest to L. madidum but differs from that species by its rough bark, longer pedicels, larger 5-valved fruits, linear seeds and ridgetop habitat.
Conservation status: Suggested status is 3RC, as defined by Briggs and Leigh (1988). Etymology: Named for the pale colour of the leaves.
2. Leptospermum madidum A. Bean, nom. nov.
Agonis longifolia C. White & Francis, Bot. Bull. Dept. Agr. & Stock, Queensland 22: 18 (1920). Type: Endeavour River, ?Persieh (holo: BRI, iso: NSW).
Leptospermum longifolium (C. White & Francis) S.T. Blake, Proc. Roy. Soc. Queensland 69: 81 (1958), nom. illeg.; non L. longifolium Cunn. in Heward, R.., Hooker’s Journal of Botany 4: 243 (1841).
Shrub or tree to 8 m high, ultsmate branches pendulous; bark smooth and deciduous throughout, white, creamy or pink; branchlets sparsely pubescent, glabrescent, stem flanges absent. Leaves alternate, sessile, concolourous, pale green, linear, 22-70 X 1~9 mm, apex acute or acuminate; appressed indumentum present on young leaves, but older leaves generally glabrous. Inflorescence axillary, consisting of several bracteolate
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646 Austrobaileya 3(4): 1992
monads; floral bracts shed before anthesis. Flowers 5-7 mm in diameter; pedicels 1 mm long, hypanthium sparsely pubescent, 2-3 mm long; sepals obtuse, margins ciliate; petals white; anthers versatile, cells parallel; style inset, stigma capitate; roof of ovary mostly glabrous but with hairs at base of style and along valve margins; ovary 3-locular. Fruit thin-walled, glabrous, hemispherical, 3-locular, 2-3 xX 2.5-5.0 mm, sepals persistent. Seeds light brown, obovoid to elliptical, reticulate, c. 0.75 < 0.3 mm; unfertilised seeds linear to narrowly cuneate.
Flowering period: July — October.
Affinities: L. madidum (as L. longifolium) was included within L. parviflorum Valeton by Thompson (1989). However, L. madidum can be readily distinguished from that species by the lack of stem flanges, pale concolourous leaves which are glabrous or only sparsely pubescent, leaves not recurved when dried, the glabrous or sparsely pubescent floral hypanthium and the larger fruits.
Two subspecies are recognisable as follows:
1. Leaves 4.5-9.0 mm wide; fruits 3.5-5.0 mm in diameter .... subsp. madidum Leaves 1.0-4.5 mm wide; fruits 2.0-3.5 mm in diameter ...... subsp. sativum
L. madidum subsp. madidum
Leaves linear to narrow-lanceolate, 38-70 * 4.5-9.0 mm, pale green. Fruits glabrous, hemispherical, 2.5—3.0 X 3.5-5.0 mim.
Selected specimens: Queensland. Cook DISTRICT: bank of Jardine River, Oct 1979, Scarth-Johnson 893A (BRI); north bank of Olive River, near mouth, Sep 1974, Tracey 14495 (BRD; 60 km west of Strathmay on Musgrave to Edward River road, Oct 1980, Clarkson 3498 (BRL.CANB,DNA,K,L,MO,NSW,PERTH,QRS), 10 km SE of Edward River Settlement, Oct 1983, Garnett ER497 (JCT); Archer River, beside main Cape York road, Jun 1988, Bean 845 (BRLNSW); Wenlock, Batavia River, Jul 1948, Brass 19699 (BRI); Big Bend in Coen River, 2 km N of Coen, Aug 1989, Jobson 733 (BRILNSW); Hann River, Qld, Aug 1975, Staples IBS2167 (BRI,;CANB,K), 15 km east of ‘Violet Vale’ HS., Aug 1978, Paijnians 2866 (BRI); Endeavour River, north arm crossing with MclIvor R.-Cooktown road, Nov 1981, Jrvine 2175 (QRS).
Distribution and habitat: L. madidum subsp. madidum is confined to Cape York peninsula, from Bamaga to Cooktown. It occurs on the banks of freshwater creeks and rivers, In sandy soils.
L. madidum subsp. sativum A. Bean subsp. noy. a L. madido subsp. madido foliis angustioribus, fructibus parvioribus differt. Typus: Northern Territory. Margaret River, 21 September 1946, S.7. Blake 17075 (holo: BRI; iso: DNA).
Leaves linear, 20-45 X 1.0-4.5 mm, pale green. Fruits glabrous, hemispherical, 2.0-2.5 x 2.0-3.5 mm.
Selected specimens: Western Australia. Picaninny Creek gorge, 15 km SE of Bungie Bungle outcamp, East Kimberley, Jul 1984, Kenneally 9301 (CANB,NSW,PERTH); Bream Gorge, Osmund Valley station, East Kimberley, Nov 1989, Afenkhorst 748 (DNA,MEL,PERTH). Northern Territory, Finnis River, Aug 1969, Byrnes 1684 (BRIJ,DNA); Jim Jim Falls, Sep 1984, Dunlop 6747 & Wightman (AD,DNA); Sawtooth Gorge, Nov 1972, Byrnes 2822 & Martensz (BRI); Wooler River, 16 km northwards of Telecom road on Marparu outstation road, Sep 1987, Clarke 1469 (DNA,NSW); Darwin area, Feb 1990, Wightman 4960 (BRI,DNA); Cobourg Peninsula, Jul 1982, Dunlop & Wightman 108 (DNA).
Distribution and habitat: L. madidum subsp. sativum occurs in the eastern part of the Kimberley region of Western Australia and throughout the northernmost part of the Northern Territory. It grows along riverbanks, and has often been recorded from sandstone gorges, but it is not confined to them.
Etymology: The subspecific epithet refers to the fact that this taxon has become widely cultivated in recent years.
Note: This taxon has been widely cultivated in northern Australia for several years. According to Brock (1988), it 1s fast growing and adapts to a wide range of well-drained solls.
Bean, Leptospermum 647
3. Leptospermum brachyandrum (F. Muell.) Druce, Bot. Soc. Exch. Club British Isles 1916 Suppl. 2: 632 (1917); Kunzea brachyandra F. Muell., Fragm. 2: 27 (1860). Type: New South Wales. ad ripas fluminis Hastings, Dr. Herman Beckler (lecto: NSW n.y.; 1solecto: A!).
Additional specimens: Queensland. CooK DIsTRIicT: Turtle Rock, 12 km SSE of Laura, Nov 1991, Bean 3805 (BRI,QRS), NORTH KENNEDY DIsTRicT: Mt Stuart, 9 km S of Townsville, Dec 1991, Bean 3867 (BRI,MEL,ORS); Cockatoo Creek area, Mt Elliot, south of Townsville, Aug 1991, Bean 3588 (BRI,;CANB,K,L,MEL,NSW,PERTH); Mingela Bluff, about 10 km E of Mingela, Sep 1989, Cumming 9294 (BRI); Cape Upstart peninsula, Jun 1967, Hinson CU4 (BRD; Stonehaven Bay, Hook Is., Jul 1985, Warrian CW706 (BRI).
Distribution and habitat: L. brachyandrum has a discontinuous distribution from northern New South Wales to northern Queensland, generally along rivers and creeks. However, in several localities in North Queensland (examples cited above), it inhabits steep rocky slopes or even cliff-faces. The hillside plants are completely smooth-barked, while the creek-dwelling plants almost always have rough bark at their bases, but in other respects, they do not appear to differ in any significant way. The respective habitats, while seemingly very different, are alike in that they are protected from fire. Perhaps this is an important determinant of its distribution.
4, Leptospermum parviflorum Valeton, Bull. Dep. Agric. Indes Neerl. 10: 39 (1907); Icon. Bogoriense 3: 93, t. 238 (1907). Type: New Guinea. Archip. Ind. G. Syap., Wichmann 52 (holo: L!).
Shrub or tree, 3-12 metres high. Bark type not recorded by collectors but appearing smooth and deciduous on herbarium specimens; young branchlets pubescent, stem flanges present, conspicuous. Leaves alternate, sessile, discolourous, linear, 20-45 < 2.0-4.5 mm, apex acute to acuminate, margins recurved in dried material, ‘dark green’ (fide Foreman LAE60470, Van Royen 4798) above, the lower surface much paler, densely silky-pubescent even on older leaves. Inflorescence consisting of 3—4 axillary bracteolate monads; bracts and bracteoles brown, oblong, shed well before anthesis. Flowers 5-7 mm in diameter; pedicels 1.0-1.5 mm long; hypanthium densely pubescent, 2.0-2.5 mm long, sepals obtuse, pubescent; petals orbicular, colour unknown; stamens c. 30, with small parallel anther cells; style inset, stigma capitate; roof of ovary glabrous except for short erect hairs at base of style and along valve margins, ovary 3-locular. Fruit thin-walled, pubescent, hemispherical, 1.5-2.0 X 2.0-2.5 mm, with valves not extending above the rim of the hypanthium, 3-locular, sepals persistent. Seeds and chaff identical, brown, cuneate, striate, c. 0.8 * 0.4 mm.
Selected specimens: Indonesia. Irian Jaya. Ransiki, Feb 1957, Avangold 2260 (L); Boepoel to Merau R., Merauke district, Aug 1954, Van Royen 4798 (A,L); between Boepoel and Tanas, Aug 1956, Leefers BW3224 (L,SING). Papua New Guinea. Kewa River, Sakoer, Jul 1941, Anta 72 (A,L,SING); Misool, Sorong, near Fakal, Sep 1948, Pleyte 1082 (BRILL,SING); 10.5 km west of Arufi, Morehead subdistrict, Jul 1974, Foreman LAE60470 (A,BRI,L,QRS); upper Wanggoe River basin, c. 47 miles [76 km] N of Weam Patrol Post, Aug 1967, Paijmans 333 (L).
Distribution and habitat: L. parviflorum is endemic to the island of New Guinea, occurring both in Irian Jaya and in Papua New Guinea. It grows on