ISSN 0085-441 NUYTSIA Volume 4 Number 2 1983 WESTERN AUSTRALIAN HERBARIUM— DEPARTMENT OF AGRICULTURE Cover A representation of Nuytsia floribunda (Labill.) R.Br. ex Fenzl — the Western Australian Christmas Tree. The journal is named after the plant, which in turn commemorates Pieter Nuijts, an ambassador of the Dutch East India Company, who in 1627 accompanied the “Guide Zeepard” on one of the first explorations along the south coast of Australia. NUYTSIA VOLUME 4 NUMBER 2 1983 WESTERN AUSTRALIAN HERBARIUM, DEPARTMENT OF AGRICULTURE, SOUTH PERTH, WESTERN AUSTRALIA CONTENTS A taxonomic revision of the tribe Chenopodieae (Chenopodiaceae) in Aus- tralia. By Paul G. Wilson 135 Publication date of Volume 4 Number 1 262 Editorial Board B. R. Maslin (Editor) J. W. Green P. G. Wilson Editorial Assistant J. W. Lee-Frampton Western Australian Herbarium, George St, South Perth, Western Australia 6151. Nuytsia 4(2): 135-262 (1983) 135 A taxonomic revision of the tribe Chenopodieae (Chenopodiaceae) in Australia Paul G. Wilson Western Australian Herbarium, George St, South Perth, Western Australia 6151. Abstract Wilson, Paul G. A taxonomic revision of the tribe Chenopodieae (Chenopodiaceae) in Australia. Nuytsia 4(2): 135-262 (1983). A taxonomic revision is presented for the tribe Chenopodieae (Chenopodiaceae) in Australia. Five genera and fifty two species are recognised viz.: Chenopodium (24 species), Dysphania (10 species), Scleroblitum (1 species), Einadia (6 species) and Rhagodia (11 species). One section, eight species, and seven subspecies are described as new; six new specific and eight new infraspecific combinations are made. A number of names are lectotypified, in some cases in a way different to that of earlier workers. The characters in the fruit that have traditionally been used for circumscribing the genera are shown to be unsatisfactory; this has led to the rearrangement of some species. The one species of Rhagodia described from outside the Aus- tralian and New ^aland region is considered to be a species of Chenopodium. Contents Page Introduction 136 Morphology 137 Key to genera 138 Chenopodium 139 Subgenus Chenopodium 141 Sectio Chenopodium 141 Sectio Degenia 143 Sectio Leprophyllum 144 Sectio Desertorum 151 Sectio Auricoma 161 Sectio Rhagodioides 164 Subgens Ambrosia 166 Sectio Ambrina 166 Sectio Orthosporum 169 Dysphania 180 Scleroblitum 197 Einadia 199 Rhagodia 213 Doubtful or incorrect records 235 Excluded names 235 Acknowledgements 235 References 236 Figures 239 Maps 243 Index to scientific names 260 136 Nuytsia Vol. 4, No. 2 (1983) Introduction A historical survey of the treatment by different authors of the various genera now included in the tribe Chenopodieae is given by Scott (1978a and b). He recognised two subtribes, the Chenopodiinae and the Rhagodiinae, and distinguished them principally on the presence of a dry membranous pericarp in the former and a fleshy pericarp in the latter. With regard to the genera found in Australia, Chenopodium, Dysphania, and Scleroblitum (as Chenopodium sect. Atriplicina) were placed in the Chenopodiinae, while Einadia and Rhagodia were placed in the Rhagodiinae. I have not found the nature of the pericarp to be a character suitable for discriminating subtribes since, as detailed elsewhere, it can be succulent or dry in species which by other criteria appear to be closely related. For this reason no subtribes are recognised in this paper. Furthermore I have redefined the genera Chenopodium and Einadia so as to include under both some species in which fruits are characteristically dry and others in which they are succulent. Further notes on the history and circumscription of some of the supraspecific categories are given under the respective genera or sec- tions. The species circumscription adopted here is rather broad (by the standards of workers on the Chenopodiaceae in the European flora). Many taxonomic problems remain, the resolution of which would often require extensive field work or the use of experimental methods, neither of which are within the scope of the present study. The names applied to the non-endemic species have been taken from recent papers by workers in Europe, U.S.S.R., and North America and the relevant taxa are not here critically revised. The synonymy provided for the species endemic to Australia is as complete as I have been able to ascertain. For other species the synonymy has been limited to names based on Australian collections or to names used in significant publications on Australian plants. The author abbreviations used after botanical names follow those suggested in the Draft Index of Author Abbreviations compiled at The Herbarium, Royal Botanic Gardens, Kew (1980). The abbreviations for journal titles is generally in agreement with G. H. M. Lawrence et al., Botanico-Periodicum-Huntianum (1968), while the abbreviations used for books have, where possible, followed the suggestions of F. A. ^ Stafleu and R. S. Cowan, Taxonomic Literature, ed. 2 (1976-). For each taxon usually two collections have been cited for each State in which it occurs. However, all specimens studied have been determined and all specimens bearing relatively precise geographical data have been mapped. Paul G. Wilson, Tribe Chenopodieae 137 Morphology Indumentum. A mealy indumentum is present to some extent in all species of Einadia, Rhagodia, and Sclerobhtum, while it is also present in some species of Chenopodium subg. Chenopodium. The mealiness consists initially of one-celled spherical or irregularly shaped globules that may be white and dull, or glossy and transparent. The spherical or irregularly shaped cells are supported on slender stalk cells and both arise from the division of an epidermal cell (Esau 1977). Brian and Cattlin (1968) imply a non-cellular morphology to those spherical globules that con- stitute the mealy indumentum of Chenopodium album (and therefore, by inference, other mealy members of Chenopodiaceae). This appears to be incorrect since other workers (i.e. Wood 1923; Osmond et al. 1969, and Esau 1977) have described and illustrated the origin of the indumentum from epidermal cells, while in some species of Chenopodium a transitional series may be found ranging from the subsessile globule to a slender multicelled trichome. In those cases where the mealy indumen- tum is very dense the stalk cells are of different lengths so that the spherical cells are arranged in layers; this is the situation in Chenopodium gaudichaudianum and Rhagodia spinescens. The spherical cell usually collapses on drying as the organ on which it has arisen matures. If the cell wall is relatively firm the spherical cell then becomes saucer-shaped, as is common in many species of Chenopodium, Einadia, and Rhagodia (e.g. C. desertorum, E. nutans and R. spinescens). If, however, the cell wall is weak the cells may collapse to form a scurfy covering which is eventually shed as irregular shaped flakes, or they may fuse together to form a silvery sheen on the leaf or branch. The latter state is adopted in Rhagodia candolleana and R. crassifolia, a circumstance which serves to distinguish these two from all other species of the genus. The mealy indumentum described above has been shown to have at least two functions, but not necessarily in the same species. Wood (1923) demonstrated that the collapsed cells in Atriplex vesicaria Benth. (which resemble those of Rhagodia candolleana) assist in the prevention of water loss by the plant, while Osmond et al. (1969) have shown that in some species of Atriplex the cells play an important part in excreting salt. These two activities are probably of only minor significance in those species of the Chenopodiaceae in which the mealy indumentum may be absent, or sparse and soon lost. For instance in Rhagodia eremaea the globular cells do not fuse together after collapsing but remain distinct as small scales which are shed as the leaf matures. In this species, therefore, the mealy indumentum would appear to have little influence on either the water or salt economies of the plants (see Hellmuth 1968, where referred to as R. baccata). In Dysphania and in Chenopodium sect. Orthosporum are found both glandular and simple, septate -acuminate hairs, characters which support the suggestion of a close relationship between the two groups. In Chenopodium sect. Ambrina both sessile glandular hairs and septate simple slender hairs are found on the branches and leaves, while distinctive geniculate hairs occur on the ovary. 138 Nuytsia Vol. 4, No. 2 (1983) In Chenopodium Sect. Auricoma vesicular mealy hairs are found, characteristic of those in section Leprophyllum. However, in addition, on the branches of the inflor- escence occur contorted tubular, multi-cellular hairs that are evidently of a similar nature to the vesicular since they grade into them. Sexuality. The species in the tribe Chenopodieae considered here range in their sexu- ality from dioecy through gynomonoecy to a monoclinous condition. In those taxa that are dioecious the terminal flower of a compound cyme tends towards the bisex- ual state; e.g. in a male plant of Rhagodia the terminal flower has a large pistillode, and in a female plant more prominent staminodes, than do the lateral flowers. In those taxa that are gynomonoecious the terminal flower of a cyme is bisexual while the lateral flowers are female; this is typically the situation in Einadia. In Dysphania the terminal flower is either bisexual or male (if the latter it possesses a prominent pistillode), while the lateral flowers are male. Pericarp. The pericarp varies from being diaphanous (in Dysphania), membranous (in some species of Chenopodium), leathery (Chenopodium species), succulent (Rhagodia etc.), or crustaceous (Scleroblitum). In some species (e.g. Chenopodium desertorum) it can be either succulent or dry in the same subspecies, evidently de- pendent to some extent on the environmental conditions. The unusual nature of the crustaceous pericarp of Scleroblitum is described under that genus where it is noted that although in this character it is unique among the Chenopodiaceae of Australia, it is similar to Monolepis, a genus of small herbs found in Asia and America. Key to genera 1. Plants shrubby, dioecious 5. Rhagodia Plants herbaceous or weakly shrubby, monoecious or flowers polygamous. ... 2 2. Herbs with basal rosette of leaves; tepals 4, cartilaginous in fruit 3. Scleroblitum Plants without a basal rosette 3 3. Plants glandular pubescent; tepals 1-4; stamens 1-2; embryo lateral or basal 2. Dysphania Plants glandular pubescent or mealy; tepals (3) 5; embryo variously positioned .4 4. Tepals 5; stamens 1-3 in terminal bisexual flowers; seed horizontal; perennial herbs (or weak shrubs) with mealy indumentum 4. Einadia Tepals (3-)5; stamens 1-5; seed horizontal or vertical; mostly annual herbs (rarely perennials or weak shrubs) with mealy or glandular indumentum or glabrous Chenopodium Paul G. Wilson, Tribe Chenopodieae 139 1. CHENOPODIUM L. Linnaeus, Sp.Pl. 218 (1753). Lectotype: Chenopodium rubrum L. (see Britton and Brown 1913). Orthosporum (R. Br.) T. Nees, Gen. FI. German, ad t. [58] (1835). Chenopodium sect. Orthospo'rum R. Br., Prod. 407 (1810). Lectotype: Chenopodium carinatum R.Br. (see Ulbrich 1934). Annual (rarely perennial) herbs, occasionally weak shrubs, mealy with minute sessile vesicular hairs which may collapse to form a scaly indumentum, glandular or glabrous. Leaves alternate, petiolate, chartaceous or somewhat fleshy, entire or vari- ously serrate or dissected. Flowers small, bisexual or female, usually clustered. Peri- anth 3 to 5 lobed, herbaceous or coriaceous, sometimes becoming cartilaginous in fruit. Stamens (l-)5, free or united at base into a saucer-shaped disc. Style usually very short; stigma 2 to 3(5). Pericarp membranous or rarely succulent. Seed lenticu- lar to sub-globular, horizontal to vertical; testa crustaceous; embryo annular to hip- pocrepiform; radicle centrifugal or (in vertical seeds) inferior. Seed shed either at- tached to or free from perianth. A cosmopolitan genus of over 70 species; 24 are recorded from Australia of which 14 are probably endemic. Classification The most recent outline of the infrageneric classification in Chenopodium is that provided by Scott (1978b). In that paper he surveys the historical aspects and pro- vides a considerable synonymy. I have differed from Scott on certain points, principally in the typification of some names and the circumscription or recognition of some sections. These matters are discussed under the relevant taxa. The Chenopodium species here recognised as being native or naturalized in Australia are classified as follows: Chenopodium L. subgenus Chenopodium sectio Chenopodium, species no. 1 Degenia Aellen, species no. 2 Leprophyllum Dumort., species nos. 3-9 Desertorum Paul G. Wilson, species nos. 10-12 Auricoma Aellen, species nos. 13-14 Rhagodioides Benth., species no. 15 subgenus Ambrosia A. J. Scott sectio Ambrina J. D. Hook., species nos. 16-17 Orthosporum R.Br., species nos. 18-24 140 Key to species Nuytsia Vol. 4, No. 2 (1983) 1. Plants somewhat mealy at least when young, not glandular 2 Plants glandular pubescent, not mealy 18 2. Shrubs 3 Herbs (sometimes perennial) 6 3. Fruit dry 4 Fruit succulent 5 4. Spinescent shrub; leaves leathery, spathulate, to 3 cm long . 15. C. nitrariaceum Unarmed soft-wooded shrub; leaves chartaceous, oblong-elliptic to hastate, to 4 (6) cm long 14. C. auricomum 5. Stigmas strongly penicillate 12. C. gaudichaudianum Stigmas papillose 11. C. curvispicatum 6. Tepals 5, eventually free, almost orbicular, very sparsely mealy, not shed with fruit; seed rounded on margin 13. C. auricomiforme Tepals 3-5, if 5 then mealy and enclosing fruit when shed 7 7. Leaves variously toothed 8 Leaves entire or lobed 13 8. Seed deeply muricate 8. C. erosum Seed smooth or minutely pitted 9 9. Perianth glabrous; seed horizontal or erect; perianth 3-5 lobed 10 Perianth mealy; seed horizontal; perianth 5-lobed 11 10. Perianth (at least of lateral flowers) shortly 3-lobed, urceolate 2. C. macrospermum Perianth segments free 1. C. glaucum 11. Seed prominently keeled, finely pitted, dull; pericarp persistent . . 7. C. murale Seed bluntly keeled, smooth to striate or weakly furrowed, pericarp readily de- tached 12 12. Leaves considerably longer than broad, usually more than 3 cm long 3. C. album Leaves about as long as broad, usually less than 3 cm long. . . .4. C. opulifolium 13. Plant foetid (smelling of rotten fish) 14 Plant not foetid 16 14. Tepals united to near apex; stamens 5 5. C. vulvaria Tepals free except near base 15 15. Stamens 1 or 2 6. C. detestans Stamens 5 lOd. C. desertorum subsp. virosum 16. Inflorescence a very slender panicle; tepals becoming black and enclosing fruit; leaf lamina mostly c. 10 mm long; indumentum very thin 9. C. hubbardii Inflorescence a narrow to broad panicle; tepals not black in fruit 17 17. Plant an erect annual; leaves mostly over 3 cm long with a very thin mealy indu- mentum; panicle large, som.etimes 50 cm or more long 3. C. album Plant prostrate to erect, usually a perennial; leaves always entire, usually less than 2 cm long; panicle up to 5 cm long .10. C. desertorum 18. Stamens 1-2; seed lenticular, erect; leaves simple 19 Stamens 5; seed horizontal to erect; leaves simple or pinnatisect 25 19. Tepals 4 20 Tepals 5 21 20. Plant delicate, erect; flowers in open panicles (W.A.) 24. C. saxatile Plant wiry, decumbent; flowers in dense axillary glomerules (New Zealand) 23. C. pusillum 21. Tepals strongly crested 22 Tepals rounded to acute on back or keeled 23 Paul G. Wilson, Tribe Chenopodieae 141 22. Perianth rostrate at apex; tepals free 21. C. cristatum Perianth truncate at apex; tepals united 22. C. truncatum 23. Tepals keeled (at least towards apex) 19. C. carinatum Tepals rounded on back 24 24. Tepals firmly united in lower half, completely covering fruit, usually black 20. C. melanocarpum Tepals free to near base, not completely covering fruit, white ... .18. C. pumilio 25. Leaves serrate; seed horizontal to oblique 16. C. ambrosioides Leaves pinnatisect; seed erect 17. C. multifidum Subgenus Chenopodium Sectio Chenopodium; A. J. Scott, Bot. Jahrb. Syst. 100:215 (1978). Chenopodium sect. Pseudohlitum J. D. Hook in Benth. et J. D. Hook., Gen. PI. 3:52 (1880). Type: C. rubrum L. Annual herbs more or less mealy. Flowers dimorphic in axillary and terminal pan- icles. Perianth 3-, 4-, or 5-merous; tepals free or united in lower half. Stamens 1-5. Seeds horizontal in terminal flowers, vertical in lateral flowers; embryo annular Cosmopolitan with four species of which one is found in Australia. 1. Chenopodium glaucum L., Sp. PI. 1:220 (1753). Type: From Europe, n.v. C. ambiguum R.Br., Prodr. 407 (1810). — C. glaucum var. ambiguum (R. Br.) J. D. Hook., FI. Tasm. 1:313 (1857). — C. glaucum subsp. ambiguum (R.Br.) Murr et Thellung ex Thellung, Mem. Soc. Natl. Sci. Nat. et Math. Cherbourg 38:196 (1912); Aellen in Hegi, 111. FI. Mitt.-Eur. ed. 2, 3/2:612 (1960). Type: Tasmania, R. Brown (BM). C. ambiguum var. majus Moq. in DC., Prodr. 13/2:67 (1849). No material cited. C. ambiguum var. minus Moq. l.c — C. glaucum subsp. ambiguum f. minus (Moq.) Aellen, Verb. Nat. Gesell. Basel 41:102 (1931). Type: Western Australia, Drummond 225 (iso:MEL). C. littorale Moq., Chenop. Monogr. Enum. 24 (1840) nom. illeg. non (L.) Thunb. (1815). Type: In nova Hollandia, Caley (holo:P). C. glaucum var. littorale Rodway, Tasm. FI. 155 (1903). Type: No specimens cited. C. glaucum L. var. paschale Fuentes, Bol. Mus. Nac. Chile 5:2 (1931) n.v., fide Aellen, Feddes Repert. 26:47 (1929). Type: From Easter Island, n.v. Annual herb, prostrate to decumbent or erect, to 0.5 m high, usually with several angular branches arising from base. Leaves somewhat fleshy; lamina elliptical to del- toid, 1-3 cm long, entire to sinuate dentate, often 3-lobed, obtuse, broadly cuneate at base_ and passing into a petiole about half length of lamina, upper surface glabrous, lower surface densely mealy. Inflorescence axillary and terminal of compact glomerules arranged in narrow paniculate cymes. Flowers glabrous, dimorphic; ter- minal flowers mostly hermaphrodite, tepals 3-4(5), free narrowly obovate, stamens 1- 4(5), stigmas subsessile, very short and thick, seed horizontal; lateral flowers female or hermaphrodite, tepals 3(4), stamens 0-1, seed erect. Pericarp thin, free or loosely attached to seed, somewhat green. Seed discoid, orbicular to oval, with rounded mar- gin, c. 1 mm diameter; testa finely reticulate, smooth or minutely pitted, reddish brown to black; embryo circular. 142 Nuytsia Vol. 4, No. 2 (1983) NEW SOUTH WALES: Lake George, Apr. 1898, E. Betche (NSW); Front Beach, L. A. S. Johnson 8248 (NSW). VICTORIA: Thurla, A. C. Beauglehole 56212 (MEL); Wingan Inlet. N. G. Walsh 35 (MEL). TASMANIA: Boomer, Mar. 1943, W. M. Curtis (HO); Stewarts Bay, J. H. Hemsley 6735 (HO). SOUTH AUSTRALIA: Encounter Bay, 23 May 1933, J. B. Cleland (AD); 25 km S of Naracoorte, D. Hunt 677 (AD). WESTERN AUSTRALIA: Fitzgerald R., A. S. George 11274 (PERTH); Herdsman Lake, K. F. Kenneally 1107 (PERTH). Distribution. Southern temperate Australia. Cosmopolitan. Map 1. Habitat. Usually in muddy eutrophic situations both inland and coastal. Cbenopodium glaucum is a polymorphic species in which numerous infraspecific taxa have been described. The ‘Australian’ plant is by some authors distinguished from the European as a distinct species (C. ambiguum) and by others as subspecies or variety of C. glaucum. Within Australia many variants are found, some of which may be of very recent introduction while others could be indigenous. The typical variant of C. ambiguum has deltoid 3-!obed leaves which are sinuate-dentate on the margin whereas in the typical variant of C. glaucum the leaves are narrow-ovate to narrow-elliptic with sinuate margins; Aellen (1960-1961) noted differences in the flower morphology but this varies considerably between populations which have similar leaf types. I consider it to be unprofitable to attempt to distinguish by name any of the numerous variants found in Australia since to do so would require a world-wide familiarity with the species and a knowledge of the application of the many infraspecific names. Some of the variants found in Australia correspond closely to those found in Europe so that subsp. glaucum also occurs in Australia if subsp. ambiguum should be recognised as distinct. Most Australian collections appear to be of a prostrate plant from boggy, often subsaline, situations; in Europe variants also occur which have an erect habit. Aellen (1960-1961) recognised C. glaucum subsp. ambiguum as being a native to Australia, New Zealand, and Easter Island, and as a recent introduction to Switzerland. The plant from Easter Island was earlier named C. glaucum var. pas- chale by Fuentes. Another rnember of section Cbenopodium (and the type of the genus) is C. rubrum L. This species was recorded from Victoria by Ewart (1931), however, the specimens* on which this record was based are C. glaucum; this error was pointed out by Willis Railway Reserve, N.Melbourne, May 1910, C. French and J. A. Tovey (MEL); Coode Island, 23 March 1912, J. R. Tovey (MEL). J' ' n Paul G. Wilson, Tribe Chenopodieae 143 Sectio Degenia Aellen, Magyar Bot. Lapok 25:56 (1927) n.v.; A. J. Scott, Bot. Jahrb. Syst. 100:216 (1978b). Lectotype: Chenopodium macrospermum J. D. Hook. (Scott op. cit.). Annual herbs, sparsely mealy. Flowers slightly dimorphic in cymose clusters pan- iculately arranged. Lateral flowers bisexual or female, perianth connate to near apex, very shortly 3-lobed; stamens 0-3; stigmas short and thick; pericarp membranous, free; seed usually vertical enclosed by sac-like perianth; embryo annular. Terminal flowers bisexual and either similar to lateral flowers or perianth divided to near base into 5 tepals; stamens 5; seed horizontal. Three species in America and Europe. One species (C. macrospermum) introduced into Australia. 2. Chenopodium macrospermum J. D. Hook., FI. Antarct. 341 (1846); Aellen, Feddes Repert. 26:42 (1929). Type citation: ‘Falkland Islands; Berkely Sound and St. Salvador Bay, near the sea; C. Darwin Esq., J. D. H.’ Type: n.v. Erect annual to 1 m high branched from base. Leaves alternate, somewhat fleshy; lamina rhombic or deltoid, c. 5 cm long, sinuate dentate, upper surface glabrous, lower surface mealy when young, petiole slender ± equal to lamina. Inflorescence of axillary sessile spike-like clusters forming terminal panicles. Flowers slightly dimor- phic. Lateral flowers: perianth very shortly 3-lobed, urceolate, closely enclosing fruit, c. 1.5 mm high, slightly succulent, glabrous, the lobes obscurely keeled; stamen soli- tary; stigmas 2 very short and subsessile; pericarp thin, readily detached; seed erect, flattened obovoid c. 1.5 mm long, reddish brown, glossy, testa faintly reticulate. Ter- minal flowers either similar to lateral flowers but with perianth shortly 3-4 lobed, or with perianth deeply 3-5 lobed and with 3-5 stamens. WESTERN AUSTRALIA: Perry Lakes, 7.5 km west of Perth, K. F. Kenneally 7189 (PERTH); Lake Waneragup, 18 April 1978, B. K. Masters (PERTH). Distribution. Western Australia, from Perth south to Bunbury. Probably a native of North and South America and an adventive in Europe (Aellen 1929, Standley 1931). Map 16. Habitat. Peat on margin of slightly brackish lakes. Chenopodium macrospermum is a very variable species. Aellen (1929) described several infraspecific taxa, of these, the plant introduced into Western Australia cor- responds most closely to C. macrospermum subsp. halophilum (Phil.) Aellen f. subviride Aellen. For a comprehensive synonymy see Aellen and Just (1943). Cheno- podium macrospermum subsp. halophilum was recorded for Australia by Probst (1949 p. 63) but without the citation of collections. The earliest record known to me is the one cited above dated 1978. Aellen (1960-1961) described the terminal flowers of the clusters as being deeply 3- 5 lobed, having 3-5 stamens and horizontal to vertical seeds. In Western Australia the terminal flowers are usually 3 lobed, rarely 4 lobed and only very occasionally deeply 5 lobed, while the stamens are usually solitary (rarely 3) and the seed consist- ently vertical. 144 Nuytsia Vol. 4, No. 2 (1983) I am grateful to Pertti Uotila (H) for providing the identification of a Western Australian collection of this species. Sectio Leprophyllum Dumort., FI. Belg. 21 (1827); A. J. Scott, Bot. Jahrb. Syst. 100:217 (1978). Type: C. album L. Chenopodium sect. Chenopodiastrum Moq. in DC., Prod. 13/2: 61 (1849) nom. illeg. (superfluous name). Type: C. album L. Chenopodium sect. Euchenopodium Ulbrich in Engler et Prantl, Nat. Pflanzenfam. ed. 2, 16c:487 (1934) nom. illeg. Type: C. album L. [Chenopodium sect. Chenopodium auct. pi. non sensu lectotypico.] Annual herbs somewhat mealy. Flowers in cymose clusters spicate or paniculate. Flowers hermaphrodite (and female), 5-merous, tepals united in lower half; stamens 5; stigmas 2-3. Pericarp membranous. Seed horizontal; embryo annular. Numerous species mostly extra-Australian in origin, many of which are now cosmopolitan weeds. Seven species are found in Australia. 3. Chenopodium album L., Sp. PL 1:219 (1753). Type: From Europe. Lectotype: LINN 313.8, photo seen (see Brenan 1954 p.6). C. lanceolatum R. Br., Prod. 407 (1810) nom. illeg. non Muehlenb. ex Willd. (1809). — C. browneanum Roem. et Schult., Syst. Veg. 6:275 (1820). Type: Banks of Paterson’s River, Oct. 1804, R. Brown (holo: BM). C. probstii Aellen in Probst, Mitt. Naturf. Ges. Solothurn 20 (8):56 (1928); Aellen, Verb. Nat. Ges. Basel 41:83 (1931). Type: Australia, N. Adelaide; Switzerland, Derendingen in wool compost. See comments below. Syntype: Ward St., N. Adelaide, 16 May 1923, J. M. Black (AD 97810510). Erect annual c. 1 cm high, usually much-branched, mealy white especially on flowers and undersurface of leaves. Stem round to angular with green stripes. Leaves thin, the lower petiolate, ovate rhombic, cuneate at base and variously dentate, the upper narrowly ovate to lanceolate entire, more or less sessile, the longer ones c. 4-6 cm long. Inflorescence largely leafless of sessile flowers in cymose glomerules aggre- gated in a large paniculate, spicate, or indeterminate arrangement. Flowers her- maphrodite; tepals 5, united in lower half, broadly ovate, obtuse, membranous on margin, weakly keeled (terminal flowers larger and more strongly keeled), green and somewhat mealy. Stamens 5, glabrous, disc absent. Pericarp membranous, glabrous, readily detached. Seed horizontal, lenticular, c. 1.2 mm diam., bluntly keeled; testa black, glossy, smooth, striate or weakly and irregularly furrowed (reticulate in var. reticulatum). Seed shed within the surrounding perianth. QUEENSLAND: Tamborine Mt., Apr. 1947, M. S. Clemens (BRI); Charleville, M. M. Biddulph 195 (BRI). NEW SOUTH WALES: Cooma, 15 Mar. 1955, E. Gauba (CBG); Canberra, N. T. Burbidge 7484 (CANB). VICTORIA: SE of Wallacedale, A. C. Beauglehole 30512 (MEL); 3 mi [4.8 km] east of Tawonga, R. V. Smith 64/67 (MEL). TASMANIA: Flinders Is., J. Whinray 23 (MEL); Macquarie Plains, Mar. 1861, W. M. Curtis (HO). Paul G. Wilson, Tribe Chenopodieae 145 SOUTH AUSTRALIA: 5 km south of Blyth, A. E. Orchard 364 (AD); Adelaide, B. Copley 1310 (AD). WESTERN AUSTRALIA: Albany, G. Perry 259 (PERTH); Manjimup, R. D. Royce 1463 (PERTH). NORTHERN TERRITORY: Harper Springs, 26 Jan. 1949, W. G. Steenson (NT). Distribution. All Australian states and territories but rare in the tropics. Cosmopoli- tan. Map 2. Habitat. Widespread as a weed in agricultural areas. The name C. album has here been used in a broad sense and may encompass sev- eral segregate species recognised by some European workers, including C. suecicum J. Murr, C. berlandieri Moq. and C. giganteum D. Don. No attempt is made here to distinguish the numerous infraspecific taxa which have been described, since such an undertaking would require the attention of a specialist who is familiar with the com- plex from different parts of the world. This is beyond the scope of this paper. A plant which is readily recognisable in fruit is Chenopodium album var. reticulatum (Aellen) Uotila (1978 p. 31). This variety may be distinguished by the re- ticulate testa; it occurs sporadically with the other variants across Australia. The application of the name C. probstii is in doubt. The plant was originally as- sumed by Aellen (1928 op. cit.) to have originated in South Australia and to have been introduced into Europe as a contaminant of wool. Jorgensen (1973 p. 314), while accepting it is a distinct species, throws doubt on its Australian origin and suggests it comes from North America. It has never been listed in an American flora and, if present in America, is presumably there included under C. album. In a study of C. album and C. suecicum in northern Europe Uotila (1978) discusses C. probstii and suggests that its taxonomy will only be understood after a thorough study has been made of the biology of the species in the C. album complex of both North and Central America. A lectotype should only be chosen after such a study is undertaken. Aellen (1960-1961) and Beauge' (1974) suggest a possible synonyomy of C. probstii with Chenopodium serotinum L. (which itself is considered to be a synonym of C. album by Beaug6). Kjellmark (1934) determined the chromosome number as 2n=54, i.e. as a hexaploid; it therefore corresponds to that of C. album and C. opulifolium, q.v. 4. Chenopodium opulifolium Schrader ex Koch et Ziz, Cat. PI. Palat. 6 (1814); DC., FI. Francaise 6:372 (1815). — C. album subsp. opulifolium (Koch et Ziz) Maire, FI. de I’Afrique du Nord 8:36 (1962). Type: From Palatinate (Pfalz), Germany, Schrader (S photo seen), see nomenclatural note below. ?C. triangulare Forsskal, FI. Aegypt. Arab. 205 (1775). Type: From ‘Taaes’. Yemen, n.v. Very similar to (and sometimes considered conspecific with) C. album. It differs from the latter species in having long-petiolate 3-lobed rhomboid leaves 2-3 cm long and wide, strongly mealy below, and in having keeled strongly mealy tepals. SOUTH AUSTRALIA: About Mortlock Experimental Station, Mintaro, 5 March 1969, D. E. Symon 6699 (ADW, CANB). 146 Nuytsia Vol. 4, No. 2 (1983) Distribution. Southern South Australia. Central and southern Europe, north Africa. Habitat. A weed of agriculture. This species is rather variable in morphology and difficult to distinguish from C. album, another highly complex taxon. The only unequivocal record is the Symon specimen cited above although other collections included by me under C. album may belong to C. opulifolium. It was first recorded in Australia by J. M. Black (1912) who cited a specimen from Henley Beach, South Australia but this was later seen by P. Aellen (1927 in litt.) who considered it to be better placed under C. album. An ad- ditional character useful for distinguishing the two species has been pointed out by Beaug6 (1974 p. 305); he notes that in the seed of C. album the micropyle has the form of a dimple (a scar) while in C. opulifolium it resembles a knob (i.e. it is con- vex). I have been unable to confirm tbis observation in material of C. opulifolium available to me. Work by Cole (1961, 1962) and Wilson (1980) has shown that both C. album and C. opulifolium are hexaploids (2n=54) unlike most other weedy species of the section which are either diploids (2n=18) or tetraploids (2n=36). Battandier (1888-1890 p. 753) states that one can find all intermediates between the two species. It would therefore be reasonable to accept a subspecies status for C. opulifolium as was done by Maire (1962 op. cit.). Nomenclature. In the publication by Koch and Ziz (1814 op. cit.) the name ‘Chenopodium opulifolium. Schrad in. lit.’ appears, without a description but with a reference to ‘Ch. album Borkh. in Rhein Mag.’ i.e. to M. B. Borkhausen (ed.), Rheinisches Magazin zur Erweiterung der Naturkunde vol. 1:115 (1793). Here there is a description of the form of Chenopodium album found in the valley of the Rhine. I have not attempted to lectotypify this species nor to confirm that the plant de- scribed by Borkhausen is the same as that to which Schrader gave the name C. opulifolium (although the description agrees). The name C. opulifolium was also validly published by Decandolle (1815 op. cit.) with reference to the publication by Koch et Ziz (1814), to ‘C. viride Loiseleur’ (1806) and to ‘C. erosum Bastarde’ (1814). Beauge" (1974) cites as type a Schrader collection from ‘Palatinate’. The name Chenopodium triangulare Forsskal (1775) was given by Beauge' (1974) as a probable synonym of C. opulifolium-, it is based on a plant from Yemen and has generally been placed in synonymy under C. murale L. (e.g. by Christensen 1922). Beaug^ points out that the latter synonymy could not be correct and that the only species found in Yemen to which the description applies is C. opulifolium. If this is so then C. triangulare is the correct name for this species. The type was not seen by Christensen and is not present in Copenhagen (herb. C); I have not attempted to search elsewhere. 5. Chenopodium vulvaria L., Sp. PI. 220 (1753). — C. olidum Curtis, FI. Londinensis lasc. 5, t. 20 (c. 1785). Type-. From Europe, n.v. Spreading annual c. 20 cm high branching from base with a strong fish-like smell (of trimethylamine). Branches soft, mealy with thin-walled vesicular hairs. Leaves alternate; lamina ovate to broadly ovate or trullate, 1-2 cm long, entire, apex acute, base cuneate, upper surface subglabrous, lower surface mealy, petiole slender c. half length of lamina. Inflorescence of dense clusters of flowers forming short compact axillary and terminal thyrses 1-2 cm long. Flowers hermaphrodite (terminal) and fe- Paul G. Wilson, Tribe Chenopodieae 147 male (lateral); perianth subglobular c. 1 mm diameter, shortly 5 lobed, enlarging with and enveloping the fruit, densely mealy outside. Stamens 5, glabrous, united at base into a cup-shaped disc, completely absent in female flowers. Pericarp membranous, papillose, somwhat adherent. Seed horizontal, lenticular with obvious keel, c. 1-2 mm diameter; testa radially lineate. NEW SOUTH WALES: Binnalong, Jan. 1922, J. J. Parry (NSW); Berridale, Nov. 1914, D. J. O’Rouke (NSW). VICTORIA: Warrnambool, H. B. Williamson 1348 (MEL); Omeo, R. V. Smith 64/29 (MEL). SOUTH AUSTRALIA: Bool Lagoon, D. Hunt 1938 (AD). Distribution. Eastern New South Wales, Victoria, south-eastern South Australia. North Temperate areas. Map 16. Habitat. Principally a weed of horticulture. Chenopodium vulvaria may be recognised by its fish-like odour and by its shortly lobed perianth which encloses the fruit. Features which distinguish it from C. de- testans are discussed under the latter species. Chenopodium vulvaria was first recorded in Australia by Ewart (1908) who stated that it was “introduced with impure agricultural seed, and now widely spread”; the earliest dated collection I have seen from Australia is the one cited above made by O’Rourke in 1914. 6. Chenopodium detestans Kirk, Trans, and Proc. N. Zeal. Inst. 9:550 (1877); Murr, Allg. Bot. Zeitschrift 16:57 (1910); Allan, FI. N. Zeal. 1:228 (1961). Type ci- tation: “Originally discovered by Mr J. D. Enys, who informs me that he has seen it in several localities in Canterbury”. Lectotype: Trelissick Basin, Canterbury, New Zealand, 4 Feb. 1876, T. Kirk 821 (holo: WELT; iso: WELT), (see Allan 1961). Prostrate to ascending annual herb with foetid odour, numerous wiry branches to 15 cm long arising from base. Leaves alternate; lamina thin, trullate, 5-10 cm long, entire or with a pair of lateral teeth, sparsely to moderately mealy with round vesicu- lar hairs. Inflorescence of short dense axillary thyrses up to 5 mm long. Flowers mostly hermaphrodite; perianth erect in young flower, becoming incurved and ob- scuring fruit; tepals (4)5, thin, broadly ovate with short broad claw, cucullate and fleshy at apex, united towards base, mealy outside. Stamens 1 or rarely 2, completely absent in female flowers, divaricate at anthesis, ± equal to tepals; filament linear glabrous; disc absent. Pericarp membranous, prominently papillose, somewhat ad- herent. Seed discoid to lenticular with rounded margin, c. 1.2 mm diam.; testa punc- tate, black. NEW SOUTH WALES: Nimitybelle near Cooma, Dec. 1896, J. H. Maiden (NSW 143153). Distribution. Near Cooma, New South Wales, probably introduced; South Island, New Zealand. Habitat. In New Zealand “lowland to montane open tussock-grassland and bare places” fide Allan (1961). 148 Nuytsia Vol. 4, No. 2 (1983) Chenopodium detestans resembles C. vulvaria in having the foetid odour of rotting fish caused by the chemical trimethylamine which is presumably located in the oil glands that are plentiful in the leaves and in the cortex of the stems. Apart from the different habit and larger leaves of C. vulvaria, it differs from C. detestans in having almost completely united tepals, usually 5 stamens united into a disc at base, and an obvious keel to the seed. The orginal description of C. detestans and that provided by Allan (1961) stated that the flowers have four stamens (a most unusual number) whereas in material I have examined from New Zealand and Australia (including the lectotype) the number in the hermaphrodite flowers has been one or rarely two. Chenopodium detestans is presumably endemic to New Zealand (the solitary re- cord from Australia, cited above, is probably an introduction), however, there is also the possibility that it is an early introduction to New Zealand from South America since it behaves as a weed and is quite distinct from other Australian or New Zealand endemic species. The absence of recent Australian collections suggests that it is not common in that country, although in the field it could have been overlooked or con- fused with C. vulvaria. Neither Kirk (1877 op. cit.) nor Allan (1961 op. cit.) suggested the probable affin- ities of C. detestans and this was not mentioned by Aellen (1960-1961) or Scott (1978b) in their conspectuses of the genus. Murr (1910) placed it in sect. Chenopodiastrum (= sect. Leprophyllum), in which decision he was followed by Maiden and Betche (1916). Cheeseman (1925) and Ulbrich (1934) suggested an affin- ity to C. vulvaria, probably being influenced by the odour the two species have in common; the latter author placed it with C. vulvaria in sect. Euchenopodium (= sect. Leprophyllum). My studies suggest that Chenopodium detestans is most closely re- lated to C. carnosulum Moq. (1849) and C. scabricaule Speg. (1902) both of which are indigenous to South America (C. carnosulum is also recorded from Mexico and Texas). In the broad sense the three taxa could reasonably be treated as being conspecific for they are very similar in vegetative and floral characters. Aellen and Just (1943) placed C. scabricaule in synonymy under C. carnosulum, however, Giusti (1970) and Planchuelo (1975) considered them to be distinct being distinguished in part by the free pericarp of the former species and adherent pericarp of the latter. Chenopodium detestans has an adherent pericarp and thus in this character is more similar to C. carnosulum', it differs from both C. carnosulum and C. scabricaule in having a foetid smell, a character which is noticeably lacking (according to available descriptions) from the two American species. 7. Chenopodium murale L., Sp. PI. 1:219 (1753). Type: From Europe, n.v. Figure 4B. C. biforme Nees, PI. Preiss. 1:636 (1845). — C. murale var biforme (Nees) Moq. in (1849). Type: ‘In arenosis prope Pointwater, Perth’, Preiss 1256 (iso: MEL). C. congestum J. D. Hook., London J. Bot. 6:280 (1847). — Rhagodia congesta (J. D. Hook.) Moq. in DC., Prodr. 13/2:51 (1849). — R. baccata var. congesta (J. D. Hook.) J. P.‘,, Tasmaniae 312 (1857); A. J. Scott, Feddes Report. 89:7 (1978). — R. Mlardieri var. congesta (J. D. Hook.) Benth., FI. Austral. 5:153 (1870). Type: ‘Hobart Town,’ Oct. 1840, Gunn 868 (holo: K). Paul G. Wilson, Tribe Chenopodieae 149 Erect branched annual to 1 m high, sparsely mealy with thin walled vesicular hairs, sometimes foetid when crushed. Stem angular. Lower and middle leaves petio- late, lamina thin, broadly triangular to ovate, 2-8 cm long, mealy when young, acute, broadly cuneate at base, coarsely and irregularly dentate with slightly incurved teeth. Inflorescence of compact cymes in rather open panicles, axillary and terminal. Flowers hermaphrodite (or female); perianth sparsely mealy outside (becoming glabrous with age); tepals 5, c. 1.5 mm long, united in lower third, broadly ovate, ob- tuse, prominently keeled and slightly cucullate at apex, membranous and minutely ciliate on margin. Pericarp prominently papillose, often brownish, persistent. Seed horizontal, lenticular, 1-1.5 mm diam., prominently keeled; testa glossy, minutely pitted; embryo circular. Seed shed with surrounding perianth. QUEENSLAND; Brisbane, A. Dietrich 2307 (NSW); Rockhampton, P. O’Shanesy 210 (MEL). NEW SOUTH WALES: Pimpara Ck., S. Jacobs 1089 (NSW); Lord Howe Is., A. Rodd 1453 (NSW). VICTORIA: Port Phillip, Apr. 1886, C. Walter (NSW); Port Fairy, 1894, H. B. Williamson (MEL). TASMANIA: Stanley, Feb. 1948, W. M. Curtis (HO); Hobart, L. Rodway 9398 (HO). SOUTH AUSTRALIA: Innes Nat. Park, C. R. Alcock 4821 (ADW); Mt Serle HS., D E. Symon 4025 (ADW). WESTERN AUSTRALIA: Midstream, M. /. H. Brooker 2103 (PERTH); Port Denison, R. A. Saffrey 1557 (PERTH). NORTHERN TERRITORY: Alice Springs, 27 Jan. 1976, A. Mitchell (NT)- Woodgreen HS, D. J. Nelson 2131 (NT). Distribution. Principally extratropical Australia. Cosmopolitan. Probably indigenous to the Mediterranean area and to southern and south-western Asia (Engstrand and Gustafsson 1973). Map 5. Habitat. A weed of agriculture and of disturbed areas, also coastal sands. Numerous infraspecific taxa have been described from Europe but no attempt has been made in this paper to relate these names to the plants found in Australia. The name Chenopodium congestum (1847) was based on a collection found on the south coast of Tasmania in which no fruits were observed by the author. Moquin (1840), although he saw only the type collection, described the young fruit as being slightly fleshy for which reason he transferred the species (but with reservations) to Rhagodia. Later, J. D. Hooker (1857 op.cit.) while accepting it as a member of this genus made it a variety of R. baccata in which decision he was followed by Bentham (1870). Neither Rodway (1903) nor Curtis (1967) mention the taxon. Scott (1978a) retained it as a variety of R. baccata although his description of the taxon applies to R. candolleana s. str. and two of the four collections he cites belong to that species; the other two I have not seen. It is curious that no mention was made by Scott of the dentate leaves of the type which very obviously distinguish it from any species of Rhagodia, while the nature of the pericarp and testa conclusively identify it as being 22493 — ( 2 ) 150 Nuytsia Vol. 4, No. 2 (1983) C. murale. The young fruit of C. murale, when dry, has a leathery pericarp and it is probably for this reason that Moquin considered it would be somewhat fleshy when mature. Chenopodium murale was presumably an early introduction to Australia. It may be readily recognised by the usually glabrous leaves with prominent antrorse teeth and by the morphology of the seed. 8. Chenopodium erosum R. Br., Prodr. 407 (1810); Moq., Chen. Mon. Enum. 33 (1840); Moq. in DC., Prodr. 13/2:68 (1849); J.D. Hook., FI. Tasm. 1:313 (1857). Type: In collibus arenos: Insularum Kents Group in fretu Bass, Dec. 1804, R. Brown (holo: BM; iso: BM). Figure 4A. [C. murale auct. non L.: Benth., FI. Austral. 5:160 (1870) p.p. as to the R. Brown specimen cited.] “C. sp. aff. suecicum J. Murr”, Willis, Handb. PI. Viet. 2:86 (1973). Erect slender or bushy annual to 1.5 m high. Stems ribbed, with red stripes, glabrescent. Leaves papery; lamina triangular to ovate 3-15 cm long, 1.5-10 cm wide, acuminate, base somewhat truncate, margin deeply incised, undersurface pubescent with thin-walled bladder-hairs when young, becoming glabrous; petiole 1/3 to 1/2 length of lamina. Inflorescence of compact terminal and axillary paniculate cymes, becoming open with, age. Flowers hermaphrodite and female, sessile; tepals 5, shortly united at base, obovate, c. 1 mm long, with broad scarious margins and a fleshy mid- rib, pubescent with thin-walled bladder-hairs when young, becoming glabrous; stamens 5, glabrous, filaments linear, disc absent; ovary glabrous. Pericarp membra- nous, dull, adherent to seed. Seed horizontal, lenticular with rounded margins, c. 1.5 mm diam.; testa black, glossy deeply muricate with radial striations. Seed eventually falling free from perianth. QUEENSLAND: Gayndah, 13 May 1917, C. T. White (BRI); Upper Logan, Anon. 284 (MEL). NEW SOUTH WALES: Moona, Walcha, A. R. Crawford 464 (MEL); Coneac, Johnson et Briggs (NSW 142705); Tinderry South Peak, J. H. Willis (MEL 80686). VICTORIA: Bonang— Gelantipy Rd., A. C. Beauglehole 37698 (NSW); Gippsland, F. Mueller (MEL 80689); 3 mi [4.8 km] NE of Suggan Buggan, J. H. Willis (MEL TASMANIA: Kent Group, Bass Strait, Dec. 1804 R. Brown (BM). SOUTH AUSTRALIA: South-west River, Kangaroo Is., J. B. Cleland (AD); Ardrossan, J. G. O. Tapper 915 (AD). Distribution. South-west Queensland to Bass Strait, southern South Australia; New Zealand. Map 6. Habitat. River banks, forest lands, often in disturbed areas. This species is one of two in the sect. Leprophyllum apparently native to Aus- tralia. Other members of the section which have been described from Australia are now considered conspecific with European species and, therefore, probably represent Paul G. Wilson, Tribe Chenopodieae 151 relatively recent introductions to that country. Chenopodium erosum itself has the aspect of an introduced Eurasian species, however, I have been unable to match it with any extra-Australian plant. Although C. erosum is found over a wide area it has been collected only in- frequently, Robert Brown’s collection of 1804 being the solitary one for Tasmania. In New Zealand it is also widespread but rare; all collections in that country were made this century (W. R. Sykes, CHR, pers. comm.). 9. Chenopodium hubbardii Aellen, Candollea 8:19 (1940); Aellen in Hegi, 111. FI. Mitt.-Eur. ed. 2, 3:616 (1960). Lectotype: Mungallala, Queensland, 31 Dec. 1930, C. E. Hubbard and C. W. Winders 6022 (holo: K n.v.; iso: BRI) lecto. nov. Erect annual herb branching from base, c. 30 cm high. Branches slender, striate, sparsely mealy when young. Leaves alternate; lamina thin, ovate to hastate with rounded apex and lobes, c. 10(30) mm long, entire, upper surface sparsely mealy, lower surface mealy grey due to the opaque discrete vesicular hairs; petiole slender from half as long to as long as lamina. Inflorescence a terminal slender panicle c. 2 cm long (lengthening to 10 cm in fruit); flowers in compact glomerules (which be- come separated with age) subtended by filiform bracts, the branches terminated by a solitary bisexual flower (the others female). Flowers depressed globular c. 1 mm diam.; tepals orbicular, sessile (or in fruit shortly clawed), densely mealy outside with grey vesicular hairs. Bisexual flower, stamens 5, filaments linear broadened at base and united into a thick disc; ovary glabrous; style very short (c. 0.1 mm); stigmas 2, spreading, 0.3 mm long. Female flower: stamens and disc absent; ovary as in bisexual flower. Fruiting perianth eventually black, depressed globular, hard, closely envelop- ing fruit; tepals rounded on back, orbicular very shortly clawed. Pericarp brown to black, finely papillose, adherent to testa. Seed lenticular, 1.5 mm diam. with rounded margin; testa finely and irregularly pitted. Seed shed with enveloping perianth. QUEENSLAND: 70 mi [112.6 km] S of Blackall, IF. J. Bissett E 233 (BRI); Morven, S. T. Blake 10911 (BRI). Distribution. South-central Queensland. A casual in Switzerland where introduced in wool. Map 7. Habitat. Usually recorded from heavy soils which become seasonally waterlogged. Chenopodium hubbardii was considered by Aellen (1940 op.cit.) to be closely re- lated to C. polygonoides and he later (1960-1961) included it in Chenopodium sect. Polygonoidea. The two species are similar in appearance and both have persistent pericarps and pitted testas but, whereas C. polygonoides has only a single stamen, C. hubbardii has five. For this reason I am treating the latter as a member of section Leprophyllum while C. polygonoides I am transferring to the genus Einadia. This formal separation at the generic rank obscures the probable close relationship be- tween the two species but the alternative, that of transferring all the species of Einadia to Chenopodium, would cause considerable systematic confusion. Sectio Desertorum Paul G. Wilson, sect. nov. Herbae perennes vel suffructices, farinosi. Flores monoid; perianthium farinosum, 5-lobum. Stamina 5, discus patelliformis, sparse puberulus. Fructus baccatus vel 152 Nuytsia Vol. 4, No. 2 (1983) siccus; pericarpium glabrum, membranaceum vel succulentum, mesocarpio aquoso. Semen horizontale, lenticulare vel depresso sphaericum; embryo annularis. Typus: Chenopodium desertorum (J. Black) J. Black. Perennial herbs or weak subshrubs largely covered with a mealy indumentum of discrete globular vesicular hairs. Flowers monoecious, those terminal to the inflor- escence and its branches male, the others female. Perianth 5 lobed, mealy outside with vesicular hairs. Stamens 5, united at base into a sparsely puberulous disc. Fruit baccate or dry; pericarp glabrous, free from seed, membranous to succulent with a watery mesocarp. Seed horizontal, lenticular to depressed spherical; embryo annular. Three species endemic to Australia. The species in this section have, in floras and herbaria, frequently been confused with each other, for they are similar in indumentum, leaf-shape, and in flower and fruit morphology. In habit they are intermediate between herbs and shrubs being either herbaceous perennials or weak shrubs with straggly branches. The fruits when succulent have a watery interior and, on drying, leave either an inflated or a thin wrinkled pericarp which is often sticky but lacks the viscous character of the meso- carp in Rhagodia. Because of their often shrubby nature and their often succulent or semi-succulent fruit, these species have sometimes either been placed in Rhagodia or confused with one of its species. In that genus they are obviously misplaced; their semi-woody habit, their monoecious flowers, and their watery fruits (when baccate) clearly set them apart, while their habit and fruit also make them out of place in the genus Chenopodium. To segregate this group of species at the generic level is unde- sirable due to the similarity between some variants of C. desertorum and such north- ern-hemisphere species of Chenopodium as C. vulvaria. I have therefore included them in that genus, a course which is least disturbing of current taxonomic practice. Within Chenopodium the species of section Desertorum have most in common with those of subsection Lejosperma Aellen et Iljin of section Chenopodium (correctly of sect. Leprophyllum) in which are included the species C. album, C. opulifolium, and C. vulvaria. 10. Chenopodium desertorum (J. Black) J. Black, FI. S. Austral. 181 (1924) et in Trans. Roy. Soc. S. Austral 49:272 (1925). — C. microphyllum var. desertorum J. Black, Trans. Roy. Soc. S. Austral. 46:566 (1922). Type citation: ‘Murray lands; Port Augusta westward to Ooldea’. Lectotype: Ooldea, Nov. 1921, D. Bates (holo: AD 97811131; iso: MEL 545186, NSW 143061) lecto. nov. Rhagodia prostrata A. Cunn. ex Moq. in DC., Prod. 13/2:52 (1849) non C. prostratum (Pallas) Roemer et Schultes (1820). Type citation: ‘In Nova-Hollandia (Cunning.l)’. Lectotype: N.E. from L.R. [i.e. Lachlan River], 5 Aug., A. Cunningham (K) lecto. nov. C. microphyllum F. MuelL, Trans. Phil. Inst. Viet. 2:74 (1858) nom. illeg. non Thunb. (1794). C. triandrum var. lanuginosum F. E. Haviland, Proc. Linn. Soc. New South Wales 36:532 (1912) based on preceding. — C. pseudomicrophyllum Aellen, Candollea 8:8 (1939) based on C. microphyllum F. Muell. Lectotype: Bacchus Marsh, 1853, F. Mueller (lower left-hand specimen of MEL 80671) (MEL) lecto. nov. C. anidiophyllum Aellen, Candollea 8:9 (1939). Lectotype: Dubbo, Feb. 1886, E. Eetche (holo: G n.v.; iso: NSW 143105) lecto. nov. C. cochlearifolium Aellen, Candollea 8:10 (1939). Lectotype: Lower Loddon R., Vic- toria, Nov. 1887, C. Walter (holo: G n.v.; iso: MEL 80675, NSW 142716) lecto. nov. Paul G. Wilson, Tribe Chenopodieae 153 Annual or perennial herbs or subshrubs, prostrate to erect, branched from the base, covered at least when young with vesicular hairs or (inflorescence) with branched or simple tubular hairs. Leaves opposite or alternate, chartaceous to some- what fleshy, subsessile or with slender petiole; lamina elliptic, deltate, hastate or or- bicular, 5-20 mm long. Inflorescence a narrow to broad panicle, shorter than or much exceeding terminal leaves, flowers clustered in sessile (basically 3-flowered) cymes; terminal flower of inflorescence and of its principal branches male or bisexual, lateral flowers female. Flowers sessile, depressed -globular, 1-2 mm diameter; tepals 5, ovate, shortly united at base, thickened in centre, scarious along margins, densely vesicular hairy outside, spreading at anthesis. Male flower: stamens 5; filaments linear, united at base into a circular disc which is sparsely vesicular pubescent within; pistillode usually minute. Female flower: staminodes absent; ovary glabrous; style 0.2-0.5 mm long; stigmas slender. Fruit enveloped by perianth; pericarp membranous to slightly succulent and then separated from seed by a watery sap, sometimes sticky. Seed horizontal, lenticular with rounded margin, 1-1.5 mm diameter, testa almost smooth to slightly rugulose or striate, black. Key to subspecies 1. Floral indumentum of glistening irregularly shaped vesicular hairs a. subsp. desertorum Floral indumentum of dull, grey to white vesicular hairs 2 2. Plant prostrate to decumbent; leaves small to 10 mm long; flowers small, c. 1 mm diameter in short inflorescences 3 Plant erect; leaves 5-20 mm long, flowers 1-2 mm diameter; inflor- escences usually exceeding terminal leaves 4 3. Leaves elliptic to orbicular, often glabrescent above; plant without a noticeable smell b. subsp. microphyllum Leaves narrowly elliptic, mealy on both surfaces; plant with foetid odour d. subsp. virosum 4. Branches slender, straight and rigid; leaves small (to 10 mm long), orbi- cular to obovate, sparsely mealy; flowers small (c. 1 mm diam), sparsely to moderately covered with sessile vesicular hairs e. subsp.rectum Branches straight or flexuose; leaves various, densely mealy (at least below); flowers with globular or branched often stipitate hairs . c. subsp. anidiophyllum 10a. subsp. desertorum Erect, rounded, much-branched herb to 20 cm high. Leaves densely covered when young with colourless, glistening, transparent, branched vesicular hairs; lamina del- toid to orbicular, 3-8 mm long. Inflorescence congested or loose, narrowly pyramidal or slender, exceeding the terminal leaves and often up to 3 cm long. Flowers 1.5-2 mm diameter; perianth with a felty indumentum of glistening transparent irregularly shaped vesicular hairs. Seed c. 1.5 mm diameter. NEW SOUTH WALES: 5 mi [8 km] W of Moulamein, C. W. E. Moore 3400 (CANB); 128 km E of Wilcannia, 14 May 1971, G. J. White (NSW). VICTORIA: Annuello, A. C. Beauglehole 55816 (MEL); Wyperfeld Nat. Park A C Beauglehole 29462 (MEL). SOUTH AUSTRALIA: Tallaringa Well, T. R. N. Lothian 3836 (AD); 10 km W of Blanchetown, D. J. E. Whibley 3741 (AD). 154 Nuytsia Vol, 4, No. 2 (1983) WESTERN AUSTRALIA: Fitzgerald R. Nat. Park, K. Newbey 4142 (PERTH); Salmon Gums, R. D. Royce 3539 (PERTH). NORTHERN TERRITORY: Ormiston Gorge area, J. R. Maconochie 2244 (CANB). Distribution. Northern South Australia south-eastwards to central and western New South Wales and north-west Victoria; rare in southern Western Australia and the Northern Territory. Map 6. Habitat. Recorded from deep sand but evidently also found in other well-drained soils. The typical variant of this subspecies is a compact perennial with felty leaves and flowers. In western Victoria a variant is found with a looser habit and large leaves with a less dense indumentum; the hairs of this variant are of the same nature as those of the type. 10b. subsp. microphyllum Paul G. Wilson, nom. et stat. nov. — Basic name C. micTophyllum F. MuelL, Trans. Phil. Inst. Viet. 2:74 (1858) nom. illeg. non Thunb. (1794). — C. triandrum var. lanuginosum F. E. Haviland, Proc. Linn. Soc. New South Wales 36:532 (1912), based on preceding. — C. pseudomicrophyllum Aellen, Candollea 8:8 (1939), based on C. microphyllum F. Muell. Lectotype: Bacchus Marsh, 1853, F. Mueller (lower left-hand specimen of MEL 80671) (holo: MEL) lecto. nov. Rhagodia prostrata A. Gunn, ex Moq., op. cit. C. cochlearifolium Aellen, op.cit. [Rhagodia spinescens auct. non. R. Br.: Benth., FI. Austral. 5:155 (1870) p.p. as to A. Cunningham specimen cited.] [C. triandrum auct. non. G. Forster: Murr, Allg. Bot. Zeitschr. 16:56 (1910); Maiden et Betche, Census N.S. Wales PI. 66 (1916).] Intricately branched prostrate to decumbent perennial with prominent taproot; not noticeably foetid. Leaves thin to slightly fleshy, elliptic to orbicular, shortly petiolate, 5-10 mm long, undersurface covered with small simple rounded white vesi- cular hairs, upper surface glabrescent. Inflorescence short, scarcely exceeding the ter- minal leaves. Flowers c. 1 mm diameter; perianth covered with small round white vesicular hairs. Seed c. 1 mm diameter. NEW SOUTH WALES: 15 mi [24.1 km] W of Rankin Springs, J. He Nardi 945 (NSW); 12 mi [19.3 km] S of Mt. Hope, C. W. E. Moore 5945 (CANB). VICTORIA: Nhill, G. D’Alton 26 (MEL); 15 mi [24.1 km] W of Shepparton, T. B. Muir 2547 (MEL). SOUTH AUSTRALIA: 5 km W of Mannum, D. G. Spooner 2737 (AD); Hincks Nat. Park, D. E. Symon 6495 (ADW). WESTERN AUSTRALIA: Lake Wagin, 1890, M. Cronin (MEL); 2 mi [3.2 km] NW of Ongerup, K. Newbey 3632 (PERTH). Distribution. Principally in south-eastern Australia from Eyre Peninsula eastwards; also in south-west Western Australia. Map 8. Paul G. Wilson, Tribe Chenopodieae Habitat. Usually in heavy soil; often in woodland. 155 This subspecies is variable both in habit and leaf shape and I have been unable to clearly separate the variants of which it is composed. The white spherical vesicular hairs combined with the small relatively thin leaves and prostrate to decumbent habit distinguish subsp. microphyllum from the other subspecies. Typification. The lectotype of Rhagodia prostrata is » specimen in herb. K bearing that name in Moquin’s handwriting; a fragment in herb. P (mounted with the holo- type specimen of R. drummondii Moq.) is presumably an isolectotype. The name R. prostrata was included by Bentham (1870) and Scott (1978a) as a synonym of R. spinescens R. Br., an obvious misplacement since it is certainly a synonym of C. microphyllum F. Muell. Although the name R. prostrata Moq. (1849) is earlier than C. desertorum (J. Black) J. Black, it cannot be transferred at the rank of species to the genus Chenopodium since there exists the validly published name C. prostratum (Pall.) Roemer et Schultes (1820). The varietal combination Chenopodium triandrum var. lanuginosum was evidently published inadvertently by F. E. Haviland in 1912 as a new name based on C. microphyllum F. Muell. It would appear that Haviland obtained the name from the herbarium of the Botanic Gardens, Sydney, which had received it from J. Murr as the manuscript name “forma lanuginosa” for a collection made by E. Betche in February 1886 in Dubbo (NSW 143105) and a collection from the Wimmera in Vic- toria (NSW 143082). The duplicates of these collections in herb. PAE (now in herb. G) were later cited by Aellen (1939) as syntypes of C. anidiophyllum. The name C. triandrum G. Forster was considered by Murr (1910) to be correct for the taxa here included under C. desertorum; however, the former species belongs to the genus Einadia although it bears strong superficial resemblances to members of the C. des- ertorum complex. Haviland (and others) applied the name C. triandrum var. lanuginosum to collections of C. desertorum subsp. anidiophyllum. 10c. subsp. anidiophyllum (Aellen) Paul G. Wilson, stat. nov. — C. anidiophyllum Aellen, Candollea 8:9 (1939). Lectotype: Coolabah, March 1907, J. H. Maiden (G ex herb. P. Aellen) lecto. nov. [Rhagodia parabolica auct. non R.Br.: Benth., FI. Austral. 5:153 (1870) p.p. as to Dalton specimen cited.] Erect perennial herb or weak subshrub c. 20 cm high with slender branches. Leaves thin; lamina ovate to broadly elliptic, deltoid or subreniform, 10-20 mm long, mealy below with round vesicular hairs, glabrescent above (in typical variant); peti- ole slender, 5-20 mm long. Inflorescence more or less equal to terminal leaves (typical variant) to well exceeding them. Perianth c. 1.5 mm diameter covered with simple or branched white vesicular hairs. Pericarp thin and close fitting or loose and somewhat succulent when fresh (wrinkled and orange when dry). Seed c. 1.5 mm diameter. QUEENSLAND: Tambo, S. L. Everist 1455 (BRI); Enniskillen, C. T. White 11674 (BRI). NEW SOUTH WALES: Condobolin, G. M. Cunningham 255 (NSW); Dubbo, Feb. 1886, E. Betche (NSW 143105, syntype). 156 Nuytsia Vol. 4, No. 2 (1983) SOUTH AUSTRALIA: Tinga Tingana, Strezelecki Creek, D. E. Symon 5845 (ADW). NORTHERN TERRITORY: 24 mi [38.6 km] S of Alice Springs, D. J. Nelson 666 (CANB). Distribution. The typical variant is found in north-central New South Wales and south-central Queensland; the “central Australian” variant is found in western New South Wales, northern South Australia and the southern portion of the Northern Territory. Map 9. Habitat. Varies according to variant involved. In the eastern limit of its distribution (near Dubbo) this subspecies is a weak per- ennial with small ovate leaves (lamina to 10 mm long) and a thin close indumentum on the flowers; towards the west the leaves increase in size (lamina to 15 mm long) and become more orbicular, tbe plant is more erect, and the indumentum becomes denser and thicker. There appear to be a number of local variants within the general transitional dine and, in addition, plants may be found intermediate in character be- tween subsp. anidiophyllum, subsp. microphyllum, and subsp. desertorum, particu- larly in the areas of overlap of these subspecies. In southern Queensland subsp. anidiophyllum frequently has a strong smell of stale fish (as is also found in subsp. virosum) but this is not always the case as has been noted by collectors. Several collectors also note that the plants are very palat- able to stock. The distribution map for subsp. anidiophyllum indicates a marked dis- junction between the central Australian and the eastern populations which is not reflected in the morphology of the plants from the two areas; specimens collected from north-eastern South Australia and north-western New South Wales are vir- tually identical to those from the “Centre”, that is, they have an erect habit with straight branches, and relatively large leaves and flowers, these have a thick vesicular indumentum. Further east the plants become more straggly with thinner and smaller leaves and flowers. The “central” Australian variant appears to be usually found on deep sand while the “eastern” variant is generally found on clays or loamy soils. I have included under subspecies anidiophyllum the central Australian plant which has thicker and usually larger leaves than that found in central New South Wales from where the type comes. Also included is a variant found in the northern Flinders Range with rather thick reniform leaves. Although these and other regional variants have features which give them a distinctive appearance there is intergradation and there is considerable morphological variation. An additional reason for not splitting up this complex further is the difficulty of expressing the subtle differences in a manner which can be interpreted by others. Some specimens have a habit and leaf shape intermediate between subsp. anidiophyllum and Chenopodium curvispicatum, they also have a faint reticulate or- namentation of the testa. It seems likely that these plants are the result of introgressive hybridization between the two taxa, particularly since they are found in areas where the putative parents also occur, that is in western New South Wales and in the Flinders Range. Some of the specimens referred to subsp. anidiophyllum are possibly intermediates between the subspecies desertorum and microphyllum but this cannot be recognised easily without field experience. Paul G. Wilson, Tribe Chenopodieae 157 Typification. Paul Aellen cited three syntypes with his description of Chenopodium anidiophyllum, these were as follows: “Victoria: Wimmera; no date and collector; sent by F. v. Mueller as C. microphyllum F.v.M. (Herb. Murr). New South Wales: Dubbo, 1886, E. Betche (Herb. Murr); Coolabab, 1904, J. H. Maiden (Herb. Murr).” The specimens were in a collection of chenopods sent in about 1909 to Josef Murr in Feldkirch, Austria by J. H. Maiden, Director of the Royal Botanic Gardens, Sydney. They were identified by Murr as Chenopodium triandrum Forst. f. (see Murr 1910). I have selected as lectotype the Coolabah collection since it is the only syntype present in Paul Aellen’s herbarium (now in herb. G). It is also an excellent representative of the plant generally referred in herbaria to C. anidiophyllum. The Dubbo syntype cited above belongs to this taxon but the syntype from Wimmera (NSW 143105) has greater affinity to subsp. microphyllum. Bentham (1870) included collections of this subspecies in Rhagodia parabolica (which it somewhat resembles in leaf-shape) while Scott (1978a) cited collections of subsp. anidiophyllum under both Rhagodia gaudichaudiana {- Chenopodium gaudichaudianum) and R. deltophylla (= R. spinescens). While fragments of vegeta- tive material of subsp. anidiophyllum and of the other species can be similar in ap- pearance, the flowers are very distinct; when in fruit, the subspecies cannot be con- fused. lOd. subsp. virosum Paul G. Wilson, subsp. nov. Herba perennis effusa foetida. Lamina fold ovata vel elliptica c. 10 mm longa, cbartacea, acute vel obtusa, ad basim anguste cuneata. Inflorescentia anguste pyr- amidalis, congesta; flores depresso globosi c. 1.5 mm diam. Typus: 15 mi [24.1 km] N of Deniliquin, New South Wales, Oct. 1965, W. E. Mulham S393a (holo: NSW). Spreading prostrate to decumbent, perennial herb with numerous branches arising from stock, smelling strongly of rotten fish (trimethylamine) when crushed. Root a woody taproot. Branches slender closely mealy. Leaves opposite or alternate, closely covered with grey vesicular hairs; lamina (ovate to) narrow-elliptic, c. 10 mm long, chartaceous, apex acute to obtuse, base narrowly cuneate and passing into the short petiole. Inflorescence a dense narrowly pyramidal terminal panicle not exceeding leaves, composed of few-flowered clusters subtended by linear bracts c. 2 mm long. Flowers sessile, depressed, c. 1.5 mm diam.; tepals oblong, obtuse, incurved, rounded on back and cucullate at apex, densely vesicular pubescent outside, somewhat en- larged in fruit. Terminal flower of inflorescence and of lateral clusters male or bisex- ual; filaments glabrous passing at base into a thick circular disc sparsely vesicular pubescent or glabrous. Lateral flowers female; staminodes and disc absent. Fruit en- veloped by the incurved tepals; pericarp membranous or slightly succulent, smooth, readily detached. Seed c. 1.5 mm diam., testa radially striate. NEW SOUTH WALES: Morundah — Urana, C. W. E. Moore 1372 (CANB); 15 mi [24.1 km] N of Deniliquin, W. E. Mulham S 393 (NSW); Wunnamurra, June 1920, J. Sloane (NSW). VICTORIA: Daylesford (?), 1878, R. Wallace 182 (MEL). 158 Nuytsia Vol. 4, No. 2 (1983) Distribution. South central New South Wales, possibly also in central Victoria. Map 9. Habitat. Clay soil. The arrangement and morphology of the flowers of subsp. virosum are the same as those found in the other subspecies of C. desertorum and it is with this species that the plant’s affinities obviously lie. The subsp. virosum is rather similar to subsp. micropbyllum in leaf shape although readily distinguished by its smell. This smell of trimethylamine is also present in C. vulvaria and C. detestans; subsp. virosum may be distinguished from C. vulvaria by the differently shaped leaves and perennial habit and from C. detestans by the flowers which in that species have only one or two stamens. The single herbarium record of subsp. virosum from Victoria is accompanied by a label in which the locality ‘Daylesford’ has been queried; until further collections be- come available its occurrence in that State must remain in doubt. The epithet ‘virosum’ is taken from the latin word virosus meaning foetid. lOe. subsp. rectum Paul G. Wilson, subsp. nov. Herba perennis erecta ad 40 cm alta cinereo-virida. Rami recti, divaricati, tenues. Lamina folii late elliptica ad late obovata vel orbicularis, 5-10 mm longa, ad apicem rotundata vel truncata. Inflorescentia tenuis; flores c. 1 mm diam. Typus: Porcupine Hill, Musgrave Range, South Australia, 29 Oct. 1966, J. Z. Weber 180 (holo: AD; iso: PERTH). Erect grey-green perennial herb or subshrub to 40 cm high. Branches straight, di- varicate, slender, sparsely covered when young with small discrete rounded vesicular hairs. Leaves alternate; lamina broadly elliptic to broadly obovate or orbicular, 5-10 mm long, when dry often leathery and bordered in red, apex rounded to truncate, lower surface sparsely to moderately covered with small sessile vesicular hairs, glabrescent above; petiole slender, half length of lamina. Inflorescence slender, equal to terminal leaves; perianth c. 1 mm diameter, sparsely to moderately covered with small grey round vesicular hairs. Fruit with membranous to slightly succulent peri- carp. Seed 1 mm diameter; testa smooth. NEW SOUTH WALES; 13 mi [20.9 km] SE of Mt Hope, C. R. Dunlop 1538 (CBG); 45 mi [72.4 km] NW of Balranald, W. E. Mulham W372 (NSW). VICTORIA: Hattah Lakes Nat. Park, 20 Feb. 1969, G. Anderson (MEL); Nandaly, H. B. Watts 376 (NSW). SOUTH AUSTRALIA: 5 km S of Maralinga, B. Copley 2698 (AD); Hambidge Nat. Park, T. R. N. Lothian 4270 (AD). WESTERN AUSTRALIA: 15 mi [24.1 km] E of Zanthus, R. D. Royce 5577 (PERTH). NORTHERN TERRITORY: Petermann Reserve, T. S. Henshall 764 (NT); Finke River, Schwartz and Schultz 14 (MEL). Paul G. Wilson, Tribe Chenopodieae 159 Distribution. Western New South Wales and north-western Victoria to Eyre Penin- sula, the North West Plains of South Australia and the southern portion of Northern Territory; also recorded from Zanthus in Western Australia. Map 8. Habitat. Deep sand. The subspecies rectum is distinctive in its erect habit and slender straight branches. The leaves are usually more leathery when dry than in the other subspecies and are often bordered by a reddish tinge. In spite of the wide and disjunct distribution of subsp. rectum the plant is rather uniform in habit and in flower size and morphology. At Maralinga in South Australia where it has been collected by T. R. N. Lothian (No. 4022, AD), a plant intermediate in indumentum between subsp. rectum and a Rhagodia species has also been found. This plant has a more shrubby habit than subsp. rectum and has sterile or partially sterile male flowers with a woolly disc; it appears likely that this plant (T.R.N. Lothian 4023, AD) is a hybrid between subsp. rectum and a species of Rhagodia. Also from Maralinga has been gathered a specimen (F. L. Hill 748, AD) which is intermediate in appearance between subsp. rectum and subsp. desertorum (a plant also found in this area). This ‘intermediate’ specimen is probably a hybrid between the two subspecies. The infraspecific epithet is taken from the Latin word rectus, straight, referring to the straight spikes. 11. Chenopodium curvispicatum Paul G. Wilson, sp. nov. Figure 4D. [R. spinescens R.Br. var. deltophylla F. Muell., Rep. PI. Babbage’s Exped. 19 (1859) p.p. not as to lectotype. (See note below).] Suffrutex debilis, dense farinaceus. Folia plerumque opposita; lamina deltoidea vel hastata, obtusa, 1-1.5 cm longa et lata, dense farinacea. Inflorescentia paniculata, cernua, 2-5 cm longa. Fiores globosi, extus dense farinacei, c. 2 mm diam.; peri- anthium 5-partitum in fructu divaricatum, 5 mm diam. Flores terminales masculini; stamina 5, glabra; discus patelliformis, sparse vesiculoso-pilosus; pistillodium minutum. Flores laterales feminei, staminodia et discus absens, ovarium glabrum. Fructus baccatus; pericarpium succulentum. Semen horizontale, lenticulare, margine rotundato; testa prominente reticulata, nigra. Typus: South Australia, 3 km E of Sanderston; growing on red-brown calcareous loam; straggling subshrub c. 70 cm high; fruit red; 30 Mar. 1959, R. Schodde 1082 (holo; AD; iso: CANB, duplicates distributed to CAL, K). Weak intricately branched spreading shrub c. 0.5-1 m high. Branches slender (the ends frequently nutant), densely mealy when young with vesicular hairs. Leaves op- posite or sub-opposite; lamina deltoid to hastate with obtuse to rounded angles, 1-1.5 cm long and wide; densely covered with an indumentum of white rounded vesicular hairs; petiole slender, somewhat shorter than lamina. Flowers polygamo-monoecious, sessile or shortly pedicellate in drooping pyramidal panicles 2-5 cm long; bracts lin- ear, caducous. Male (or bisexual) flowers terminal to inflorescence and to its branches; perianth 5-partite, globular, c. 2 mm diameter, densely covered outside with white stipitate vesicular hairs, glabrous within; staminal filaments linear-terete, united into a broad saucer-shaped disc which is sparsely vesicular hairy within; pis- tillode globular, usually minute but sometimes equal to a normal ovary and fertile. 160 Nuytsia Vol. 4, No. 2 (1983) with stigmas subulate, erect and usually minute. Female flowers found below male; perianth similar to that of male flower, enveloping fruit until fully mature when it hardens and opens out to 5 mm diameter, often becoming red above; staminodes ab- sent; ovary globular, glabrous, style short with a pair of slender papillose stigmas. Fruit a berry; pericarp succulent, filled with sap within, red (orange on drying). Seed lenticular with rounded margin, c. 1.5 mm diameter; testa prominently reticulate with a honeycomb matrix, black. NEW SOUTH WALES: Balranald, 7 July 1971, G. M. Alehin (NSW); 30 mi [48.3 km] W of Euston, T. R. Whaite 1838 (NSW). VICTORIA: Hattah Lakes Nat. Park, 23 March 1969, G. Anderson (MEL); Raak Salt Plain, A. C. Beauglehole 16089 (MEL). SOUTH AUSTRALIA: 38 km N of Morgan, N. N. Donner 311 (AD); 22 km SW of Renmark, Hj. Eichler 13742 (AD). WESTERN AUSTRALIA: 20 km W of Naretha, D. W. Goodall 822 (PERTH); 8 mi [12.9 km] S of Norseman, L. A. S. Johnson W182 (NSW). Distribution. Southern Australia from south western New South Wales and north western Victoria westwards to south eastern Western Australia. Map 10. Habitat. Evidently in well-drained soils often associated with limestone. This species has been confused in herbaria and literature with several other species, principally Chenopodium gaudichaudianum, C. desertorum, and Rhagodia spinescens. Mueller (1859) included specimens of C. curvispicatum (along with those of R. spinescens) when he described R. spinescens var. deltophylla (cf. collection “Desert of the Murray River”, Feb. 1847, F. Mueller, MEL 87962, a syntype of that variety); Bentham (18/0) cited the Mueller collection and at least three other speci- mens of C. curvispicatum under R. spinescens] J. M. Black (1948) included charac- ters pertaining to C. curvispicatum under both R. spinescens var. deltophylla and C. desertorum (as is also evident from his herbarium); while Scott (1978a) cited collec- tions here referred to C. curvispicatum under both R. gaudichaudiana and R. deltophylla. Chenopodium desertorum may be readily distinguished from C. curvispicatum by its smooth seed; it is also a small plant whereas C. curvispicatum is a straggly shrub. Rhagodia spinescens is a dioecious shrub with stiff straight compact inflorescences, it has a small perianth which never encloses the fruit and the berry is viscous when dry Chenopodium gaudichaudianum, while similar to C. curvispicatum in habit, has a broader more branched inflorescence; it differs most obviously in the stigma which is markedly penicillate. The specific epithet curvispicatum refers to the drooping spike-like panicles which are typical of this species. 12. Chenopodium gaudichaudianum (Moq.) Paul G. Wilson, comb, nov.— Rhagodia gaudichaudiana Moq., Chenop. Enum. 11 (1840). Type: Bale des Chiens marins, Gaudichaud (holo: P n.v.; iso: BM). Figure 7F. R. coralliocarpa S. Moore, J. Bot. 35:168 (1897). Type: Kilkenny Soak, June 1895, S Moore (holo: BM). Paul G. Wilson, Tribe Chenopodieae 161 Intricately branched scrambling shrub to 2 m high. Branchlets, leaves and inflor- escence densely mealy with discrete cream-coloured vesicular hairs. Leaves alternate, chartaceous (when dry), deltoid to narrow hastate with slender petiole 3-5 mm long, in all 10-20 mm long. Inflorescence a pyramidal panicle 5-10 cm long with disjunct clusters of flowers, the main axis and branches flexuose. Flowers polygamo-mon- oecious, subsessile, those terminating the inflorescence or its branches male (with small pistillode) or bisexual, the others female. Perianth 5-partite, depressed globular c. 1.5 mm diam. at an thesis and 3.5 mm in fruit, eventually spreading and c. 7 mm diameter, persistent, densely covered outside with vesicular hairs. Bisexual flowers: stamens 5, filaments linear-terete, united into a broad saucer-shaped disc which is glabrous or vesicular puberulous; ovary depressed globular, style slender 0.5 mm long with a pair of delicate strongly penicillate stigmas c. 0.6 mm long. Fe- male flowers without staminodes. Fruit a berry, at first enveloped by perianth but which eventually becomes spreading; pericarp pale and membranous or orange to red and succulent with sap filled interior (not viscous), eventually falling from perianth. Seed lenticular c. 1.4 mm diameter with rounded margin; testa deeply reticulate with a rough surface, dark reddish brown. SOUTH AUSTRALIA: Yudnapinna HS., N. N. Donner 3205 (AD); Tarcoola, E. H. Ising 1408 (AD); Maralinga, 21 Sept. 1960, H. Turner (AD). WESTERN AUSTRALIA: Roebourne, Aug. 1884, F. S. Carey (MEL); Narrine, Sept. 1903, W. V. Fitzgerald^iNSW); 23 mi [37 km] S of Learmontb, A. S. George 1258 (PERTH); Miss Gibson Hill, SW of Warburton, A. S. George 4043 (PERTH). Distribution. Karratha to Geraldton in Western Australia, south-east to the northern edge of the Nullarbor Plain, eastwards to northern Eyre Peninsula. Map 1. Habitat. Sand, clay or loam, often in slightly saline situations. This species was recorded by Bentham (1870) and by Maiden (1916) as occurring in New South Wales but the collections on which these records were based are of other species, principally C. curvispicatum. Scott (1978a) recorded it from central and eastern South Australia and from New South Wales, but the relevant collections are either C. desertorum or C. curvispicatum. The strongly penicillate stigmas of C. gaudichaudianum clearly distinguish it from all other species of Chenopodium and all species of Rhagodia, while its scrambling habit with flexuose branches make it readily recognisable in the field. It is uniform in habit and morphology throughout its range and does not appear to hybridize with other species. The male flowers of Chenopodium gaudichaudianum have a widely spreading peri- anth at anthesis but this subsequently closes, while in female flowers the perianth re- mains closed during anthesis, with only the stigmas protruding, then opens widely under the mature fruit. Sectio Auricoma Aellen, Feddes Repert. 69:69 (1964); Aellen in Hegi, 111. FI. Mitt.- Eur. ed. 2, 3:577 (1960) nom. illeg. (without latin diagnosis). Type: C auricomum Lindl. Herbs or soft-wooded shrubs, mealy with vesicular hairs and (at least in the inflor- escence) villous with tubular hairs. Leaves chartaceous. Flowers in compact glomerules forming dense cylindrical spikes paniculately arranged, bisexual or fe- male by abortion of stamens; tepals 5, united at first in lower half; stamens 5, fila- Nuytsia Vol. 4, No. 2 (1983) ments linear, disc absent; ovary glabrous, styles very short, stigmas 2 and slender. Pericarp papery. Seed horizontal (rarely oblique or erect), lenticular; embryo annu- lar. Two species endemic to Australia. This section is very similar to sect. Leprophyllum particularly with the addition to sect. Auricoma of C. auricomiforme which forms a connecting link between them, as has been noted by Murr et Thellung (1915). It differs from sect. Leprophyllum prin- cipally in the presence of tubular hairs in the inflorescence. 13. Chenopodium auricomiforme Murr et Thellung, Viertiljahrsschr. Naturf. Ges. Zurich 60:432 (1915); Aellen in Hegi, 111. FI. Mitt.-Eur. ed. 2, 3:615 (1960). Type: ‘Kammgarnfabrik Derendingen bei Solothurn, auf Kompost von australischer Schafvolde’, 1915, R. Probst (holo: G). Figure 4c. [C. album auct. non L.: Benth., FI Austral. 5:160 (1870) p.p. as to Barton specimen cited.] Erect annual herb to 1 m high. Stem and branches rather thick, striped green and yellow, sparsely mealy when young. Leaves chartaceous; lamina 3-nerved, narrowly ovate to ovate, rarely hastate (upper ones lanceolate), entire or sparsely and coarsely toothed, 2-10 cm long, weakly cuspidate, glabrescent above, mealy below when young. Inflorescence a terminal panicle with slender densely flowered branches made up of compact glomerules subtended by filiform bracts, axis villous with tubular hairs. Flowers bisexual (or female by abortion of stamens), sessile or subsessile, spherical, c. 1 mm diam.; perianth almost glabrous (a few vesicular and tubular hairs), (4)5-lohed to middle or lower (often initially syntepalous becoming choritepalous with age), lobes (or tepals) imbricate, broadly elliptic to almost circu- lar, sometimes abruptly constricted at base into a short claw, yellow-green with fleshy keel and scarious margins, ciliate; stamens 5; filaments linear, glabrous; disc absent. Ovary glabrous; style very short; stigmas 2, slender. Fruit free from perianth; pericarp membranous, papillate, pale brown, adherent. Seed horizontal, lenticular, c! 1 mm diameter, rounded on margin; testa minutely puncticulate. QUEENSLAND: Minerva, S. T. Blake 7927 (BRI); Pittsworth, C. T. White 6659 (BRI). WALES: Bow via Merriwa, July 1923, H. Sturt (NSW); Premer, June 1912, H. McMaster (NSW). SWITZERLAND: Derendingen, 12 Aug. 1927, P. Aellen (NSW). Distribution. South-eastern Queensland and north-eastern New South Wales- a cas- ual in Europe. Map 11. ’ Habitat. Black soil plains and heavy loams, often in low-lying situations. This species was first recognised by Murr and Thellung who discovered it in Switzerland as an introduction with wool from Australia. Aellen (1960-1961) con A to be also a native of New Zealand but it is not recorded by Allan (1940 and 1961) as either native or introduced in that country nor is it present in the D.S.I.R. herbarium in Christchurch, N.Z. (CHR). Murr and Thellung (1915) corisidered it to be intermediate in morphology between C. album and C. auricomum and subsequently Murr (1926) suggested it might be a hybrid between these species. Paul G. Wilson, Tribe Chenopodieae 163 Both species were confused with C. auricomiforme by previous authors for Bentham (1870) cited specimens under C. album and F. Mueller (1869 p. 11) under C. auricomum. The earliest record is a pre 1848 collection by L. Leichhardt from Liverpool Plains (MEL 88487). Chenopodium auricomiforme is clearly a distinct species and shows very little variability throughout its range. Chenopodium auricomiforme was placed by Aellen (1960) in section Polygonoidea along with C. trigonon, C. stellulatum, C. polygonoides, and C. hubbardii. However, it differs greatly from the first three species both in habit and in number of stamens. It compares most closely to C. auricomum, particularly in hair and perianth charac- ters, and I have therefore transferred it to sect. Auricoma. 14. Chenopodium auricomum Lindley in Mitchell, J. Exped. Int. Trop. Austral. 94 (1848); Moq. in DC., Prod. 13/2:460 (1849); Benth., FI. Austral. 5:159 (1870); Aellen in Hegi, 111. FI. Mitt.-Eur. ed. 2, 3:616 (1960). Type: Collected along the Narran R., 14 March 1846, T. L. Mitchell (holo: CGE n.v.; ?iso: MEL, NSW). ?C. auricomum f. subglabrum Aellen in Probst, Mitt. Naturf. Ges. Soloth. 20 (8):56 (1928); Aellen in Hegi, l.c. Type: Derendingen, Switzerland, n.v. Soft-wooded bluish grey shrub to 2 m high. Branches slender, costate, thinly mealy. Leaves alternate, oblong-elliptic to hastate, 2-4(6) cm long including a slen- der petiole c. 1 cm, entire, chartaceous, apex and basal lobes rounded, upper surface glabrescent, lower surface with a thin mealy indumentum of grey or very pale yellow discrete globular vesicles. Inflorescence grey mealy to very pale yellow in a broad ter- minal panicle c. 10 cm long with ovoid to cylindrical branches composed of dense 3-5 flowered glomerules subtended by small spathulate bracts; axis pubescent with tubular hairs. Flowers bisexual, or female by abortion of stamens (sometimes with minute staminodes), sessile, depressed globular, c. 1 mm diam.; perianth united in lower 1/2 to 2/3, entirely pubescent outside (or only in distal half) with irregularly shaped short-tubular and bladder hairs, lobes imbricate, broadly obovate, membra- nous, ciliolate; stamens 5, filaments linear and neither thickened at base nor united into a disc; ovary globular, glabrous, style very short, stigmas slender. Fruit envel- oped by perianth and deciduous with it; pericarp chartaceous, free, white and opaque. Seed horizontal (rarely oblique or erect), lenticular, c. 1.5 mm diam., margin rounded; testa minutely reticulate. QUEENSLAND: Cunnamulla, S. T. Blake 10629 (BRI); Noondoo Stn., S. L. Everist 814 (BRI). NEW SOUTH WALES: Depot Glen, S. Jacobs 3076 (NSW); Mt King Stn., R. A. Perry 5m (NSW). SOUTH AUSTRALIA: 48 km NE of Billa Kalina Stn., M. Fagg 379 (AD); Murnpeowie Stn., D. E. Symon 5653 (ADW). WESTERN AUSTRALIA: DeGrey River district, N. T. Burbidge 1585 (PERTH): Mt Augustus Stn., 8 July 1970, D. G. Wilcox (PERTH). NORTHERN TERRITORY: Supplejack Stn., T. S. Henshall 2306 (NT); Mt Andado Stn., T. S. Henshall 601 (NT). Distribution. Western Queensland, central and western New South Wales, northern South Australia, northern Western Australia, Northern Territory. A casual in Europe. Map 11. 164 Nuytsia Vol. 4 , No. 2 (1983) Habitat. Clay soil and heavy loam in depressions and areas of periodic flooding. This species was placed by both Bentham (1870 l.c.) and Ulbrich (1934) in the same section as C. album, but Aellen (1960 l.c.) removed it to the monotypic section Auricoma, in which decision he was followed by Scott (1978b). The shrubby nature of C. auricomum as well as the presence of tubular hairs suggests that it should be considered sectionally distinct from the annual species related to C. album. Bentham’s (1870) suggestion that C. auricomum might be conspecific with C. furfuraceum Moq. (1840) and C. acuminatum Willd. (1799) is unacceptable since Chenopodium furfuraceum is a synonym of Rhagodia candolleana Moq. (1840) q.v., while C. acuminatum is an annual species with a sharply keeled margin to the seed, it is native to Asia and not found in Australia. Owing to its palatability to stock this species has virtually disappeared from large areas of the pastoral country. It is also edible to humans and Max Koch (1895 in sched.) records it as being eaten by Aborigines in western New South Wales. Sectio Rhagodioides Benth., FI. Austral. 5:158 (1870); J. D. Hook, in Benth. et J. D. Hook., Gen. PI. 3:52 (1880); Volkens in Engler et Prantl, Nat. Pflanzenfam. III. la:61 (1892); Ulbrich in Engler et Prantl, Nat. Pflanzenfam. ed. 2, 16c:489 (1934); A. J. Scott, Bot. Jahrb. Syst. 100:214 (1978). Type: C. nitrariaceum (F. Muell.) Benth. Spinescent shrubs with a thin mealy indumentum which eventually collapses and coalesces to form a scaly silvery sheen. Flowers sessile in a spinescent panicle, uni- sexual or bisexual; tepals 5 united towards base, enveloping mature fruit; ovary pu- bescent; stamens 5, glabrous; disc absent. Pericarp membranous, pubescent. Seed horizontal to vertical; embryo annular. A monotypic section indigenous to Australia. This section differs from sect. Auricoma principally in the spinescent habit, the weak mealy indumentum which becomes scaly with age, and the pubescent pericarp, characters which, individually, are indefinite in their application but which together demarcate C. nitrariaceum as a very distinct taxon. Scott (1978b) included in this section Rhagodia ulicina (Gand.) Paul G. Wilson (as a species of Chenopodium), however, R. ulicina is a dioecious plant with a fleshy pericarp and was originally placed in the genus Chenopodium only through Gandoger’s ignorance of its morphology. 15. Chenopodium nitrariaceum (F. Muell.) F. Muell. ex Benth., FI. Austral. 5:158 (1870) ‘nitrariacea’; F. Muell., Icon. Austral. Salsol. PI. t. 28 (1890); S. J. Oostroom et T. J. Reichgelt, Gorteria 1:21 (1962). — Rhagodia nitrariacea F. Muell., Trans. Phil. Inst. Viet. 2:72 (1858); F. Muell. Sec. Gen. Rep. 14 (1854) nomen. Lectotype: Upper left-hand specimen on sheet bearing labels 1) ‘Ad ripam fluvii Murray’, Feb. 1847, and 2) ‘Avoca’, 2 Dec. 1852, both leg. F. Mueller (MEL 80673), see note below, lecto. nov. C. lycioides Gand., Bull. Soc. Bot. France 66:224 (1919). Type: Warracknabeal, Vic- toria, 8 Oct. 1903, F. M. Reader (holo: LY). Paul G. Wilson, Tribe Chenopodieae 165 Divaricately branched spinescent shrub c. 1 m high. Branches slender, covered with a grey metallic sheen due to fusion of collapsed vesicular hairs. Leaves alter- nate, frequently clustered at nodes; lamina coriaceous, spathulate with rounded apex passing at base into a short petiole, in all 10-30 mm long, sparsely mealy with minute branched collapsed vesicular hairs. Inflorescence a terminal panicle with spinescent branches to 15 cm long and wide or reduced to a single branchlet. Flowers subtended by small linear to ovate bracts, clustered in irregular glomerules, unisexual and di- oecious or bisexual. Perianth depressed globular to broadly turbinate, 1.5-2 mm di- ameter; tepals 5, united in lower half, pubescent outside with small branched tubular hairs. Male flowers; stamens 5; filaments ligulate, glabrous; disc absent; pistillode c. 1 mm long, with minute erect stigmas. Female flower: staminodes minute c. 0.1 mm long or absent; ovary pubescent above with branched tubular hairs; style short or ab- sent; stigmas 2, slender, to 1.5 mm long, pilosulose. Fruit enveloped by perianth, dry; pericarp membranous, white, pubescent above, free. Seed horizontal to vertical, de- pressed globular or concavo-convex, to sub-reniform, c. 1 mm diam.; testa punctate, reddish-brown to black, glossy; embryo hippocrepiform to annular. QUEENSLAND: Noondoo Stn., S. L. Everist 831 (BRI). NEW SOUTH WALES: Bourke, May 1918, J. L. Boorman (NSW); 8 mi [12.9 km] SW of Tulpa, C. W. E. Moore 5433 (CANB). VICTORIA; Dimboola, 1947, E. T. Muir (MEL); Borung, 13 Nov. 1903, F. M. Reader (MEL). SOUTH AUSTRALIA: 3 km W of Hawker, B. Copley 218 (AD); 48 km W of Tallaringa Well, T. R. N. Lothian 3847 (AD). NORTHERN TERRITORY: 7 mi [11.3 km] E of Docker R. Settlement, P. K. Latz 968 (AD); 12 mi [19.3 km] SE of Tempe Downs HS., N. M. Henry 585 (NT). Distribution. Central Australia south eastwards to north western Victoria, central and western New South Wales, south central Queensland. A casual in Switzerland, Germany, Holland and England. Map 15. Habitat. Clay soil in seasonally waterlogged situations. This species exhibits considerable variation in the shape of the flowers and in the nature of the stamens and ovary. In some plants the flowers are either all male (with a very small pistillode) or all female, in other plants the apparently male flowers have well-developed pistillodes which sometimes form seed (but the stigmas remain miriute), while in yet others the flowers are bisexual. The orientation of the seed also varies. I have not been able to find characters by which the several variants that ap- pear to be involved could be separated, partly because the herbarium material is usually imperfect. Typification. Mueller (1858) did not cite specimens with his description of Rhagodia nitrariacea but he gave as its distribution ‘Throughout the interior of Australia, from Arnhem’s Land to Lake Torrens, the Murray River and its tributaries’. The only type material in herb. MEL is of four specimens mounted on one sheet (MEL 80673) on which are two labels indicating that the specimens were collected along the Avoca River on 2 December 1852 and along the Murray River in February 1847. I have chosen the upper left-hand specimen as the lectotype but there is no indication as to which label it relates. A syntype in herb. K was collected by Mueller along 22493 — ( 3 ) 166 Nuytsia Vol. 4, No. 2 (1983) Sturts Creek, Western Australia, presumably in 1856. This had been identified as R. nitrariacea by Mueller but was cited by Bentham (1870) under R. crassifolia; it is a specimen of R. eremaea. Chenopodium nitrariaceum bears a superficial resemblance to Rhagodia ulicina but the two differ markedly in hair type and in flower and fruit characters. Subgenus Ambrosia A. J. Scott, Bot. Jahrb. Syst. 100:211 (1978) Type: C. ambrosioides L. Sectio Ambrina J. D. Hook, in Benth. et J. D. Hook., Gen. PI. 3:51 (1880). Type: C. multifidum L. Ambrina Spach, Hist. Nat. Veg. 5:295 (1836). nom. illeg. (includes type of Roubieva). Type: C. multifidum L. Roubieva Moq., Ann. Sci. Nat. Bot. ser. 2,1:292 (1834). — Chenopodium sect. Roubieva (Moq.) Asch. et Graebner, Syn. Mitteleur. FI. 5/1:21(1913). Type: C. multi- fidum L. Annual or perennial herbs with yellow glandular hairs, aromatic. Leaves herbaceous, entire to pinnatifid. Flowers in dense axillary glomerules, bisexual and female. Perianth united in lower half; stamens 3-5; stigmas 2-5. Pericarp membra- nous, covered in part with geniculate glandular hairs. Seeds horizontal to vertical; embryo hippocrepiform to sub-annular. About 8 species from South and Central America some of which are now cosmo- politan weeds. Two species are found in Australia. The sect. Ambrina has been by some authors (e.g. Ascherson et Graebner 1913, Aellen 1960-1961, Scott 1978b) lectotypified on the species C. ambrosioides. How- ever, Spach (1836 op. cit.) when describing the genus Ambrina, included in it both C. multifidum L. (as A. pinnatisecta Spach) and C. ambrosioides (among other species). Moquin (1840) while accepting the genus placed C. ambrosioides in Ambrina sect. Adenois (which he stated to be the typical section) and C. multifidum in the genus Roubieva. The latter name was based on Roubieva Moq. (1834) with the one species R. multifida (Chenopodium multifidum L.). The generic name Ambrina is therefore illegitimate being superfluous and is to be typified by the type of the name which should have been adopted, i.e. C. multifidum. J. D. Hooker (1880) clearly based the sectional name Ambrina on Spach’s generic name and gave it the same broad cir- cumscription. I consider that C. ambrosioides and C. multifidum belong to the same section so the effect of a lectotypification different to that of other recent authors (who have selected C. ambrosioides) is not significant. I have accepted the name Chenopodium sect. Ambrina in the sense of J. D. Hooker (1880). It was first used by J. D. Hooker in Flora Novae-Zelandiae 213 (1853) with- out reference to an author or to a basionym. He included two species, C. carinatum R.Br. and C. pumilio R.Br., and gave as a description ‘seed vertical’. The name if treated as new is illegitimate, being superfluous since it includes the lectotype of the section Orthosporum R.Br. (1810). Hooker probably intended it to be based on the genus Ambrina and was following Moquin (1840) who included both C. carinatum Paul G. Wilson, Tribe Chenopodieae 167 and C. pumilio in Ambrina sect. Adenois (the typical section of that genus). It is de- batable whether Hooker’s two word description in his 1853 publication is acceptable as a diagnosis, particularly since his description for the unnamed group, which con- tained the rest of the species he placed in the genus, was ‘seed horizontal, rarely (in C. glaucum sometimes) vertical.’ This being the case, and in order to retain current nomenclature where possible, I am treating the 1853 publication of the name as being invalid. Chenopodium sect. Ambrina therefore dates from 1880 and is based on Ambrina Spach. The species of section Ambrina are peculiar in having geniculate hairs on their ovaries. This character, along with their distinctive smell, clearly distinguish them from all other Chenopodium species found in Australia. 16. Chenopodium ambrosioides L., Sp. PI. 219 (1753). Type citation: ‘Habitat in Mexico, Lusitania’. Type: n.v. Perennial herb c. 1 m high with ascending branches, strongly aromatic, glabrous or villous towards apex. Leaves herbaceous; lower ones elliptic to narrowly elliptic, coarsely serrate or incised, narrowed at base into a short petiole, in all c. 10 cm long; upper ones entire and smaller; under-surface prominently covered with sessile globular amber-coloured oil-filled hairs. Flowers sessile in small glomerules forming leafy or leafless panicles, sometimes spicate. Terminal flower of glomerule bisexual; perianth 5-lobed to near base, glabrous, lobes cucullate and enclosing flower; stamens 5, glabrous, filaments ligulate, anthers exserted; ovary pubescent in upper half with shortly stipitate geniculate glandular hairs, style short with 3-4 slender stigmas. Lateral flowers female; perianth depressed spherical shortly 5-toothed and enclosing flower. Pericarp membranous, free, glandular pubescent above Seed hori- zontal or oblique, obtusely lenticular, c. 1 mm diam., almost smooth, glossy; embryo hippocrepiform. QUEENSLAND: Moreton Island, L. Durrington 1464 (BRI); Carnarvon Gorge, T. J. McDonald 215 (BRI). NEW SOUTH WALES: Mt Harris Station, 1 May 1952, E. F. Constable (NSW); Scarborough Park, Koyarah, E. F. Constable 5940 (NSW). VICTORIA: Recorded for eastern Victoria by Willis (1973), no specimens seen. SOUTH AUSTRALIA: Paradise, Adelaide, A. G. Spooner 3973 (AD). WESTERN AUSTRALIA: Fremantle, P. G. Wilson 8687 (PERTH). Distribution. Widespread in mainland Australia. Cosmopolitan, native of tropical America. Map 10. Habitat. Disturbed situations, often bordering streams or creeks. I have not attempted to distinguish in this paper the various closely related species recognised by Aellen (1960), nor have I considered the infraspecific taxa enumerated by Aellen (1929) and by Aellen and Just (1943). Much of the Australian material agrees well with C. anthelminticum L. as delimited by Aellen (C. ambrosioides var. anthelminticum (L) A. Gray), and a few specimens from Rockhampton, Queensland, correspond to Aellen’s description (1960-1961) of C. spathulatum Sieber (C. ambrosioides f. spathulatum (Sieber) Aellen 1929) which he records from Sydney. 168 Nuytsia Vol. 4, No. 2 (1983) The former species is distinguished by its slender spike-like inflorescence branches in which the bracts of the flower clusters are very small or absent, while the latter species is recognised by its spathulate bracts. Comments on this species complex by C. F. Reed in Flora of Texas (1969) and by J. A. Duke in Flora of Panama (1961) ap- pear to be apposite to the situation in Australia; Duke remarks on C. ambrosioides: ‘polymorphically perplexing, this species, or some of its subspecies, varieties or forms, is quite cosmopolitan’, while Reed, after commenting on two varieties, states that: ‘combinations of all characters vary to a degree that separating them is very difficult’. Voroschilov (1942) and Aellen (1960-1961) recognised the following three taxa of the C. ambrosioides complex as being present in Australia: C. integrifolium Vorosch., J. Bot. URSS 27:42 (1942) n.v.; Aellen in Hegi, 111. FI. Mitt.-Eur. ed.2, 3:596 (1960). C. spathulatum Sieber ex Moq. in DC., Prod. 13/2:73 (1849); Aellen in Hegi, loc. cit. C. suffruticosum Willd. subsp. remotum Vorosch. op. cit. n.v.; Aellen in Hegi, op. cit. 595. I have not attempted to discriminate any of the above taxa. This species is cultivated in some countries for its oil which is considered to have anthelmintic properties while the leaves are used for making the drink known as Mexican tea. The variety antbelminticum is also cultivated and its oil, as the varietal epithet suggests, has a vermifugal effect, which is more pronounced than that of the typical variety. An account of the oil content of the plant found in Australia is given by Shapter (1941). 17. Chenopodium multifidum L., Sp. PI. 220 (1753). — Roubieva multifida (L) Moq., Ann. Sci. Nat. Bot. ser. 2, 1:292 D834). — Ambrina pinnatisecta Spach, Hist. Nat. Veg. 5:296 (1836). Type citation: ‘Habitat in Bonaria’ (Buenos Aires). Type: n.v. C. multifidum f. spathulatum Aellen, Amer. Midi. Naturalist 30: 50 (1943). Type not cited. Perennial herb, prostrate, decumbent or erect, to 1 m high, much branched from the base, aromatic. Branches sparsely to moderately covered with cottony hairs. Leaves pinnatisect, to 6 x 3 cm; segments linear with recurved margins, under sur- face densely covered with pale yellow glandular hairs and sparsely so with cottony hairs. Flowers predominantly female, sessile in few flowered glomerules forming spicate panicles. Bisexual flower: perianth hemispherical, shortly 5-lobed; stamens 5, filaments ligulate, glabrous; ovary narrowly turbinate, with shortly stipitate amber- coloured geniculate glandular hairs in upper half; stigmas 3-5, sessile, slender. Fe- male flower: perianth obovoid (pouch-like), minutely lobed and enclosing pistil. Fruiting perianth c. 2.5 mm long, coriaceous and reticulate-veined. Pericarp mem- branous, tree. Seed erect, lenticular with rounded margin; testa reddish brown, glossy, puncticulate; embryo hippocrepiform. QUEENSLAND: Near Warwick, Nov. 1958, Shirley (BRI). NEW SOUTH WALES: East Tamworth, R. H. Goode 226 (NSW); Hay, Feb. 1970. B. M. Alehin (NSW). Paul G. Wilson, Tribe Chenopodieae 169 VICTORIA: Seymour, R. V. Smith 64/105 (MEL); Coode Is., 23 Mar. 1912, J. R. Tovey (MEL). SOUTH AUSTRALIA: Pt. Adelaide, D. E. Symon 1233 (ADW); Semaphore, 19 June 1940, C. G. Samuel (ADW). WESTERN AUSTRALIA: Bunbury, Sept. 1942, C. A. Gardner (PERTH); Harvey, Dec. 1967, K. S. Cole (PERTH). Distribution. Southern mainland Australia. Native of South America and now widely distributed. Map 12. Habitat. Disturbed areas; often in coastal sand. The branches of some specimens have the upper leaves entire and spathulate and it was evidently on this variant that the f. spathulatum Aellen (1943) is based. As will be noted in the sectional synonymy this species is the type of the names Chenopodium sect. Ambrina and Chenopodium sect. Roubieva. These two sections have been considered distinct by some authors (e.g. Aellen 1960-1961, Scott 1978b) with C. ambrosioides (L.) Spach accepted as the type of sect. Ambrina and C. multi- fidum of sect. Roubieva. However, even if this typification is adopted, the two sec- tions appear to be not deserving of separate status since sect. Roubieva is then only distinguished by its erect seeds and pouch-like perianth, characters which appear to be rather insignificant compared with those the two sections have in common, par- ticularly the glandular leaves and the curious geniculate hairs on the ovary. Sectio Orthosporum R.Br., Prod. 407 (1810); Benth., FI. Austral. 5:162 (1870); J. D. Hook, in Benth. et J. D. Hook., Gen. PI. 3:51 (1880) see note 1 below; Volkens in Engler et Prantl, Nat. Pflanzenfam. 3, la:61 (1892); Aellen, Verb. Naturf. Ges. Basel 44:308 (1933); Ulbrich in Engler et Prantl, Nat. Pflanzenfam. ed 2, 16c:494 (1934); Aellen in Hegi, 111. FI. Mitt.-Eur. ed. 2, 3:576 (1960); Scott, Bot. Jahrb. Syst. 100:213 (1978). Lectotype: Chenopodium carinatum R.Br. (see Ulbrich 1934). See note 2 below. Blitum sect. Orthosporum (R.Br.) C. Meyer in Ledeb., FI. Alt. 11 (1829); Moq. in DC., Prod. 13/2:81 (1849). See note 3 below. Orthosporum (R.Br.) C. Meyer ex T. F. L. Nees, Gen. PI. FI. German, ad t. [58] (1835). See note 4 below. Orthospermum (R.Br.) Opiz, Seznam rostlin Kveteny Ceske 70 (1852). See note 5 below. Chenopodium sect. Ambrina J. D. Hook., FI. Nov.-Zel. 213 (1853) nom. invalid, non J. D. Hook. (1880). Type; C. carinatum R.Br. Chenopodium sect. Carinata Standley, N. Amer. FI. 21/1:11, 27 (1916) nom. illeg. (superfluous name). Type: Chenopodium carinatum R.Br. Note 1: J. D. Hooker (1880) and Volkens (1892) included Chenopodium tenue Colla from Chile in the section Orthosporum. However, this species is conspecific with C. multifidum L., a member of section Ambrina. Note 2: A. J. Scott (1978b) nominated Chenopodium pumilio R.Br. as the lectotype of section Orthosporum but since the choice of lectotype had already been made by Ulbrich (1934) this earlier selection stands. 170 Nuytsia Vol. 4, No. 2 (1983) Note 3: C. A. Meyer (1829 op.cit.) described Blitum sect. Orthosparum without reference to the name Chenopodium sect. Orthosporum R.Br. He did, however, in- clude in his section the two species described by Brown under sect. Orthosporum and therefore it can be assumed that the names are nomenclatural synonyms. Moquin (1840) recognised Meyer’s section but excluded both the species described by Brown; instead he placed them in Ambrina sect. Adenois. This error was corrected in a later publication (Moquin 1849). Note 4; The generic name Orthosporum (R.Br.) C. Meyer ex T. Nees (1835 op.cit.) was based on Blitum sect. Orthosporum (R.Br.) C. Meyer (1829 op.cit.) but without any mention of Robert Brown or of the two species Brown had placed in Chenopodium sect. Orthosporum. A. J. Scott (1978b p. 209) considered the generic name Orthosporum to be illegitimate and gave as type O. rubrum (L.) T. Nees ( = Chenopodium rubrum L.); this was one of the two species named and illustrated by Nees under Orthosporum, the other being O. bonus-henricus (L.) T. Nees (= C. bonus-benricus L.). However, since the generic name Orthosporum is indirectly based on Chenopodium sect. Orthosporum R. Br. it has the same type which is C. ca- rinatum R.Br. The name Orthosporum is therefore legitimate and correct should the “carinatum group” be considered to warrant separate generic status. Note 5: The generic name Orthospermum w'as attributed by Opiz (1852 l.c.) to Robert Brown and he gave as a synonym “Orthosporum C. A. Meyer”. It would therefore appear that the first name is an orthographic variant of the second, and that it was based on Brown’s sectional epithet either directly, or indirectly through Meyer’s combination under Blitum, or through the generic name Orthosporum C. Meyer ex T. Nees. Annual herbs, prostrate, ascending, or erect, pilose with segmented hairs and with sessile or stipitate gland-tipped hairs, strongly aromatic. Flowers ebracteate, small in dense axillary clusters (condensed thyrses) or paniculate (C. saxatile), bisexual and feniale. Tepals 4-5, erect, united towards base, variously thickened and enlarged in fruit; stamens 0-1(2) opposite anterior or (in 4-merous flowers) anterior-lateral sepal; stigmas 2, subsessile, slender. Pericarp membranous. Seed erect, lenticular; funicle very short; embryo hippocrepiform around apex of seed; radicle inferior. Fruit de- ciduous with perianth, an abcission zone forming at apex of pedicel. Distribution. Australia and New Zealand; almost cosmopolitan through recent introduction. Seven species, six of which are endemic to Australia and one to New Zealand. The habit, hair types, and inflorescence structure of the species in this section are similar to those found in Dysphania section Tetrasepalac, the principal differences between these sections of the two genera being found in the seeds. In Dysphania sect. Tetrasepalae the seed is globular or subglobular and the embryo either lateral or basal. In Chenopodium sect. Orthosporum the seed is lenticular and the embryo always hippocrepiform around its apex. Hybridisation commonly occurs between the species of this section (apart from C. pusillum and C. saxatile) and several names have been given to the crosses or backcrosses; these are listed below with critical comments. Since the species them- selves exhibit considerable variability the number of different variants is immense and it is pointless to attempt to name them. Where possible, herbarium specimens of apparetit hybrid origin have been designated by a formula, but since parentage is often difficult to determine (unless a suite of specimens has been collected) often only the more obvious parent has been indicated. Paul G. Wilson, Tribe Chenopodieae 171 In addition to the complexities noted above it is apparent that intergradation is present between at least some of the pairs of species, in particular, the following: 1. C. pumilio — C. carinatum 2. C. carinatum — C. melanocarpum 3. C. cristatum — C. melanocarpum These are discussed below. Names of hybrids — sect. Orthosporum A. Chenopodium xbontei Aellen, Verb. Naturf. Ges. Basel 44:317 (1933). Considered by Aellen to be a hybrid between C. carinatum and C. cristatum which, with the hybrid, were found at Derendingen in Switzerland. Aellen divided it into two varieties as follows: nothovar. cristatiforme Aellen, op. cit. p. 318. Lectotype: Derendingen, Switzerland, 9 Sept. 1923, R. Probst (G) lecto nov. Syntype: Luterbach, Switzerland, 16 Aug. 1933, R. Probst (G). Stated by Aellen to be a hybrid between C. carinatum var. holopterum (i.e. var. ca- rinatum) and C. cristatum. The two specimens cited above appear to have been de- rived from C. cristatum and another species, possibly C. carinatum. nothovar. submelanocarpum Aellen, op. cit. Lectotype Derendingen, Switzerland, 5 Sept. 1926, R. Probst (G) lecto nov. Syntype: Kettwig, Germany, 5 Oct. 1930, L. Bonte (G). Stated by Aellen to be a hybrid between C. carinatum var. melanocarpum (= C. melanocarpum) and C. cristatum. The two specimens cited appear to have had such an origin. B. Chenopodium xchristii Aellen, op cit. p. 317. Considered by Aellen to be a hybrid between C. carinatum var. holopterum (i.e. var. carinatum) and C. pumilio. Aellen divided it into two varieties as follows: nothovar. intermedium Aellen, op. cit. 317. Type: “Western Australia: 120 km south of Perth, 1901, leg. Cecil Andrews, 1th collect, no. 697”. (holo: K n.v.). The description and illustration provided by Aellen suggest that this is a variant of C. pumilio. nothovar. semiconnatum Aellen, op. cit. 317. Type: “In Nova Holandia ad portum Jackson, 1830, leg. Gaudichaud” (holo: G n.v.). The Gaudichaud specimen is one of the two syntypes of Ambrina carinata var. parvifolia Moq. (1840). It is probably a variant of C. pumilio. C. Chenopodium trigonocarpum Aellen, op. cit. 41:99 (1930). Lectotype: Derendingen, Switzerland, 31 July 1929, R. Probst (G) lecto. nov. 172 Nuytsia Vol. 4, No. 2 (1983) In addition to the lectotype, Aellen cited one other specimen from Derendingen (not seen) and stated that it was also known to him from New Zealand and Australia. In a later paper Aellen (1933) placed C. trigonocarpum in synonymy under C. ca- rinatum var. holopterum (= var. carinatum). The lectotype appears to be a hybrid between C. melanocarpum and probably C. cristatum or between C. melanocarpum and another hybrid involving C. cristatum. 18. Chenopodium pumilio R.Br., Prod. 40 (1810); Benth., FI. Austral. 5:163 (1870); Aellen, Verb. Naturf. Ges. Basel 44:314 (1933).— Bbtum pumilio (R.Br.) C. A. Meyer ex Steudel, Norn. Bot. ed. 2, 1:210 (1840); Moq. in DC., Prod. 13/2:82 (1849).— Ambrina pumilio (R.Br.) Moq., Chenop. Monog. Enum. 42 (1840). Type: Kangaroo Island, R. Brown (holo: BM n.v.). Blitum glandulosum Moq. in DC., Prod. 13/2:82 (1849) sensu lectotypica fide Aellen (1933).— Chenopodium glandulosum (Moq.) F. MuelL, Fragm. 7:11 (1859). C. pumilio f. glandulosum (Moq.) Aellen, Verb. Naturf. Ges. Basel 44:315 (1933); Aellen in Hegi, 111. FI. Mitt.-Eur. ed. 2,3:598 (1960). Type citation: “In Nova-Hollandia” “(v.s. in h. Desf., Poir., DC. et Webb.)”. Type: n.v. Chenopodium carinatum f. rubellum Aellen, Feddes Repert. 24:345 (1928). C. pumilio f. rubellum (Aellen) Aellen, Verb. Naturf. Ges. Basel 44:315 (1933). Type ci- tation: “Johannisburg (advent, aus Austr.) 1922, leg. C. E. Moss! no 6702 (Herb. Pretoria).” Type: n.v. Chenopodium x christii n-var. intermedium Aellen, Verb. Naturf. Ges. Basel 44:317 (1933). Type: 120 km S of Perth, Western Australia, C. Andrews 1st. Coll, no 697 (holo: K n.v.). Chenopodium x christii n-var. semiconnatum Aellen, Verb. Naturf. Ges. Basel 44:317 (1933). Type citation: “In Nova Hollandia ad portum Jackson, 1830 lee Gaudichaud'\ Type: G-DC n.v. Chenopodium pumilio var. oblongifolium J. Black, FI. S. Austral, ed. 2, 289 (1948) nom. illeg., sine desc. latina. [Chenopodium carinatum auct. non R.Br.: F. MuelL, Icon. Austral Sals PI t 32 19??’29g a972)'p^T'^^^ ^ [Blitum carinatum auct. non (R.Br.) Steudek 252 f. 8-10 (1909).] Reichenbach, Ic. FI. Germ. 24: 118, t. Annual or short-lived perennial herb to 30 cm high and wide, usually with several stems arising from base, aromatic. Stems slender, simple or branched, sparsely to moderately pilose with both slender segmented and sessile to stipitate gland-tipped hairs. Leaves herbaceous; lamina narrow to broad elliptic to ovate, 10-20 mm long obtuse, entire to sinuate or obtusely lobed, sparsely pilosulose with simple and gland-tipped hairs; petiole slender, c. half to equal length of lamina. Flowers in com- pact axillary glomerules, subsessile or on slender pedicels up to 2 mm long, ovoid, c. 0.5 mm high; tepals 5, erect, naviculiform, united towards base, herbaceous, thin sparsely pilosulose at apex or along midrib. Fruiting perianth usually crustaceous and white, c. 1 inm high; tepals erect, narrowly naviculiform, rounded on back, con- tiguous at apex but often disjunct in middle to expose fruit. Pericarp diaphanous dull grey or translucent, sparsely papillose. Seed lenticular 1 mm long keeled or rounded on margin. §^78^.^^^^^^' Longreach, S. T. Blake 6615 (BRI); Bengalla, L. Leichhardt (MEL 173 Paul G. Wilson, Tribe Chenopodieae NEW SOUTH WALES: 78 mi [125.5 km] W of Cobar, J. C. De Nardi 225 (NSW); Broken Hill, P. L. Milthorpe 176 (NSW). VICTORIA: Kul Kyne Forest, A. C. Beauglehole 40651 (MEL); Creswick, Whan 196 (MEL). TASMANIA: Buckland, 24 Apr. 1912, R. A. Black (HO); Cape Barren Is., J. S. Whinray 733 (AD). SOUTH AUSTRALIA: Millswood, D. N. Kraehenbuehl 1878 (AD); Queenscliffe, Kangaroo Is., J. G. 0. Tapper 1285 (MEL). WESTERN AUSTRALIA: Wyola, T. E. H. Aplin 2203 (AD, PERTH); Pemberton, M. Koch 2558 (PERTH). NORTHERN TERRITORY: Burt Plain, G. Chippendale (NT 8033). Distribution. Principally southern Australia; as an introduced weed in many parts of the world and as an occasional weed in northern Australia. Map 13. Habitat. Heavy loamy soils, sometimes slightly saline; frequently an agricultural weed. The name Chenopodium carinatum was used by Bentham (1870) to include ma- terial correctly placed in C. pumilio. He was followed in this regard by Mueller (1889-1891), J. M. Black (1924) and Ewart (1931). It was not until 1933, when Aellen published a revision of the section Orthosporum, that the separate identity of both species became widely accepted, although some Floras continue to unite them. Chenopodium carinatum and C. pumilio are evidently closely related. It would ap- pear that in Australia with the widespread dissemination of C. pumilio since the coming of agriculture (in which industry it features as a prominent weed) it has spread into areas previously only occupied by C. carinatum. Integradation between the two is apparent from a study of herbarium material. The situation is compli- cated by the considerable variation, found between different populations of C. pumilio, in habit, leaf shape, and perianth structure. In spite of this variability I have not recognised any infraspecific taxa since the variants do not appear to occupy discrete areas and there appears to be a smooth gradation between them. In Chenopodium pumilio, and possibly in other members of the section, the an- terior tepal of hermaphrodite flowers subtends the solitary stamen and is initially larger than the others. At anthesis the swollen base of the staminal filament becomes turgid and causes the stamen and anterior tepal to spread widely. Later all tepals be- come equal in size and assume an erect posture. The perianth usually becomes carti- laginous and white in fruit but this change does not always occur and would appear to depend on the conditions under which the plant is growing. 19. Chenopodium carinatum R.Br., Prod. 407 (1810); Benth., FI. Austral. 5:162 (1870) p. pte; Aellen, Verh. Naturf. Ges. Basel 44:312 (1933) exc. var. mel- anocarpum. — Salsola carinata (R.Br.) Sprengel, Syst. Veg. 1:923 (1825). — Ambrina carinata (R.Br.) Moq., Chenop. Enum. 41 (1840). — Blitum carinatum (R.Br.) C. A. Meyer ex Steudel, Nom. Bot. ed. 2, 1:210 (1840); Moq. in DC., Prod. 13/2:82 (1849). Type: Hawkesbury R., New South Wales, R. Brown (holo: BM). 174 Nuytsia Vol. 4, No. 2 (1983) C. cristatum var. holopterum Thellung, Vierteljahrsschr. Naturf. Ges. Zurich 64:724 (1919). — C. holopterum (Thellung) Thellung et Aellen in Probst, Mitt. Naturf. Ges. Soloth. 20(8):57 (1928). — C. carinatum var. holopterum (Thellung) Aellen, Verh. Naturf. Ges. Basel 44:312 (1933). Type citation: ‘Mit dem Typus der Art und mit Ch. carinatum, 1917, Probst. ’ Type: n.v. Ambrina carinata var. parvifolia Moq., Chen. Mon. Enum. 42 (1840) pro parte as to lectotype.— Pbtum glandulosum var. parvifolium (Moq.) Moq. in DC., Prod. 13/2:82 (1849). Lectotype: D’Urville (P, n.v.) see Aellen (1933). C. carinatum f. foliosum Domin, Biblio. Bot. 22:65 (1921). Type: Queensland, Brisbane River, A. Dietrich 968 (iso: NSW). Annual herb, decumbent to erect, branching from the base, aromatic. Stems simple or branched, 30 cm long, pilosulose with both slender and sessile to stipitate gland-tipped hairs. Leaves herbaceous; lamina ovate to broadly ovate, 5-20 mm long, shortly lobed to coarsely serrate, glandular puberulous and pilosulose below on veins; petiole slender, rather shorter than lamina. Flowers subsessile, densely congested in fruit; tepals 5, erect, united towards base, narrow-oblong, naviculiform, hirtellous on keel. Fruiting perianth white, thickened and coriaceous; tepals prominently keeled especially towards apex, truncate or acute, sparsely hirtellous on keel. Pericarp di- aphanous, smooth. Seed lenticular c. 0.5 mm long, keeled over radicle and channelled over cotyledons. QUEENSLAND: Hughenden, S. T. Blake 6181 (BRI); Wandoon, C. E. Hubbard 5072 (BRI). NEW SOUTH WALES: Kunghur, E. F. Constable 6578 (NSW); 1.5km N of Wollombi, L. A. S. Johnson and B. G. Briggs 3287 (NSW). VICTORIA: Suggan Buggan, A. C. Beauglehole 36646 (MEL); Near Deddick, 21 Jan. 1948, J. M. Bechervaise (MEL) Distribution. Eastern Queensland and New South Wales, far eastern Victoria. Map 4. Habitat. Frequently an agricultural weed or an invader of disturbed soils. There has been much confusion over the application of the name Chenopodium carinatum, this arose through Moquin (1840, 1849) not having seen the type of that species or of C. pumilio and therefore including material of the latter under the for- mer name. This confusion was continued by Bentham (1870) who, while recognising both species placed most specimens of each under C. carinatum and included in it the plant later recognised as the species C. melanocarpum. The situation was clari- ried by Aell^ (1933) who, unfortunately, had an individual approach to the international Code of Botanical Nomenclature and who, for his understanding of the species, relied to a large extent on plants arising in Europe from seed introduced there in wool. While I recognise C. carinatum as a distinct species, hybrids with C. pumilio, C. melanospermum, and C. cristatum are common, probably largely through the move- ment of seed of these species due to agriculture and commerce; this is particularly apparent in Europe where numerous hybrids have been documented and in some cases named. The species exhibits considerable geographic variation in leaf and peri- anth form m particular in the tepals which range from being slightly to strongly Paul G. Wilson, Tribe Chenopodieae 175 keeled. When strongly keeled the fruiting perianth has a form approaching that of C. melanocarpum. Specimens with a particularly prominently keeled and hardened perianth from central and South Australia and from north-western Western Aus- tralia have, ill herbaria, sometimes been placed in C. carinatum and sometimes in C. melanocarpum. I have placed them with the latter taxon, as did Aellen who named this variant f. leucocarpum, although I recognise that the plants with this form of fruit have evolved in several independent lines: some have affinity to C. carinatum, some (a portion of the western element) probably to C. melanocarpum, while others which 1 have segregated under f. leucocarpum may represent a hybrid between C. ca- rinatum and C. melanocarpum. 20. Chenopodium melanocarpum (J. Black) J. Black, Trans. Roy. Soc. S. Austral. 58:173 (1934). — C. carinatum var. melanocarpum J. Black, Trans. Roy. Soc. S. Aus- tral 46:566 (1922); Aellen, Verb. Naturf. Ges. Basel 44:313 (1933); Aellen in Hegi, 111. FI. Mitt.-Eur. ed. 2, 3:599 (1960). Lectotype: Broken Hill, New South Wales, Sept. 1918, per T. G. B. Osborn (AD 96732197) lecto. nov. [C. carinatum auct. non R.Br.: Benth., FI. Austral. 5:162 (1870) p.p. as to Drummond 165.] [C. holopterum auct. non (Thell.) Thell. et Aellen: J. Black, FI. S. Austral. 683 (1929) “C. holocarpum Thell. et Aellen”. See note below.] Annual herb, prostrate, with numerous stems arising from the base, aromatic. Stems simple, slender, to 30 cm long, pilosulose with both slender segmented and sessile to shortly stipitate gland-tipped hairs. Leaves slightly fleshy; lamina broadly elliptic c. 15 mm long, bluntly lobed to almost entire, glandular puberulous and pilosulose below along veins; petiole slender about half length of lamina. Flowers subsessile to shortly pedicellate -(densely congested in fruit); tepals 5, erect, united in lower half, narrow-oblong, keeled, hirtellous; stigmas 2, subsessile, slender. Fruiting perianth bluntly stellate in cross-section, crustaceous, black (or pale fawn in f. leucocarpum, firmly united in lower half and completely covering fruit; tepals strongly and bluntly keeled, sparsely hirtellous on keel. Seed lenticular, 0.5 mm long, keeled on margin over radicle, channelled over cotyledons; pericarp diaphanous, dull, smooth or minutely puberulous towards apex, somewhat adherent to seed. This species comprises two forms which may be distinguished by the colour of their fruiting perianths. 20a. forma melanocarpum Fruiting perianth hlack. QUEENSLAND: 35 mi [56.3 km] SW of Eulo, M. law 43 (BRI); Boatman Station, S. L. Everist 2792 (BRI). NEW SOUTH WALES: Mt. Derriwong, G. M. Cunningham 774 (NSW); 12 mi [19.3 km] E of Turlee, J. De Nardi 340 (NSW). SOUTH AUSTRALIA: Mann Range, 22 Aug. 1954, J. B. Cleland (AD); Amoorinyirra Hill, D. J. E. Whibley 1124 (AD). WESTERN AUSTRALIA: 80 mi [128.7 km] E of Carnarvon, C. A. Gardner 6051 (PERTH); Mileura Stn., P. G. Wilson 8987 (PERTH). 176 Nuytsia Vol. 4, No. 2 (1983) NORTHERN TERRITORY: Palm Valley, A. C. Beauglehole 10403 (PERTH); 4 mi [6.4 km] N of Alice Springs, R. Swinbourne 563 (NT). Distribution. Arid and semi-arid areas of Australia south of 22° latitude. Map 7. Habitat. Well-drained situations either in sand or on sides of rocky hills. The synonymy cited above is indicative of the difficulty experienced by different workers in fitting a nomenclatural framework onto a variable and freely interbreeding group. Bentham (1870 op.cit.) placed material of C. melanocarpum (and of C. pumilio) under C. carinatum, while Black (1922 op.cit.) still recognising it as part of that species, gave it varietal status, in which decision he was followed by Aellen (1933, 1960-1961). Later Black (1934 op.cit.) raised var. melanocarpum to the rank of species and in this he has been followed by workers in most Australian herbaria. The course adopted by Black is preferable since the typical form of C. mel- anocarpum differs appreciably in perianth and leaf shape from C. carinatum. The spelling “C. holocarpum” was inadvertently used by J. M. Black (1924) in- stead of C. holopterum. This error was later noted by Black in Trans. Roy. Soc. S. Austral. 58:174 (1934) when he published the name C. melanocarpum as a new com- bination. 20b. forma leucocarpum (Aellen) Paul G. Wilson, comb, nov.— C. carinatum var. melanocarpum f. leucocarpum Allen, Verb. Naturf. Ges. Basel 44:313 (1933). Neotype, here chosen: Ashburton River, Minderoo, Oct. 1905, A. Morrison (holo- PERTH; iso: K). . v • Fruiting perianth pale fawn in colour. Distribution. North western Western Australia (Port Hedland-Exmouth Gulf area), central and South Australia, eastern Queensland, New South Wales and Victoria’ Map 4. Habitat. As for the typical form. No collections were cited by Aellen for this forma, however, a duplicate at herb K of the Morrison specimen which has been designated as a neotype has been deter- mined by Aellen as f. leucocarpum. 21. Chenopodium cristatum (F. Muell.) F. Muell., Fragm. 7:11 (1864); Benth., FI. Austral. 5:163 (1870); F. Muell., Icon. Austral. Salsol. PI. 31 (1890); Aellen, Verb. Naturf. Ges. Basel 44:310 (1933).— Blitum cristatum F. Muell., Trans. Phil. Inst. Viet. 2:73 (1858). Lectotype. Raro in planitiis sterilioribus subsalsis prope montis Flindersianos, Oct. 1851, F. Mueller (holo: MEL 19998) iso: PERTH) lecto. nov. Annual herb, prostrate to ascending with several stems arising from base, aro- matic Stems slender, mainly simple, pilosulose with both slender segmented and sessile to shortly stipitate gland-tipped hairs. Leaves slightly fleshy; lamina elliptic to broadly elliptic, entire to coarsely toothed, sparsely puberulous. Flowers subsessde, tepals 5, erect, acuminate, strongly keeled and hirsute, ciliate, united in lower half. FruAing perianth spherical, c. 2 mm diameter, strongly 5-winged, white (rarely grey to black), tepals cartilaginous, strongly crested (vertically winged), semi- Paul G. Wilson, Tribe Chenopodieae 17^ orbicular, rostrate, completely enclosing the fruit. Pericarp diaphanous dull, papil- late towards apex or all over. Seed lenticular, c. 0.5 mm long, the margin slightly keeled over radicle and grooved over cotyledons. QUEENSLAND: Oakwood Stn., L. S. Smith and S. L. Everist 846 (BRI); Thargomindah, Apr. 1885, Spencer (MEL). NEW SOUTH WALES: “Birdwood” NE of Pooncarie, R. P. Green 177 (NSW); W. of Stoney Crossing, D. L. W. Henderson 327 (NSW). VICTORIA: 23 km NNW of Manangatang P.O., A. C. Beauglehole 55812 (MEL); nr. Lakes Ranfurly and Hawthorn, May 1937, J. H. Willis (MEL). SOUTH AUSTRALIA: Koonamore, Hj. Eichler 12492 (AD); “Markaranka”, Morgan, D. E. Symon 3557 (ADW). WESTERN AUSTRALIA: Esperance, R. A. Rose 3005 (PERTH); W side Fraser Ra., P. G. Wilson 5898 (PERTH) NORTHERN TERRITORY: Trephina Gorge, B. G. Briggs 1229 (NSW); 2 mi [3.2 km] S of Weepita Bore, G. Chippendale (NT 2499). Distribution. Arid and semi-arid regions of Australia south of 20° latitude. Map 14. Habitat. Frequently in red sandy loam, often long creek lines. This species may be readily recognised when in fruit by the strongly crested ros- trate tepals. It is most similar to C. truncatum which may be distinguished by the firmly united tepals and the absence of a rostrate apex to the perianth. As is noted in the species description, the fruiting perianth while normally white is sometimes grey to black in colour. In some cases these melanized plants have obvi- ously arisen through hybridisation with C. melanocarpum (i.e. when the grey fruited form has been collected with the two parents); in other cases the fruit shape is typical of C. cristatum while the colour is black and no plants of C. melanocarpum have been observed in the area. 22. Chenopodium truncatum Paul G. Wilson, sp. nov. Herba prostrata vel adscendens cum pilis simplicibus et pilis glandulosis subsessilibus pilosula, caulis numerosis ex caudice exorientibus. Folia modice carnosa; lamina elliptica obtusa c. 10 mm longa, Integra vel leviter undulata, glanduloso puberula. Flores in glomerulis axillaribus depositi; tepala 5 versus apicem connata, c. 0.5 mm longa; stamina 0-1, glabra. Perianthium fructiferum cartilagineum, late obtriangulare, c. 2 mm latum, apice applanato, marginis dentatis vel breviter laceratis, tepalis firme connatis. Pericarpium diaphanum, nitidum, glabrum. Semen erectum, lenticulare, c. 0.7 mm longum, margine rotundato. Typus: 65 mi [104.6 km] W of Henbury HS on road to Kings Canyon, 10 Dec. 1968 P. K. Latz 255 (holo: AD; iso: CBG, NT). Annual or (short-lived) perennial herb, prostrate or ascending, with numerous stems arising from rootstock, aromatic. Stems slender, pilosulose with both slender segmented and sessile to shortly stipitate gland-tipped hairs. Leaves slightly fleshy; 178 Nuytsia Vol. 4, No. 2 (1983) lamina elliptic c. 10 mm long, obtuse, entire or slightly undulate, glandular puberulous all over and pilosulose below along veins; petiole slender about half length of lamina. Inflorescence of discrete axillary glomerules. Flowers ebracteate, subsessile, bisexual and female (the terminal always bisexual), ovoid, c. 0.5 mm high; tepals 5, united to near apex, hirtellous on midrib; stamen 0-1, glabrous, anther shortly exserted. Fruiting perianth cartilaginous, broadly obtriangular in plain view, flat above (or the lobes slightly ascending), lateral margins toothed or shortly lacerate, shortly hirtellous, white or black, c. 2 mm wide, the tepals firmly united to apex. Pericarp diaphanous, glossy, glabrous. Seed erect, lenticular c. 0.7 mm high, margin rounded (neither grooved nor keeled). QUEENSLAND: Thylungra, S. L. Everist 5717 (CANB); Oakham Stn., Windorah, 28 Aug. 1959, R. S. Stranger (BRI). NEW SOUTH WALES: Near Mt Murchison, Jan. 1887, R. H. Kennedy (MEL); Calindry Stn., J. H. Leigh W183 (NSW). SOUTH AUSTRALIA: NE corner of Lake Gairdner, B. Lay 341 (AD); 61 km E of Dalhousie Springs, T. R. N. Lothian 1917 (AD); 10 km W of Quinyambi HS, D. J. E. Whitley 5557 (AD). NORTHERN TERRITORY: Old Andada Homestead, A. C. Beauglehole 27967 (NT); 5 mi [8 km] SE of Harper Springs Homestead, G. Chippendale (NT 6468). Distribution. South-west Queensland, eastern New South Wales, northern and central South Australia, the southern half of the Northern Territory. Map 3. Habitat. Usually associated with deep red sand. Specimens of this species have, in Australian herbaria, generally been placed with Chenopodium cristatum, in some cases with the suggestion that they may be hybrids with C. melanocarpum. Chenopodium truncatum differs from C. cristatum most ob- viously in having a fruiting perianth which is truncate above (not with a rostrate apex), the tepals firmly united to apex (not free) and not so markedly crested; the pericarp is glabrous and glossy (not papillate and dull) and the lenticular seed has a rounded margin (not keeled or grooved). From C. melanocarpum it differs in having almost entire leaves (not strongly lobed) and in having crested tepals. Seven of the collections (representing about 15 per cent of the total) consisted of mixed material; in five C. cristatum was also present, in one C. melanocarpum, and in the other C. pumilio. This suggested that C. truncatum was of hybrid origin. However, most sheets of this species contained more than one specimen and in no case was there any evidence of introgression. In addition hybrids between C. cristatum and C. mel- anocarpum which I have observed on sheets of mixed material have a different ap- pearance, being intermediate in morphology in both leaf and fruit characters. The illustration provided by Aellen (1933) for the fruiting perianth of C. xbontei var. suhmelanocarpum resembles the fruiting perianth of C. truncatum (except for the union of the tepals in the latter). I have examined the two syntypes of var. submelanocarpum and find them to be intermediate between C. melanocarpum and C. cristatum as was stated by Aellen. The specific epithet refers to the flat upper surface of the fruiting perianth. Paul G. Wilson, Tribe Chenopodieae 179 23. Chenopodium pusillum J. D. Hook., Handb. N. Zeal. FI. 231 (1864); Cheeseman, Man. N. Zeal. FI. ed. 2, 407 (1925); Allan, FI. N. Zeal. 1:228 (1961). Type citation: ‘Sandy shores oF Lake Taupo’. Type: Colenso (K, n.v.). [C. pumilio auct. non R.Br.: J. D. Hook., FI. Nov.-Zel. 1:214 (1853).] Small decumbent annual with wiry stems radiating from base, c. 10 cm diam., pi- lose with simple segmented hairs and with sessile or subsessile gland-tipped hairs, aromatic. Leaves somewhat fleshy; lamina suborbicular to broadly ovate, obtuse, c. 5 mm long, entire or sparsely and bluntly dentate, sparsely puberulous above, glandular puberulous below; petiole c. half length of lamina. Flowers in compact axillary glomerules, shortly pedicellate; tepals 4, erect, c. 0.5 mm long, membranous apart from the narrowly oblong midrib, united in lower half, pilose, eventually en- larging to 1 mm long but not obscuring fruit. Stamen 1. Pericarp membranous, free from seed. Seed compressed obovoid to lenticular, 0.5-0.8mm long; testa glossy, finely puncticulate, dark reddish brown. NEW ZEALAND: Christchurch, gravel ballast, 23 Feb. 1949, A. J. Healy (CHR); Lake Lyndon, T. Kirk 1079 (CHR); Marlborough, on formed road, 23 Apr. 1952, L. B. Moore (CHR). Distribution. New Zealand. Habitat. ‘In sandy and open places’ (Allan 1961); frequently a coloniser of disturbed sites. This species has been confused with Chenopodium pumilio which it resembles. The two may be distinguished by the much smaller and more orbicular leaves of C. pusillum, the 4-merous perianth, and by the membranous margin to its lobes which do not become hardened in fruit. Chenopodium pumilio is also found in New Zealand (presumably as an introduction). When growing in the same environment as C. pusillum, the two are very similar in habit and general appearance. 24. Chenopodium saxatile Paul G. Wilson, sp. nov. Herba annua erecta subtilis aromatica ad 30 cm alta, pilis debilibus glanduliferis et pilis simplicibus rigidis modice pilosa. Caulis et rami tenues. Folia alterna; lamina chartacea, ovata vel elliptica 10-20 mm longa, integra, sinuata, vel obtuse lobata. In- florescentia paniculata, ramis flexuosis; Bores aggregati. Tepala 4, erecta, naviculiformia, breviter pilosa; stamina 0-2, filamento non pulvinato. Pericarpium diaphanum ad semen adherens, glabrum vel minute glandulosum. Semen erectum, lenticulare, c. 0.6 mm altum; testa minute puncticulatum; embryo hippocrepiformis ad apicem semine positus; radicula inferior. Typus: 24 km E of Depot Springs HS., 28°54’S, 121° 20’E, Western Australia; base of breakaway; 27 Aug. 1970, P. G. Wilson 8909 (holo: PERTH; iso: K). Delicate erect aromatic annual herb to 30 cm high, moderately pilose all over with weak gland-tipped and stiffer simple segmented hairs. Stem and branches slender. Leaves alternate; lamina thin ovate to elliptic, 10-20 mm long, entire, sinuate, or ob- tusely lobed; petiole slender c. half length of lamina. Inflorescence paniculate eventually occupying most of plant, composed of open dichasia with flexuose branches which become abbreviated towards apex of plant, the flowers sessile in small clusters. Flowers bisexual and female, c. 1 mm high (c. 1.5 mm in fruit); tepals 180 Nuytsia Vol. 4, No. 2 (1983) 4, erect, naviculiform, shortly pilose, fleshy with membranous margins; stamens 0-2, slightly exceeding perianth, filament linear (not pulvinate at base), anther broadly cordate c. 0.25 mm long; stigmas 2 filamentous, united into a short style. Utricle en- closed in perianth which remains herbaceous in fruit; pericarp dull, diaphanous and adherent to seed, glabrous, or minutely glandular. Seed erect, lenticular, c. 0.6 mm high; testa minutely puncticulate; embryo hippocrepiform around apex of seed, rad- icle inferior. Fruit shed while enclosed in perianth. Distribution. In the central portion of the Eremaean Botanical Province of Western Australia. Map 15. Habitat. On rocky outcrops and at the base of breakaways. WESTERN AUSTRALIA: Mt Magnet, Sept. 1903, W. V. Fitzgerald (NSW); Pass of the Abencerrages, Rawlinson Ra., A. S. George 12137 (PERTH); near Mt Squires, 27 Aug. 1891, R. Helms (AD). Superficially this species would appear to be related to the 4-merous species in the genus Dysphania (e.g. D. rhadinostachya) but in seed character it is obviously differ- ent, being very similar to Chenopodium pumilio and its allies of the section Orthosporum. Yet even here its placement is uncertain since the other species of this section have flowers arranged in compact glomerules, have one stamen, and normally a 5-merous perianth. Aellen (1939 in sched.) was of the opinion that it was a member of the Chenopodium botrys complex, but those species (and others of the section Botryoides) differ in having 5-merous perianths and horizontal seeds. In 1956 Brenan described the sect. Margaritaria which contained the single species Cheno- podium congolanum (Haum.) Brenan; this species is similar to C. saxatile except that it has whitish sub-globose hairs (as well as glandular) and only one stamen, in addition the terminal flower of each axillary cyme has a horizontal (not vertical) seed. Thus it appears that C. saxatile does not fit satisfactorily into any of the pres- ently recognised sections. Rather than create a further section, which would be monotypic, I have expanded the circumscription of the section Orthosporum to in- clude this species. The axillary inflorescence (a dichasium) is similar in morphology to the compact glomerule typical of the sect. Orthosporum', its different appearance being due to the more open lateral branches which, towards the base of the plant, themselves branch and become flexuose. The glandular hairs found in C. saxatile appear to be modifications of the normal multicellular trichome; the gland itself being very small and often scarcely observ- able. This situation is very different to that in the other species of the section where the glandular hairs consist of a large globular gland cell, either sessile or on a short stipe. The specific epithet refers to rocky situations in which Chenopodium saxatile is found. 2. DYSPHANIA R. Br. R.Bj., Prod. 411 (1810); Moq. in DC., Prod. 13/2:86 (1849); Benth., FI. Austral. 5:164 (1870); J. D Hook in Bentham et J. D. Hook., Gen. PI. 3:14 (1880) “Illecebraceae”; Payn Engler et Prantl, Nat. Pflanzenf. 3, lb:92 (1889) “Caryophyllaceae”; Pax et Holtmann in Engler et Prantl, Nat. Pflanzenf. ed. 2, 16c:272 (1934) Paul G. Wilson, Tribe Chenopodieae 181 “Dysphaniaceae”; Aellen in Hegi, III. FI. Mitt.-Eur. ed. 2, 3:748 (1961) “Dysphaniaceae”; Eckhardt in Melchior, A. Engler’s Syllabus der Pflanzenf. ed. 12, _ 2:96 (1964) “Dysphaniaceae”; Airy Shaw in Willis, Diet. FI. PL Ferns ed. 7.387 (1966) “Dysphaniaceae”; A. J. Scott Bot. Jahrb. Syst. 100:217 (1978). — Chenopodium sect. Dysphania (R.Br.) Aellen, Bot. Jahrb. Syst. 63:486 (1930); J. Black, Trans. Roy. Soc. S. Austral. 58:173 (1934). Type.' D. littoralis R.Br. Chenopodium sect. Tetrasepala Aellen, Bot. Jahrb. Syst. 63:490 (1930); J. Black, Trans. Roy. Soc. S. Austral. 58:172 (1934). — Dysphania sect. Tetrasepalae (Aellen) A. J. Scott, Bot. Jahrb. Syst. 100:218 (1978). Lectotype: C. inflatum Aellen (see A. J. Scott loc. cit.). Dysphania sect. Caudatae A. J. Scott, Bot. Jahrb. Syst. 100:218 (1978). Type; D. plantaginella F. Muell. Annual or short-lived aromatic perennial herbs with subsessile glands or simple and gland-tipped multicellular hairs, aromatic. Leaves alternate, simple, entire or variously dissected, fleshy or thin. Inflorescence of compact cymose glomerules of flowers (thyrses), these either axillary, spicoid or paniculate in arrangement. Flowers ebracteate, minute (0.5 mm long), bisexual and female; bisexual flowers (often functionally male) terminal to the glomerule or to each side of the glomerule; female flowers lateral. Tepals 1-4 (3 or 4 in terminal flowers of glomerule), free or united at base; limb cucullate and often inflated of spongy. Stamens 1-2, anterior-lateral in position, somewhat exceeding tepals at anthesis; filament slender thickened at base, disc absent. Ovary ellipsoid, erect; stigmas 1-2, linear, delicate. Fruit a nut; pericarp diaphanous, smooth to papillate. Seed erect, oblique, or horizontal, globular to ellip- soid or laterally compressed; testa crustaceous, smooth; embryo lateral or basal, situ- ated beneath a groove (“embryo-groove”), radicle either inferior or superior. Ten species all endemic to mainland Australia. The taxonomic position of Dysphania has been a matter of doubt or disagreement amongst botanists since Brown (1810) first described the genus. Brown considered it to have affinity to the family Chenopodiaceae, in which he placed the genus but evi- dently with reservations. Moquin (1849) also included it in the Chenopodiaceae (as Salsolaceae) and suggested affinities with Monolepis and Blitum. Bentham (1870) suggested that it was closely related to the section Orthosporum of Chenopodium (a view with which I concur). J. D. Hooker (1880) included it in the Illecebraceae and Pax (1889), who was evidently influenced by J. D. Hooker, in the Caryophyllaceae. However, Pax (1927) and later Pax and Hoffmann (1934) placed it in its own family, the Dysphaniaceae, which they considered to be intermediate in morphology be- tween (Chenopodiaceae and (Caryophyllaceae. This systematic arrangement was adopted by Aellen (1961), Eckhardt (1964 loc.cit.) and Airy Shaw (1966), who all rec- ognised its affinity to the Chenopodiaceae. Aellen had earlier (1930) made Dysphania a section of Chenopodium as did Black (1934 l.c.). The close relationship between Chenopodium and Dysphania appears to have always been recognised by workers in Australia who were familiar with some endemic species of the former genus that were very similar in habit and floral morphology to those of the latter. This relationship was not so evident to European systematists who would have compared species such as Chenopodium album with Dysphania and found them to be very different. The confusion created by the rather extreme pos- itions taken by several workers was finally resolved by Eckardt (1967, 1968, 1969) and by Scott (1978b). Eckardt showed that the floral morphology of Dysphania cor- responds to that of Chenopodium hut is quite different to that found in the 22493 — ( 4 ) 182 Nuytsia Vol. 4, No. 2 (1983) Illecebraceae (or Caryophyllaceae). Even so the circumscription of Dysphania re- mained unsatisfactory for it included only species with three or fewer perianth seg- ments. The addition to the genus by Scott (1978b) of the two species placed by Aellen (1930) in Chenopodium sect. Tetrasepala was a big improvement for it then became a discrete taxon clearly separable from Chenopodium. Scott recognised three sections in Dysphania; these were based on the number of perianth segments (1-2, 3, or 4) and the arrangement of the flower clusters (axillary or spicate). These sections appear to be natural ones (although the tepal numbers given by Scott are not always correct) but with so few species it appears to be unnecessary to provide an infrageneric classification. The genus Dysphania is closely related to Chenopodium sect. Orthosporum (as was recognised by Bentham, 1870). This section has similar glandular hairs and glomerules of small flowers with 4-5 tepals and one stamen. The only characters which suggest that the two genera should be recognised as distinct are found in the seed. In Dysphania the seed (when erect) has either a basal or lateral embryo (or it is intermediate in position); when lateral the radicle is either inferior or superior. In Chenopodium sect. Orthosporum the erect, lenticular seed always has a hippocrepi- form embryo curved around its apex. The position of the embryo in the Dysphania seed, is indicated by a corresponding depression in the testa; this I am referring to as the embryo-groove. The radicle is situated in a pouch formed by an invagination of the testa; this pouch is usually slightly protruding and, since at this point the testa is thicker, it also appears darker in colour. The orientation of the embry-groove and the position of the pouch are characters useful in recognising the different species. Key to species 1. Tepals 1-3 Tepals 4 2. Flowers in glomerules axillary to foliage leaves 3 Flowers in spike-like inflorescences 8 3. 4. 5. 6. 7. 8. Tepals (of lateral flowers of glomerules) 1-2; seed erect, ellipsoid to obovoid or laterally flattened Tepals 3, seed napiform to subglobular 5. D. littoralis Tepals united at base and shed with the enclosed seed 5 Tepals free; seed and tepals shed separately 8 Seed lenticular, embryo curved around its base 3. D. valida Seed obvoid or ellipsoid, embryo lateral 1. D. glomulifera Seed strongly flattened (usually slightly twisted), obovate to deltoid; pericarp almost smooth. piatycarpa Seed ellipsoid, obovoid, or lenticular; pericarp smooth or glandular-papillate . . 7 Pericarp smooth, puncticulate, or minutely papillate; seed ellipsoid to obovoid or lenticular 1. D, glomulifera Pericarp prominently papillate; seed asymmetrically narrow-obovoid (the em- bryo side flat) 2. D. glandulosa Seed compressed globular, embryo oblique to erect 7. D, sphaerosperma Seed ellipsoid, erect q Paul G. Wilson, Tribe Chenopodieae 183 9. Tepals free from each other, rounded or inconspicuously keeled on back 6. D. plantaginella Tepals united into a hard cup-shaped tube in lower half, prominently (and hori- zontally) keeled 8. D. simulans 10. Plant prostrate or decumbent, many stems arising from base; seed (i.e. pericarp) glossy, compressed globular; inflorescence slender spicate 10. D. kalpari Plant erect with one main stem; seed dull (rarely glossy), almost globular; inflor- escence paniculate 9. D. rhadinostachya 1. Dysphania glomulifera (Nees) Paul G. Wilson, comb. nov. — Atriplex glomulifera Nees in Lehm., PI. Preiss 1:634 (1844); Moquin in DC., Prod. 13/2:103 (1849); Benth., FI. Austral. 5:177 (1870); Ulbrich in Engler et Prantl, Nat. Pflanzenf. ed. 2, 16c:515 (1934). Type: “In cultis ad caput fluvii Cygnorum, Januario a 1840”. Preiss 1257 (holo: LD; iso:P). D. myriocephala Benth., FI. Austral. 5:165 (1870).— C. myriocephalum (Benth.) Aellen, Bot. Jahrb. Syst. 63:488 (1930). Lectotype: Western Australia, J. Drummond 206 (holo: K n.v.; iso: MEL, P) lecto. nov. D. benthamiana Domin., Biblio. Bot. 89:103 (1925). Type: “Flooded ground south of Wills Creek, Howitts Expedition”, Dr Murray (iso: MEL 19878). Prostrate, decumbent, or erect annual or short-lived perennial herb c. 25 cm diam., to 15 cm high, numerous slender stems arising from base, sparsely covered with sess- ile or stipitate gland-tipped hairs. Leaves fleshy, crowded towards the apex; lamina narrow to broad elliptic, (1)5(10) mm long, passing into a slender petiole c. half its length. Glomerules of flowers dense, present in most leaf axils. Terminal flower (of glomerule) bisexual; tepals 3, obovate, cucullate, c. 0.7 mm long, green, fleshy except for scarious margin, glabrous or with a few marginal glandular hairs, shortly united at base or free; stamens 2 shortly exserted at anthesis; style solitary or paired. Lat- eral flowers (of glomerule) female; tepals 1 or 2 with a white bladdery strongly cucul- late limb and a filamentous to linear claw; stamens absent; ovary ellipsoid, delicate; styles 1 or 2. Fruiting perianth: tepals 0.7-1 mm long; limb inflated and fungoid covering the fruit. Pericarp diaphanous dull, closely adhering to seed, puncticulate, papillate, or granulate. Seed erect, bluntly ellipsoid, compressed obovoid, or some- what lenticular with rounded margins, c. 0.5 mm long; testa smooth reddish brown; embryo slightly curved extending along the dorsal lateral margin beneath the em- bryo-groove; radicle inferior with the radicular pouch forming a small dark protuber- ance at base of seed. Typification. The type of Dysphania glomulifera (i.e. Preiss 1257) came from near Guildford (c. 15 km east of Perth) in cultivated ground near the Swan River (‘in cultis ad caput fluvii Cygnorum’). It is a precise match with the lectotype of D. myriocephala (Drummond 206) which probably came from the same general area since it is only near here that the particular variant this collection represents has been found. Aellen (1938) recognised that the plant described by Nees (1844) as Atriplex glomulifera did not belong to that genus but since Aellen assumed the name to be a synonym of D. littoralis R.Br. (1810) he did not make a new combination under Dysphania. Very few replicates of Preiss no. 1257 appear to exist; neither Moquin (1849) nor Bentham (1870) saw this number (they relied on Nees’ descrip- tion when writing their own works) and both assumed the species to be correctly placed in Atriplex although a puzzling member. Moquin (1849) based his descrip- tion of D. littoralis on Drummond 206, the lectotype of D. myriocephala; therefore his description applies to D. glomulifera rather than to the former species. 184 Nuytsia Vol. 4, No. 2 (1983) Key to subspecies Seed compressed obovoid to bluntly lenticular; pericarp smooth (rarely papillate) Seed ellipsoid; pericarp granulate la. subsp. glomulifera lb. subsp. eremaea la. subsp. glomulifera Figure 6A. D. myriocephala Benth., op. cit . — Chenopodium myriocephalum (Benth.) Aellen, op. cit. Lateral flowers of glomerules: tepals 1(2); claws filiform to linear, readily separating from pedicel (and from opposite member where present); styles (1)2. Seed broadly compressed obovoid to bluntly lenticular; pericarp smooth or minutely puncticulate (rarely papillate). QUEENSLAND: Moonie Ponds, S. L. Everist 3718 (BRI); Cunnamulla, P. Martensz 62 (CANB). NEW SOUTH WALES: Near Booligal, E. M. Canning 3859 (CBG); Coolabab, W. E. Mulham W857 (NSW). VICTORIA: 15 mi [24.1 km] NE of Hattah, A. C. Beauglehole 40569 (PERTH)- Lake Eildon, 14 May 1973, J. H. Willis (MEL). SOUTH AUSTRALIA: Loxton, Hj. Eichler 12326 (AD); Lenswood, A. G. Spooner 1 *7 A / A \ ■* 5174 (AD). WESTERN AUSTRALIA: Guildford, 12 Aug. 1901, C. R. P. Andrews (NSW)- Leederville, Mar. 1967, 1. S. Hutchinson (PERTH). Distribution. Near Perth in Western Australia, south-eastern and north-eastern South Australia, Victoria, eastern New South Wales and Queensland. Map 17. Habitat. Principally along banks of freshwater streams or rivers and in muddy areas. Some ecotypes in forest areas, rocky hillsides, or as weeds of cultivation. Ib. subsp. eremaea Paul G. Wilson, subsp. nov. Figure 6B. D. benthamiana Domin., op. cit. Flores laterales: tepala 2, limbis multo inflatis fructum obtegenti; stylus plerumQu© solitarius. Semen obtuse ellipsoideumj pericsrpium granulatum. Birdsville, S. T. Blake 12224 (BRI); Boorara Stn., R. C. Carolin 4035 (SYD). Paul G. Wilson, Tribe Chenopodieae 185 NEW SOUTH WALES: Wansaring, L. A. S. Johnson 547/125 (NSW); Wanheroo Hills, P. G. Milthorpe 1915 (NSW). SOUTH AUSTRALIA: Coglin River, Basendow 7 (AD); Millars Ck. Stn., R. J. Chinnock 2573 (AD). WESTERN AUSTRALIA: 22 mi [35.4 km] W of Browne Ra., A. S. George 5417 (PERTH); 18 mi [29 km] E of Meekatharra, T. E. H. Aphn 2460 (PERTH). NORTHERN TERRITORY: Redbank Gorge, G. Chippendale (N T 4878); Mt Doreen Stn., P. K. Latz 2002 (NSW). Distribution. South-West and central Queensland, north-west New South Wales, northern South Australia, central Western Australia, southern portion of the North- ern Territory. Map 17. Habitat. Damp sand near freshwater. The typical variants of the two subspecies of D. glomulifera are quite distinct (at least as to microscopic characters) but there are numerous other variants which blur the separateness and provide additional characters that extend the morphological complexity. Since the distinguishing features are so small (and fragile) and appear to be subject to environmental modification, it appears unwise to treat the two taxa at the species level or to formally recognise further infraspecific groups. Near the Murray River in Victoria are found two variants of subsp. glomulifera-, in one the pericarp is smooth and in the other it is papillate. There do not appear to be other morphological characters which might support the recognition of these vari- ants as distinct taxa. The variant with papillae on the pericarp may be readily dis- tinguished from D. glandulosa by the shape of the seed. The variant of subsp. glomulifera found in central and eastern New South Wales and in central and eastern Queensland, has a bluntly lenticular seed. This variant may be clearly separated from D. valida by its free (or almost free) tepals and by the embryo being lateral, not basal, in position. The subspecies eremaea represents the plant referred by Aellen (1930) to Cheno- podium blackianum, and by J. M. Black (1948) to its nomenclatural synonym Dysphania littoralis. Most Australian herbaria have followed Black in their misappli- cation of the latter name. Domin (1925) appears to have recognised the two taxa as distinct and described as a new species, Dysphania henthamiana, the plant here called subsp. eremaea. Unfortunately Domin incorrectly made D. myriocephala a synonym of D. littoralis (as ‘D. litoralis’) although he cited no material of that taxon; he also failed to recognise the critical characters which distinguished the species in this group. Dysphania glomulifera subsp. eremaea is, in fact, quite distinct from D. littoralis for in the latter species all flowers have three tepals firmly united at their base and deciduous together with the enclosed fruit, while the seed is sub-globular and the embryo horizontal. The illustration of ‘Dysphania litoralis’ provided by F. Mueller (1889-1891 t. 36), was evidently intended to portray the plant here referred to D. glomulifera subsp. eremaea. Unfortunately it would appear that the artist either exercised considerable licence or obtained material from several species and combined their different fea- tures in the one plant. In addition, the seed has been drawn incorrectly; the embryo 186 Nuytsia Vol. 4, No. 2 (1983) is indicated as being on the opposite side to that of the longitudinal groove (whereas it lies beneath it) and the radicular pouch is placed in an apical position (instead of basal) although the radicle itself is correctly illustrated as being inferior. The infraspecific epithet is derived from the Greek word eremia, a desert, alluding to the typical habitat of this plant. 2. Dysphania glandulosa Paul G. Wilson, sp. nov. Figure 6C. Herba prostrata; caules numerosi graciles ad basim exorientes. Folia parva, lamina anguste elliptica c. 3 mm longa. Flores in glomeratis axillaribus in fructu c. 1 mm diam. Flos terminalis hermaphroditus; tepala 3, libra; stamina 2; styli 2. Flores lat- erales feminei; tepala (1)2, libra, c. 0.5 mm longa, delicatissima, fructu imperfecte obtegentia. Pericarpium diaphanum, papillis prominentibus glanduliformibus ornato. Semen asymmetrice semi-obovoideum (latere embryonis piano), c. 0.3 mm longum, apice rotundato. Typus: 28 km NE of Nambi Homestead which is 65 km NNE of Leonora (28°8'S, 121°43'E); prostrate in red loam; 29 Aug. 1968, P. G. Wilson 7503 (holo: PERTH). Prostrate annual herb c. 20 cm diam., numerous slender stems arising from base and giving rise to many lateral branches, sparsely pilosulose all over (except inflor- escence) with simple and glandular multicellular hairs. Leaves small, slightly fleshy; lamina narrow-elliptic, c. 3 mm long; petiole equal to lamina. Inflorescence of axillary glomerules as in D. glomulifera; glomerules (in fruit) c. 1 mm diam. Ter- minal flower of glomerule bisexual; tepals 3, free, the two anterior-lateral 0.4 mm long in fruit and larger than the posterior; stamens 2 opposite anterior-lateral tepals; styles 2, slender, c. 0.1 mm long. Secondary terminal flowers (when present) similar to terminal but with two tepals and one stamen. Lateral flowers female, tepals (1) 2, free, 0.5 mm long (in fruit), very delicate and incompletely covering fruit; limb strongly cucullate, slightly inflated, emarginate with a scarious margin; claw linear ± equal to limb. Pericarp diaphanous, covered with prominent gland-like papillae. Seed erect, asymmetrically semi-obovoid (the embryo-side flat), c. 0.3 mm long, apex rounded; testa smooth, reddish brown; embryo-groove longitudinal along flat- tened side of seed; embryo lateral, erect, radicle inferior. Infructescence disintegrating at maturity into separate tepals and fruits. WESTERN AUSTRALIA: Glenorn Street, August 1938, N. T. Burbidge (PERTH); Belele Station, D. W. Goodall 3351 (PERTH). Distribution. West-central area of Western Australia. Map 18. Habitat. Loam; seasonally waterlogged areas. Dysphania glandulosa may be distinguished from D. glomulifera (to which it ap- pears to be most closely related) by its relatively small leaves, its small tepals (which do not completely cover the apex of the fruit), its strongly papillate pericarp, and by the shape of its seeds. One variant of D. glomulifera subsp. glomulifera also has pa- pillae on the pericarp but in this plant the seed is broadly compressed-obovoid or len- ticular whereas in D. glandulosa it is asymmetrically obovoid (flattened on the side bearing the embryo-groove). The specific epithet alludes to the gland-like appearance of the papillae on the surface of the pericarp. Paul G. Wilson, Tribe Chenopodieae 3. Dysphania valida Paul G. Wilson, sp. nov. 187 Herba perennis, prostrata, caulibus aliquot per radicem orientibus. Folia crassiuscula; lamina anguste oblonga vel anguste elliptica, integra, 10-15 mm longa. Flores in glomeratis axillaribus 3-5 mm diam. Flos terminalis masculinus; tepala 3 ad basim breviter connata; stamina 2; pistillodium minutum. Flores laterales feminei; tepala 2, in fructu c. 1.3 mm longa, ad basim breviter connata, limbo inflato ob- ovoideo cucullato, unguide breviter oblongo; styli 2. Pericarpium diaphanum laeve. Semen erectum, lenticulare, c. 0.6 mm altum; embryo semicircularis basalis; radicula cotyledonesque erectae. Typus: Warrego District, near Morven; hard red box-sandalwood country; common; 6 April 1941; C. T. White 11895 (holo: BRI 061936). Short-lived herbaceous perennial, prostrate, to 60 cm diameter, almost glabrous to sparsely pilose with short, multicellular, simple or gland-tipped hairs. Stems several, slender, arising from rootstock. Leaves somewhat fleshy; lamina entire, narrow-ob- long to narrow-elliptic, obtuse, 10-15 mm long; petiole slender c. half length of lamina. Glomerules of flowers dense, axillary, 3-5 mm diam. Terminal flower (of glomerule) functionally male; tepals 3 shortly united at base; stamens 2 opposite an- terior-lateral tepals; pistillode minute, c. 0.2 mm high. Lateral flowers (of glomerule) female; tepals 2, at fruiting stage with an inflated obovoid cucullate limb scarious at the margin and a short broad claw united to opposite member at base, in all c. 1.3 mm long, glabrous or sparsely pilose with gland-tipped hairs; styles 2, short and delicate. Pericarp diaphanous, smooth, adherent to seed. Seed erect, lenticular, c. 0.6 mm long; testa smooth; embryo semicircular occupying lower 2/3 of seed; radicle and cotyledons erect; raphe extending from apex to base of seed. Infructescence breaking up at maturity to release fruit enclosed in the persistent tepals. QUEENSLAND: Morven, S. T. Blake 10914 (BRI); Eulo, August 1967, M. Law (BRI). Distribution. Southern Queensland (south of 26° latitude). Map 19. Habitat. Collections have been noted as coming from “cultivation paddock in forest soil”, hard red box -sandalwood country, and “timbered, rocky, sandy hilltop”. This species is similar to D. littoralis in its habit and in its firmly united tepals; it differs most obviously in the female flowers possessing only two tepals, in the len- ticular seed and in the basal embryo. From D. glomulifera it differs in having larger tepals and again in its seed shape and embryo position. The specific epithet valida is a Latin word meaning strong, and alludes to the firmly united tepals. 4. Dysphania platycarpa Paul G. Wilson, sp. nov. Figure 6D. Herba prostrata sparse pilosulosa. Folio fere glabra; lamina elliptica 5-10 mm longa. Flores in glomeratis axillaribus in fructu c. 2-5 mm diam. Flos terminalis hermaphroditus; tepala 3, libra, c. 1 mm longa, prominente cucullata; stamina 2; styli 2, perbreves. Flores laterales feminei; tepala 1 vel 2, c. 0.7 mm longa. Semen erectum, applanatum, obovatum vel deltoideum, c. 0.4 mm longum, leviter tortum vel lateribus concavis; embryo lateralis, radicula inferior!. Pericarpium minute granulatum. 188 Nuytsia Vol. 4, No. 2 (1983) Typus: West of Western Australia — Northern Territory border at 23°10'S; edge of claypan, prostrate, 26 July 1967, A. S. George 8927 (holo: PERTH). Prostrate annual herb to 20 cm diam., sparsely pilosulose with simple or glandular multicellular hairs, numerous slender mostly simple stems arising from base. Leaves slightly fleshy, almost glabrous; lamina elliptic, obtuse, 5-10 mm long; petiole slen- der, ± equal to lamina. Glomerules of flowers dense, axillary, c. 2.5 mm diam. in fruit. Terminal flower of glomerule bisexual; tepals 3, free, c. 1 mm long, prominently cucullate, claw filiform; stamens 2 opposite and ± equal to anterior-lateral tepals; styles 2, very short; secondary terminal flower similar to but slightly smaller than terminal with two tepals and one stamen. Lateral flowers female; tepals 1 or 2, strongly cucullate in fruit with delicate white inflated limh and a filamentous claw, in all c. 0.7 mm long. Pericarp diaphanous, minutely granulate, intimately adherent to seed. Seed erect, obovate to deltoid, flat, c. 0.4 mm long, slightly twisted or with faces sunken; testa dull reddish brown; embryo lateral, radicle inferior. Infructescence readily breaking up when mature into separate tepals and seeds. QUEENSLAND: Birdsville, S. T. Blake 12224 (BRI); Currawilla, S. L. Everist 3977 (CANB). NEW SOUTH WALES: Broken Hill district, A. Morris 890 (ADW, NSW); Sturt Nat. Park, W. E. Mulham W937 (NSW). SOUTH AUSTRALIA: 13 km NW of Quinyambie H.S., N. N. Donner 3593 (AD); Abminga, E. H. Ising 2397 (AD). WESTERN AUSTRALIA: W of W.A.— N.T. border at 23°10’S, A. S. George 8927 (PERTH). NORTHERN TERRITORY: Mulga Park Stn., T. S. Henshall 134 (MEL); Andado Stn., PK Late 6826 (NT). Distribution. Central Australia; South-west Queensland, extreme western New South Wales, central, northern and north-east South Australia, extreme west-central West- ern Australia, south eastern Northern Territory. Map 20. Habitat. In clay or mud by fresh water. Dysphania platycarpa may be readily distinguished from related species (i.e. D. glomulifera, D. glandulosa, and D. littoralis) by the slender claws to the tepals and by the peculiar fruit. This fruit, unlike that of all other Dysphania species, is flat but slightly twisted (or sunken on the hroad faces), and the pericarp is minutely granu- late. The specific epithet is derived from two Greek words platy, broad, and carpos, fruit, with reference to the shape of the seed. 5. Dysphania littoralis R.Br., Prod. 412 (1810). -Chenopodium blackianum Aelien, Bot. Jahrb. Syst. 63:487 (1930) based on above, non C. littorale (L.) Thunb. (1815). Type: Port I [near Gladstone], Queensland, 6 August 1802, R. Brown (holo: BM; iso: K), see comment below. Figure 7B. Prostrate annual (or perennial?) herb with fleshy taproot, c. 30 cm diam., sparsely pilosulose with simple or gland-tipped multicellular hairs, numerous slender, mostly simple, radiating stems arising from base. Leaves somewhat fleshy; lamina elliptic Paul G. Wilson, Tribe Chenopodieae 189 obtuse, 5-10 cm long, petiole slender somewhat shorter than lamina. Glomerules of flowers axillary, dense, c. 2 mm diam. Terminal flower (of glomerule) male; tepals 3, one posterior and two anterior-lateral, shortly united at base, obovate, the limb in- flated, in all c. 1.3 mm long; stamens 2 opposite the anterior-lateral tepals, anther c. 0.3 mm long; pistillode minute c. 0.2 mm high with a pair of short styles. Secondary terminal flower (when present) similar to the terminal but with only one stamen. Lateral flowers female; tepals 3 (4), at fruiting stage obovoid, very inflated (fungoid), without a scarious margin, white, claw narrow oblong somewhat shorter than the limb, in all c. 0.8 mm long; style 1 (rarely 2), slender. Pericarp diaphanous, smooth, glossy, closely adherent to the testa. Seed napiform to depressed globular, horizontal to oblique, 0.3-0. 4 mm high, slightly furrowed over the horizontal embryo; testa smooth, reddish brown, raphe extending from radicle to base of seed. Infructescence breaking up when mature to release the seed enclosed in the three persistent tepals. QUEENSLAND: Gladstone, A. Dietrich 2417 (MEL); Curtis Is., S. T. Blake 22525 (BRI). NEW SOUTH WALES: Idalia, South Wanaaring, P. L. Milthorpe and G. M. Cun- ningham 4768 (NSW). WESTERN AUSTRALIA: Upper Murchison, 1890, /. Tyson (MELU). Distribution. Northern New South Wales; south eastern and east coastal Queensland; possibly the Murchison River district of Western Australia (see com- ment below). Map 18. Habitat. The coastal variant in tidal mud flats; the inland variant in loam in open forest. The name Dysphania littoralis has been used loosely to include D. glomulifera or, when the latter species has been recognised (usually as D. myriocephala), has been applied to the inland plant here referred to D. glomulifera subsp. eremaea. The two species are quite distinct: in D. littoralis the lateral flowers have three (rarely four) tepals which remain united at their base and the seed is turnip-shaped with a hori- zontal to oblique embryo; in D. glomulifera (and in the related species D. glandulosa, D. platycarpa, and D. valida) the lateral flowers have one or two tepals and the seed and embryo are erect. The illustration under the name ‘D. litoralis’ provided by F. Mueller in his Icon. Austral. Salsol. PL tab. 36 (1890) was evidently derived from several collections, possibly none of which were of this species. For further discussion see under D. glomulifera.. The specimen collected by Isaac Tyson from the upper Murchison River is the only record of this species from Western Australia, while it has not been recorded from the Northern Territory, South Australia, or western Queensland. The type of Dysphania littoralis was collected by Robert Brown on coastal mud- flats in the tropics near Gladstone in Queensland. Material which matches the type comes from such habitats in Queensland along its east coast. A variant very similar to the type form has been collected at inland localities in southern Queensland and north-west New South Wales and here grows in quite different habitats such as ‘mixed open forest, sandy soil’ and ‘hard red soil’. It is possible that the inland 190 Nuytsia Vol. 4, No. 2 (1983) ecotype should be recognised as a distinct taxon but, since I am unable to find clear morphological differences on which such a recognition could be based, I have in- cluded both variants under the same name. Typification. George Bentham (1870) stated that Robert Brown had collected the type while running from some Aborigines, a comment that may relate to a note in Brown’s unpublished diary in which he records an attack having taken place at Keppel Bay on 16 August 1802. However, in his manuscript Flora, Brown states that the collection was made at 'Port F on 6 August 1802, which information is also given on the labels of the holotype and isotypes in the herbaria BM and K. Port I is near the present port of Gladstone and about 50 km SE of Keppel Bay. 6. Dysphania plantaginella, F. Muell., Fragm. 1:61 {1858}.— Chenopodium plantaginellum (F. Muell.) Aellen, Bot. Jahrb. Syst. 63:487 (1930). Lectotype: Sturts Creek, Australia sub-centralis, March 1856, F. Mueller (MEL 19916) lecto. nov Fig- ure 7E. Annual herb with several prostrate to ascending stems arising from base. Stems sparingly branched, pilose with multicellular hairs. Leaves sparsely pilose and with sessile glandular hairs; lamina elliptic to broadly elliptic, entire, obtuse, 1-2 cm long, base cuneate passing into a short petiole. Inflorescence of terminal, shortly peduncu- late, very narrowly cylindrical usually erect spikes composed of continuous (or interrupted) globular dichasial cymes; bracts solitary beneath each globular cluster, with an inflated cucullate rostrate limb and a slender claw, in all c. 1 mm long. Flowers with a slender pedicel 0.3-0.5 mm long. Terminal flower (of dichasial cyme) bisexual; tepals 3, free but remaining attached to pedicel, erect, limb concave and strongly inflated with a rounded to spreading keel, ciliolate otherwise glabrous, closely enveloping seed; stamen solitary opposite an anterior-lateral tepal; style slen- der, solitary. Lateral flowers (of dichasial cyme) numerous, female, otherwise similar to terminal flower. Pericarp diaphanous, faintly areolate otherwise smooth. Seed el- lipsoidal, erect, c. 0.5 mm high; embryo erect, lateral and basal beneath a longitudi- nal groove, the radicle inferior; testa smooth, reddish brown. Infructescence breaking up when mature at base of pedicels to release seed enclosed within the three tepals. NEW SOUTH WALES: 23 km NNW of Conoble Rly Stn., J. Pickard 1943 (NSW). SOUTH AUSTRALIA: Koonamore Stn., Hj. Eichler 17177B (AD); 2 mi [3.2 km] SE of Oodnadatta, A. C. Beauglehole 20839 (AD). WESTERN AUSTRALIA: Nita Downs Stn., R. A. Saffrey 1680 (PERTH); 32 km W of Gascoyne Junction, P. G. Wilson 8424 (PERTH). NORTHERN TERRITORY: 30 mi [48.3 km] N of Willowra HS, G. Chippendale (NT4739); 6 mi [9.7 km] S of Hermannsburg Mission, P. K. Latz 793 (AD). Distribution. Central New South Wales north westwards through central and north- ern South Australia, the Northern Territory to northern and north western Western Australia. Map 18. Habitat. Sand, either coastal or on the margin of waterholes and inland saltlakes. Dysphania plantaginella is rather variable in its morphology, the dichasial cymes sometimes forming discrete globular clusters with rounded dorsal margins to the tepals, and sometimes forming an apparently continuous spike, with slightly spread- Paul G. Wilson, Tribe Chenopodieae 191 ing keels to the tepals. In addition, whereas most collections appear to have erect spikes, in a few these are quite prostrate. These variants are not clearly defined so it seems unwise to establish infraspecific taxa to accommodate them. In view of the variety of habitats in which this species is found several ecotypes might be involved. Dyspbania plantaginella is evidently closely related to D. sphaerosperma but dif- fers most obviously in seed shape and in the nature of the pericarp. In D. plantaginella the seed is ellipsoidal and the pericarp areolate (marked out in a fine reticulate pattern) whereas in D. sphaerosperma the seed is compressed-globular and the pericarp minutely granular-papillate. 7. Dysphania sphaerosperma Paul G. Wilson, sp. nov. Figure 6E-F. Herba prostrata vel ascendens; caules numerosi ad basim exorientes. Folia pilosulosa; lamina elliptica obtusa integra vel undulata 0.5-1. 5 cm longa. Inflorescentia spicata; spicae breviter pedunculata, anguste cylindraceae ad 15 cm longae, cymarum congestorum compositae. Flores fere sessiles vel pedicello crasso c. 0.1 mm longo instruct!. Flos terminalis hermaphroditus (vel femineus); tepala 3, c. 1mm longa, libra; stamen solitarium; stylus solitarius gracilis. Pericarpium diaphanum, minute granulatum. Semen compresse globulare c. 0.5 mm alte; embryo semicircularis obliques vel erectus, radicula inferiore. Tepala fructum cingentia per- sistentia. Typus: Upper reaches of Turner River, Woodstock Stn., south of Port Hedland, Western Australia; travertine ridge near river, ascendent herb, 28 April 1958, N. T. Burbidge 5924 (holo: CANB 53651; iso: AD). Annual herb with several prostrate to ascending stems arising from base. Stems sparingly branched, pilosulose with multicellular hairs; lamina elliptic, obtuse, entire or undulate 0.5-1. 5 cm long, passing into a short petiole. Inflorescence of terminal, shortly pedunculate, narrowly cylindrical spikes to 15 cm long, composed of con- gested dichasial cymes of 3 to several flowers; bracts (beneath each cyme) with an in- flated cucullate rostrate limb and a slender claw, in all c. 1 mm long. Flowers almost sessile or with a short thick pedicel c. 0.1 mm long. Terminal flower (of cyme) bisex- ual (or female); tepals 3, c. 1 mm high in fruit, free but remaining attached to pedi- cel, deeply cucullate and inflated with a rounded inflated limb and a short broad claw, glabrous except for a few sessile glands towards the margins; stamen solitary opposite an anterior-lateral tepal; style solitary, slender. Lateral flowers (of cyme) similar to terminal flower but female. Pericarp diaphanous, minutely granulate pap- illose. Seed compressed globular c. 0.5 mm high; embryo semicircular, oblique or erect, lying beneath a deep median groove, the radicle inferior; testa smooth, reddish brown. Infructescence breaking up when mature with flowers dehiscing at base of pedicels releasing the seeds enclosed in the three tepals. WESTERN AUSTRALIA: Wolf Meteorite Crater, A. C. Beauglehole 47376 (PERTH); Dirk Hartog Is., A. S. George 11413 (PERTH). NORTHERN TERRITORY: Curtin Springs Stn., T. S. Henshall 728 (NT); Lake Neale, P. K. Latz 2673 (NT). Distribution. Shark Bay area on the west coast of Western Australia eastwards to the southern area of the Northern Territory. Map 21. Habitat. Possibly always associated with gypseous or calcareous soil, frequently around salt lakes. 192 Nuytsia Vol. 4, No. 2 (1983) Dysphania sphaerosperina is similar in appearance to D. plantaginella-, it differs in having rounded inflated tepals (not keeled), a very short thick pedicel, and subglobular seeds with a minutely granulate-papillose pericarp. Several herbarium collections consist of plants with only female flowers. The specific epithet is derived from the Greek words sphaera, a sphere, and sperma, a seed, referring to the almost globular seeds. 8. Dysphania simulans F. Muell. et Tate ex Tate, Trans. Roy. Soc. S. Austral. 8:71 (1886). — Chenopodium simulans (Tate) F. Muell., Sec. Syst. Census 50 (1889); J. Black, Trans. Roy. Soc. S. Austral. 58:174 (1934). Type: Gypsum and salt flats in the spring country, Cootanoorina, M. Murray (holo: AD; iso: MEL, NSW, PERTH). C. osbornianum Aellen, Bot. Jahrb. Syst. 63:488 (1930). Type: “Koonamore Head Station, 1914”, T. G. B. Osborn (holo: G n.v.; iso: AD96630144). Annual herb with several decumbent to ascending branches arising from its base and from below the inflorescences, 10-30 cm high, pubescent with simple multicellular hairs on branches, petioles, and inflorescence axes, and sessile glandular hairs on leaves and flowers. Leaves early deciduous; lamina chartaceous, elliptic, 1-2 cm long, with 2-4 pairs of prominent obtuse lobes, apex obtuse to acumi- nate, base cuneate and passing into a slender petiole somewhat shorter than lamina. Inflorescence an erect narrowly cylindrical spike 5-20(30) cm long, 4 mm diam., sess- ile at points of branching of stem, composed of densely arranged normally 7-flowered dichasial cymes; bracts (beneath each cyme) with an inflated strongly cucullate ros- trate limb and a slender claw, in all c. 1 ram long. Terminal flower (of cyme) large and sessile, bisexual; tepals 3, cucullate, c. 1 mm long, united in lower half into an in- durated strongly papillose cup-shaped tube; stamen solitary in axil of an anterior- lateral tepal, sty'Ies 2, slender. Lateral flowers (of cyme) female otherwise similar to, although smaller than, terminal flower. Fruiting perianth cartilaginous (especially at the cup-shaped base), at first firmly attached to axis; tepal limbs produced into spreading horizontal hyaline wings, in all 2 mm high and wide. Seed erect, broadly pear-shaped, c. 0.6 mm high; embryo vertical extending along one side and around base of seed; Radicle superior occupying the small dark-coloured terminal apiculum; raphe extending from apex to base. Infructescence tardily breaking up, the seed en- closed in the cartilaginous perianth which eventually splits into its separate components. QUEENSLAND: Lake Crocker, 28 June 1939, R. L. Crocker (AD); Poeppel Corner D. E. Boyfand 237 (NSW). NEW SOUTH WALES: 63 mi [101.4 km] SE of Wanaaring, W. E. Mulham W445 VICTORIA: 28.8 km W of Nowingi, 29 July 1979, J. M. Browne (MEL). SOUTH AUSTRALIA: Koonamore Stn., M.D. Crisp 709 (AD); Oodnadatta, 8 Apr. 1950. E. E. Lord (MEL). WESTERN AUSTRALIA: 37 km N of Agnew, D. Symon 9941 (ADW); 30 km NE of Nambi HS., P. G. Wilson 7499 (PERTH). TERRITORY: 11 km SW of Erldunda Stn., T. S. Henshall 46 (NT); Mt Wedge Stn., P. K. Latz 2146 (AD). Paul G. Wilson, Tribe Chenopodieae 193 Distribution. Central Australia; south-west Queensland, north-west New South Wales, north-east South Australia, east central Western Australia, southern portion of the Northern Territory. Map 21. Habitat. Saline (halitic), gypseous, or calcareous soil, often on margin of salt lakes. The densely arranged cymes form long terminal ‘mouse-tail’ inflorescences which, because of their erect and sessile habit, give the plant a candelabrum aspect. This, and the indurated cup-shaped base to the perianths, make Dysphania simulans a very distinctive species. The illustration by Graff provided by Mueller in his Icon. Austral. Salsol. PL tab. 34 (1890) well portrays the species; however, it shows the embryo incorrectly with the cotyledons occupying the terminal apiculum whereas, in fact, this area is filled by the radicle. The orientation of the embryo is thus the reverse to that found in D. plantaginella (which has a similarly shaped seed); this is because the embryo differs in its position by 180° to that which it occupies in D. plantaginella causing the hilum to be moved near the apex of the seed and the raphe to elongate. The illustration (tab. 34) showing an incorrect orientation of the embryo appears to have been di- rected by Mueller from a specimen of A. Henry from Georgina River (MEL 19933) for there are some pencil sketches by Mueller in which he evidently instructed Graff on the fine detail of the fruit. 9. Dysphania rhadinostachya (F. Muell.) A. J. Scott, Bot. Jahrb. Syst. 100:218 (1978 ). — Chenopodium rhadinostachyum F. Muell., S. Sci. Rec. 2:98 (1882); F. Muell., Icon. Austral. Salsol. PI. t. 33 (1890) p.p. Type: Finke River, 1881, H. Kempe 413 (holo: MEL 20098). Figure 7C-D. Chenopodium rhadinostachyum var. paniculatum F. Muell. et Tate, Trans. Roy. Soc. S. Austral. 16:345 (1896) nomen. Chenopodium inflatum Aellen, Bot. Jahrb. Syst. 63:490 (1930). — D. inflata (Aellen) A. J. Scott, Bot. Jahrb. Syst. 100:218 (1978). Lectotype: Bulloo Range, 29 August 1923, MacGillivray (holo: G n.v.; iso: ADW 17774, BRI 010647) lecto. nov. Erect herb to 30 cm high with a single main stem, copiously branched above the base, moderately pilose all over with short glandular and longer segmented hairs, aromatic. Leaves thin; lamina elliptic, 10-20 mm long, undulate or with a few obtuse lobes, base cuneate and passing into a slender petiole. Inflorescence of axillary and terminal leafy or spike-like thyrses which occupy nearly the whole plant and are composed of compact dichasial cymes of 3-7 flowers subtended by a bract. Bracts of the inflorescence either foliaceous and shortly exceeding cymes or very reduced, nar- rowly ovate acuminate c. 2 mm long. Pedicels very short, turbinate, c. 0.2 mm long, all apparently arising from main axis. Flowers bisexual and female (the terminal flower of cyme always bisexual, the lateral sometimes so); tepals 4, erect, c. 1 mm long, incurved and cucullate, united in lower third, rounded or shortly and bluntly horizontally keeled on back, inflated in fruit, glabrous to hirsute, ciliate; stamen 0-1, about twice length of perianth; styles 2, slender, long exserted. Pericarp diaphanous, dull, sometimes slightly scurfy, somewhat loosely adherent to seed. Seed subglobular, horizontal to erect, c. 0.5 mm diam., slightly keeled and grooved over the semicircu- lar embryo, radicle and cotyledons superior; testa reddish brown; raphe extending from apex to base of seed. Infructescence breaking up with flower dehiscing at apex of pedicel, the fruit loosely enclosed in perianth. 194 Nuytsia Vol. 4, No. 2 (1983) Key to subspecies Tepals hirsute 9a. subsp. rhadinostachya Tepals glabrous or sparsely hirtellous, ciliate 9b. subsp. inflata 9a. subsp. rhadinostachya Tepals hirsute, rounded on back or slightly keeled. Seed oblique to erect; pericarp dull. Inflorescence much-branched with the bracts smaller than or exceeding the glomerules. QUEENSLAND: Woodstock, S. T. Blake 6499 (BRI); Mt Isa, R. L. Specht and R. W. Rogers 30 (BRI). SOUTH AUSTRALIA: Mt Ilbillee area, A. C. Beauglehole 25520 (AD); Mt Morris, Hj. Eichler 17333(AD). WESTERN AUSTRALIA: Dampier, 30 March 1971, H. Hukin (PERTH); Rawlinson Range, P. Wilson 2378 (AD). NORTHERN TERRITORY: Mt Olga, G. Chippendale (NT 4665); Mann Range, D. J. Nelson 362 (NT). Distribution. Western Queensland, northern South Australia, north-west to east- central Western Australia, and southern half of the Northern Territory. Map 19. Habitat. Typically a species of rocky slopes in skeletal soil. 9h. subsp. inflata (Aellen) Paul G. Wilson, stat. nov. — Chenopodium infiatum Aellen, Bot. Jahrb. Syst. 63:490 (1930). Type: loc. cit. Tepals glabrous or sparsely hirtellous, ciliate, rounded on back. Seed horizontal or slightly oblique; pericarp dull (or rarely glossy). Inflorescence of long slender spikes often with slightly disjunct glomerules; bracts inconspicuous. QUEENSLAND: Boulia, S. T. Blake 6475 (BRI); 51 mi [82.1 km] W of Yaraka, L. Everist 7368 (BRI). NEW SOUTH WALES: Byrock, E. McBarron 14706 (NSW). WESTERN AUSTRALIA: Mt Anderson Stn., R. D. Royce 6919 (PERTH); 74 km E of Carnarvon, P. G. Wilson 8393 (PERTH). NORTHERN TERRITORY: Stirling Swamp, T. S. Henshall 510 (NT); 110 km N of Alice Springs. J. Z. Weber 995 (AD). Distribution. Central Queensland, northern New South Wales, north-western West- ern Australia, and southern half of the Northern Territory. Map 20. Habitat. Evidently usually in skeletal soil but also recorded from ‘hard red mulga country’ in Queensland. Paul G. Wilson, Tribe Chenopodieae 195 Dysphania rhadinostachya has been confused with D. kalpari which may be dis- tinguished by its decumbent habit and erect glossy seeds. The subspecies inflata varies considerably in habit; part of this variation is due to age and environmental conditions and part to the presence of several distinct forms which are not recog- nised nomenclaturally in this paper. In Western Australia is found a plant which is superficially similar to the lectotype of subsp. inflata (from Queensland) but in which the pericarp is almost glossy; to this variant belongs the lectoparatype of Chenopodium inflatum collected by Morrison at the Ashburton River (PERTH). This variant is further discussed under D. kalpari. The characters which distinguished the two subspecies are far from constant and it is evident that these taxa grade into each other. For this reason I have not kept them as separate species as did both Aellen (1930) and Scott (1978b). The plate in Mueller (1890 op. cit.) cited above under Chenopodium rhadinostachyum appears to contain illustrations of two species; the small habit drawing on the left-hand side is probably of D. kalpari while the drawing of the in- florescence is of D. rhadinostachya. The illustrations of the seed incorrectly display the position of the embryo and it is evident that both Mueller and Robert Graff, the artist, had difficulty in establishing the orientation of this minute org an The name Chenopodium rhadinostachyum var. paniculatum does not appear to have been validly published. It was applied by Tate to a collection made by R. Helms in 1891 on the Elder Exploring Expedition and, according to the text, was collected at camp 53. 1 have seen only one specimen with that name; the label accompanying it was written by Tate and the collection was made by Helms at camp 41. This speci- men has a paniculate inflorescence typical of D. rhadinostachya but it is evident that Tate assumed it to represent a new variety because specimens collected at camp 53 were identified by him as being the typical form of Chenopodium rhadinostachyum whereas they are, in fact, the new species D. kalpari. 10. Dysphania kalpari Paul G. Wilson, sp. nov. Figure 7A. Herba annua prostrata vel decumbens. Folia lamina profunde undulata, 1-5 cm longa. Inflorescentia: spicae breviter pedunculatae, erectae, graciles, ad 15 cm longae. Flores hermaphroditi et feminei; tepala 4, erecta obovata, c. 1 mm longa in base connata, breviter hirsute; stamen 0-1; styli 2, graciles. Semen verticale compresse sphaericum, c. 0.5 mm diam.; testa laevis, nitida; embryo semi-circularis basalis, radicula cotyledonibusque superioribus. Typus: West end of Hopkins Lake (south of Sir Frederick Range), Western Aus- tralia, ± 24°20'S, 128°40'E; 1 Aug. 1962; D. E. Symon 2354 (holo: PERTH; iso: AD, ADW). Prostrate to decumbent annual herb with several stems arising from base, moder- ately pilose all over with sessile or shortly stipitate glandular hairs and longer seg- mented hairs, aromatic. Leaves thin, lamina elliptic, deeply undulate, 1-5 cm long; petiole slender, half to as long as lamina. Inflorescence of axillary and terminal, shortly pedunculate, erect, slender spikes to 15 cm long, made up of compact dichasial cymes of 5-7 flowers subtended by bract. Bracts stipitate, limb ovate, ros- trate, in all c. 2 mm long, not exceeding cyme. Flowers bisexual and female (the ter- minal flower of cyme always bisexual); pedicel very short and thick, c. 0.2 mm long. Tepals 4, erect, obovate, c. 1 mm long, united in lower third, rounded on back. 196 Nuytsia Vol. 4, No. 2 (1983) shortly hirsute; stamen 0-1, shortly exceeding perianth; styles 2, slender. Pericarp di- aphanous (so thin as to he not apparent). Seed vertical, compressed spherical with rounded margins (slightly keeled over embryo), c. 0.5 mm diam.; testa reddish brown smooth and glossy; embryo semi-circular, basal, radicle and cotyledons superior, raphe passing from upper margin to base. Infructescence breaking up when mature with flowers dehiscing at apex of pedicel, the seed loosely enclosed in perianth. QUEENSLAND: Boatman Stn., S. L. Everist 2935 (BRI); 7.8 mi [12.6 km] N of Galway Downs, G. Trapnell E58 (BRD- NEW SOUTH W.\LES: 70 km N of Bourke, S Jacobs 2061 (NSW); 8 mi [12.9 km] S of Barrier Hwy on Cobar-Ivanhoe road, C. W. E. Moore 6012 (CANB). SOUTH AUSTRALIA; 48 km W of Tallaringa Well, T. R. N. Lothian 3847 (AD); 30 mi [48.3 km] W of Welbourne Hill Stn., R. A. Perry 5534 (AD, MEL, NSW, NT, PERTH). WESTERN AUSTRALIA: Mundiwindi, B. G. Briggs 3601 (NSW); 44.5 mi [71.6 km] E of Carnegie Stn., A. R. Fairall 1999 (PERTH). NORTHERN TERRITORY; 16 mi [25.7 km] NW of Hamilton Downs, G. Chippendale (NT 1779); 12 mi [19.3 km] W of Stuart Hwy on Yuendemu Rd, J. R. Maconochie 60 (NT). Distribution. Widespread in the inland areas of Australia in all mainland States ex- cept Victoria. Map 22. Habitat. Loam flats and red sand plains; often associated with mulga {Acacia aneura). Dysphania kalpari has been confused with D. rhadinostachya from which it may be distinguished by its decumbent habit with several stems arising from the base, its erect glossy seeds, and its “rats-tail” spikes. The habitat preferences of the two species are quite different, those of D. rhadinostachya being skeletal soils often on rocky slopes while D. kalpari is found on mulga {Acacia aneura) flats or sand plains. Intergradation with Dysphania rhadinostachya subsp. inflata appears to occur, at least with the variant of the latter found in the Pilbara region of Western Australia, and it is possible that the lectoparatype of Chenopodium inflatum from the Ashburton River area is a member of such an intergrade. A note by F. T. Turvey with a collection made near Ernabella, South Australia, in 1968 (NSW 103747), states that “The natives gather the ripe seeds and make them into a meal, and at this time mix them in with the honey from honey-ants (also in season). This is relished as a food.”. On a specimen from Warburton Mission (R. A. Gould, 3 April 1967, PERTH) it is recorded that the Aboriginal name is Kalpari and that it is an important seed-food plant of ngatatjara, pitjantjatjara and pintupi natives. A further specimen (W. H. Moyle, 1 July 1958, PERTH) has with it a note to say that it is called Kalpari and that it is sought after by Aborigines in the Walter James Range to make flour. The specific epithet is taken from the Aboriginal name for the plant in central Australia. Paul G. Wilson, Tribe Chenopodieae 197 3. SCLEROBLITUM Ulbrich Ulbrich in Engler et Prantl, Nat. Pflanzenfam. ed. 2, 16c: 495 (1934); Aellen in Hegi, 111. FI. Mitt.-Eur. ed. 2, 3: 660 (1960). Type: S. atriplicinum (F. Muell.) Ulbrich. Chenopodium sect. Atriplicina Aellen, Verb. Naturf. Ges. Basel 41:99 (1931); A. J. Scott, Bot. Jahrb. Syst. 100:211 (1978). Type: C. atriplicinum (F. Muell.) F. Muell. (=S. atriplicinum). Somewhat succulent very sparsely mealy herb. Stems numerous arising from root- stock, unbranched. Leaves alternate simple, flat, the basal ones in a rosette. Flowers in compact glomerules axillary to basal and stem leaves. Tepals 4, free; stamen soli- tary in terminal flower, absent in others; ovary glabrous, stigmas 2, dorsiventral. Pericarp hard and thick. Seed erect, completely filling utricle; embryo inverted hip- pocrepiform (around apex of seed); endosperm sparse around embryo; perisperm copious, central. One species, endemic to Australia. This genus is not closely related to any other Australian genus but appears to have most in common with species of the genus Monolepis (northern hemisphere) and Chenopodium sect. Thullingia (South America). 1. Scleroblitum atriplicinum (F. Muell.) Ulbrich in Engler et Prantl, Nat. Pflanzenfam. ed. 2, 16c:495 (1934); Aellen in Hegi, loc. cit. — Blitum atriplicinum F. Muell., Trans, et Proc. Viet. Inst. Advancem. Sci. for 1854-55: 133 (1855). — Chenopodium atriplicinum (F. Muell.) F. Muell., Fragm. 7:11 (1869); F. Muell. Icon. Aust. Salsol. PI. t. 30 (1890). Lectotype: Cudnaka, Oct. 1850, F. Mueller (MEL) lecto. nov. Euxolus enervis F. Muell., Fragm. 1:140 (1859). — Amaranthus enervis (F. Muell.) F. Muell. ex Benth., FI. Austral. 5:216 (1870). Type: In planitiebus ad flumina Darling et Murray, J. Dallachy (holo: MEL). Amaranthus tenuis Benth., FI. Austral. 5:216 (1870). Type citation: ‘Lower Darling River, Herb. F. Mueller’ (iso: MEL). Annual (or short-lived perennial?) somewhat succulent herb. Taproot prominent. Stems numerous slender, prostrate or ascending, arising from rootstock, normally unbranched. Leaves very sparsely mealy with thin-walled stipitate vesicular hairs, eventually glabrous. Basal leaves in a rosette, lamina thin, 3-nerved, hastate-acumi- nate c. 3 cm long with divaricate acuminate basal lobes; Petiole slender 3 to 8 cm long with a broad clasping base; cauline leaves alternate, well separated, the upper ones sub-sessile and narrowly elliptic. Flowers very shortly pedicellate; tepals 4, free, erect, imbricate, ovate, acute, c. 1.5 mm long, somewhat glumaceous, mealy in lower half with stipitate vesicular hairs; dorsal sepal larger than others and when mature several nerved. Terminal flower of glomerule bisexual; stamen solitary opposite dor- sal sepal, filament slender, glabrous, not thickened at base; anther caudate, c. 0.7 mm long; ovary narrowly ellipsoidal, glabrous with a pair of opposite vascular traces which pass up into the dorsi-ventral long slender stigmas. Ovule completely filling ovary. Lateral flowers of glomerule female without staminodes. Fruiting perianth cartilaginous; tepals narrowly oblong acuminate, c. 2.5 mm long, thickened keeled and sigmoid at base. Utricle erect, lenticular; pericarp hard and thick; epicarp mem- branous; mesocarp white hard and rugose, endocarp dark reddish brown, crus- 22493— (s) 198 Nuytsia Vol. 4, No. 2 (1983) taceous, rugose on the outside. Seed erect, completely filling utricle and fused to the endocarp; testa membranous, embryo inverted hippocrepiform (around apex of seed) radicle inferior; perisperm central, copious, vitreous. Mature fruit shed with sur- rounding perianth. QUEENSLAND: Berbera, C. T. White 12231 (BRI); Clover Downs, R. Roe 3 (BRD- NEW SOUTH WALES: Pinipara Ck., S. Jacobs 1003 (NSW); 3 mi [4.8 km] S of Nyngan, G. M. Cunningham 766 (NSW). VICTORIA; Gunbower, Oct. 1914, E. Curtis (NSW); Lake Hindmarsh, Oct. 1899 C. Walter (NSW.) SOUTH AUSTRALIA: Yourambulla Caves, B. Copley 2241 (AD); 31 mi [49.9 km] N of Overland Corner, D. E. Symon 3681 (ADW). Distribution. Southern Queensland, central and southern New South Wales, north west Victoria and eastern South Australia. Map 3. Habitat. Heavy soils usually clay, frequently in situations that are seasonally water- logged. Although this plant has been referred to by collectors as an annual it has a sub- stantial taproot which may enable it to live through more than one season. The genus Scleroblitum was originally separated from Chenopodium largely be- cause of the cartilaginous fruiting perianth; this in itself does not appear to be gen- erically significant (especially as hardening of the tepals does not always take place), however, the pericarp is very different from that in other Australian members of the family. This pericarp has previously been interpreted as the testa and its distinctive features overlooked: the developing seed fills the ovary at a very early stage causing the testa to be in intimate contact with the ovary wall; this then presents difficulties in distinguishing the two structures. The ovary wall is discernible, shortly after an- thesis, as being composed of three layers; in the fruit these layers form very distinct structures of the pericarp, an outer membranous epicarp in which a pair of faint vas- cular strands pass up to the two style arms, a white crystalline mesocarp which be- comes hard and rugose, and a dark reddish brown crustaceous endocarp. The endo- carp is rugose on its outer surface (where in contact with the white mesocarp) and smooth on the inner surface where it fuses with the membranous testa. The whole structure is usually enclosed within the persistent cartilaginous perianth, however, in some cases (possibly due to continuing moist conditions) the tepals remain small and glumaceous. Bentham (1870) described this species under three names, Chenopodium atriplicinum, Amaranthus enervis and A. tenuis. The last two names were based on material in which the fruiting perianth had not hardened. He also described the plant (C. atriplicinum) as being slightly glandular-pubescent, a description repeated by Black (1924, 1948) and by Scott (1978a). It was evidently for this reason that Scott placed Chenopodium sect. Atriplicina in subgenus Ambrosia, along with the glandular species of the genus, instead of in subgenus Chenopodium, in which are placed the species with a mealy indumentum. In the latter subgenus, the sect. Atriplicina comes closest to the sect. Thullingia in which the only species, C. antarc- ticum (J. D. Hook.) J. D. Hook, is very similar, it also being a prostrate sparsely mealy plant with a basal rosette of leaves, simple stems, glomerules of flowers with 2 Paul G. Wilson, Tribe Chenopodieae 199 to 4 tepals and solitary stamens, and erect seeds with a subannular embryo. The genus Scleroblitum is, in addition, very similar in fruit and seed morphology to the northern hemisphere genus Monolepis to which it is probably also closely related. 4. EINADIA Raf. Rafinesque, Flora Telluriana 4:121 (1838); Dalla Torre et Harms, Gen. Siph. 145 (1900), pro syn. sub Suaeda; Ulbrich in Engler et Prantl, Nat. Pflanzenfam. ed. 2, 16c: 555, 558 (1934) pro syn. sub Suaeda sect. Schanginia\ A. J. Scott, Feddes Repert. 80:3 (1978). Type: Einadia linifoUa (R.Br.) Raf. Chenopodium sect. Polygonoidea Aellen, Feddes Repert. 69:69 (1964) et in Hegi, 111. FI. Mitt.-Eur. Ed. 2, 3/2:577(1960) sine desc. latina. Type: Chenopodium poly- gonoides (Murr) Aellen. Herbaceous or weakly woody perennials, often prostrate or scandent, branching from the base. Branches slender, ridged. Leaves opposite or alternate, petiolate, lin- ear to broadly hastate, c. 2-3 cm long, flat and slightly fleshy, ± covered when young (at least below) with spherical vesicular hairs. Flowers small, gynomonoecious in small cymose clusters forming racemose or open panicles, often congested; pedicels (when present) constricted at apex. Terminal flower of cluster bisexual; lateral flowers female without staminodes. Tepals (4) 5, herbaceous, sometimes succulent in fruit. Stamens 1-3, often spreading at anthesis with the subtending tepal, ± equal to tepals, glabrous; filament slightly pulvinoid at base; disc absent. Ovary glabrous; style absent or very short; stigmas 2, slender, papillose. Fruit a nut with thin dry pericarp or depressed globose (rarely ovoid) berry with inflated pericarp (succulent outside, water within), not enveloped by perianth. Seed horizontal, depre.ssed globose to lenticular with rounded margin; testa crustaceous; embryo annular, radicle centrifugal; perisperm central and copious. A genus of six species endemic to Australia and New Zealand. Rafinesque (1838 op. cit.), in his original description, segregated the genus Einadia from Rhagodia on the basis of the presence of only one or two stamens instead of five. In this he was relying solely on Robert Brown’s descriptions of the seven species of Rhagodia since he had seen no material. Brown (1810) only in R. linifolia referred to the stamen number (as being 1 or 2), evidently he assumed it to be 5 in the other species as indicated in his generic description. Rafinesque, therefore, only transferred this one species to Einadia and was unaware of its close relationship (and common stamen number) with R. nutans and R. hastata which had also been de- scribed by Brown. The generic name Einadia was subsequently ignored or misap- plied until Scott (1978a) took it up. Although Scott recognised the marked difference between those species of Rhagodia with polygamous flowers and reduced stamen number (which he placed in Einadia) and those with dioecious flowers and five st am ens (which he retained in Rhagodia), he did not comprehend the close relation- ship of the Einadia group with those Australian and New Zealand species of Cheno- podium that have similar floral characters and similar habit but which lack a succu- lent pericarp. The similarity is such that with non-fruiting material it is often im- possible to recognise to which genus it should be assigned. The difficulty is enhanced by the fact that the succulent pericarp does not always develop in the group of species with normally baccate fruits and, further, that hybridization appears to take place between related species placed in the different genera. I am therefore ex- panding the genus Einadia to include those Australian and New Zealand species of 200 Nuytsia Vol. 4, No. 2 (1983) Chenopodium and Rhagodia with a herbaceous perennial habit, small flat leaves with discrete vesicular indumentum and polygamous 5-merous flowers with reduced stamen number. The distinction between Einadia and Chenopodium is not easy to define, partly because of the great variability found in the latter genus. Einadia comes closest to a group of species in Chenopodium sect. Leprophyllum (e.g. C. hubbardii) and another in sect. Desertorum (e.g. C. desertorum). From the former section the species of Einadia differ in being perennials (not annuals) and in having 1-3 (not 5) stamens, while from the latter section they differ in absence of disc (pubescent disc in sect. Desertorum) and again reduced stamen number. The species of Einadia differ from those of Rhagodia sensu stricto in being her- baceous usually sprawling perennials with gynomonoecious flowers, a reduced stamen number and no disc. The fruit, if succulent, is “inflated”, that is it has a fleshy outer and a watery inner layer; or drying it does not become viscous. The in- dumentum is of spherical hairs (which may collapse to become saucer — or shallowly cup-shaped) and not of irregularly shaped hairs, nor does it collapse to form a scaly sheen as it does in some Rhagodia species. It is not clear why Dalla Torre et Harms (1900 op. cit.) made Einadia a synonym of Suaeda-, possibly they assumed, as did Ulbrich (1934 op. cit.), that Einadia linifolia was a synonym of Suaeda linifolia Pallas, which is not the case; the two species are very distinct as are their respective genera. Key to species 1. Fruit succulent 2 Fruit dry 2. Leaves deltoid or broadly hastate with rounded apex, dark green . . 5. E. hastata Leaves linear to ovate, deltoid, narrowly hastate or sagittate, acute or obtuse. grey to green 3 3. Leaves ovate, obtuse, base usually sagittate (N.Z.) 2. E. triandra Leaves linear to broadly triangular, base often hastate 1. E. nutans 4. Tepals in fruit with a hard circular (often black) limb; leaves narrowly oblong- elliptic to narrowly hastate 6. E. polygonoides I epals in fruit oblong to obovate, herbaceous or indurated; leaves broadly ovate or elliptic to orbicular or deltoid 5 5. Leaves broadly elliptic to orbicular (N.Z.) 4. E. allanii Leaves broadly ovate to deltoid, often hastate at base 3. E. t'rigonos 1. Einadia nutans (R.Br.) A. J. Scott, Feddes Repert. 89:3 (1978).— Rhagodia ^tans R.Br., Prod. 408 (1810). Lectotype. Ad rupes saxa in Oyster Cove in fluvio Derwent, Feb. 1804, R. Brown (holo: BM) lecto. nov. Figure 4E. R. linifolia R.Br., Prod. 408 (1810). — E. linifolia (R.Br.) Raf., FI. Tellur. 4:121 (1838). Lectotype: Broad Sound, Sept. 1802, R. Brown (holo: BM; iso: K, NSW) lecto. nov. R. chenopodioides Moq., Chenop. Monogr. Enum. 11 (1840). Lectotype: Port Jackson, 1830, Gaudichaud (holo: P) lecto. nov. Chenopodium australasicum Moq., Chenop. Monogr. Enum. 20 (1840) Type ci- tation: “Nova Hollandia, Caley’\ Type: P, n.v. Paul G, Wilson, Tribe Chenopodieae 201 Scrambling perennial with woody base. Branches slender, ridged, sparsely covered when young with small discrete bladder hairs. Leaves opposite or alternate, lamina chartaceous to sub-coriaceous, linear to elliptic or narrowly to broadly triangular, en- tire 10-20(40) mm long, rounded to truncate or hastate at base, sparsely to moder- ately mealy when young, eventually glabrescent (at least above), abruptly petiolate. Inflorescence a narrow racemose panicle 2-4 cm long; flowers in glomerules, either compact or pedicellate, if terminal then bisexual, otherwise usually female. Perianth depressed globular, 0.7-1.5 mm diam., not enveloping the ovary, mealy with bladder hairs to glabrescent; tepals (4)5, oblong, somewhat enlarging and becoming red, fleshy and sometimes reflexed in fruit. Bisexual flower: stamens 1-2(3), spreading or erect at anthesis; filament ligulate, thickened at base, glabrous; anthers included, c. 0.3 mm long; disc absent; ovary ovoid, glabrous, stigma 2 sessile or subsessile, divari- cate, terete, 0. 1-0.2 mm long. Female flower: staminodes absent; ovary as in bisexual flower. Fruit: pericarp inflated succulent, orange or red, depressed (or narrowly ovoid), c. 4 mm diam., exceeding perianth, surface faintly colliculate; seed lenticular with rounded margin, 1-1.5 mm diam.; testa rugulose to areolate-reticulate, black. Distribution. Central and eastern Australia including Tasmania. This species exhibits considerable variation with several of the variants having broad regional distributions. In many cases these variants are partially sympatric although evidently ecologically separated. Transition between the variants is readily apparent from study of the herbarium material. Only four of the variants have been recognised in this paper leaving a large number of specimens which it is only possible to determine to the species level. To attempt a more complete taxonomic analysis would appear to be unwarranted due to the considerable plasticity of the plants, and their transition in the field from one variant to another. Vegetative or non-fruiting material is sometimes difficult to distinguish from E. trigonos subsp. trigonos and specimens of the two species were confused by Bentham (1867) and by later authors; it seems likely, from the observation of herbarium material, that the two hybridize but the study of field evidence is required to support this. Hybridization between E. nutans and E. hastata may also occur since several collections appear to be intermediate in morphology. The name Rhagodia nutans was applied for some time to the New Zealand plant now known as Einadia triandra (e.g. by Cheeseman 1884 and 1914). The latter species is very similar to some variants of E. nutans and, if native to Australia, would probably be included with it under the earlier name E. triandra. One of the variants of E. nutans found in southern Australia is so similar to the New Zealand plant as to make identification somewhat dependent on locality data. Einadia nutans subsp. nutans is found in New Zealand as a recent introduction and is there very distinct in appearance and ecology from the native E. triandra. It forms dense hummocks or scrambles over hedgerows so as to kill the surrounded shrubs (A. J. Healy in sched. CHR 91541). The characters by which E. nutans may be distinguished from E. trigonos are discussed under the latter species. The name Rhagodia chenopodioides Moq. was considered by Bentham (1870), who had not seen the type, to be possibly a synonym of R. billardieri (= R. baccata) and was included by Scott (1978a) under this latter name. The type is, however, a nar- row-leaved varient of Einadia nutans subsp. nutans. The terminal flower of a branch of the inflorescence is normally bisexual but may be functionally male through the weak production of stigmas. In most specimens studied this flower had normal stigmas and produced a seed. 202 Nuytsia Vol. 4, No. 2 (1983) Key to subspecies 1. Leaves linear to narrow oblong Ic. subsp. linifolia Leaves either broader than above or hastate to sagittate at base 2 2. Leaves narrowly sagittate, sub-coriaceous; seed c. 1.5 mm diameter (central Australia) Id. subsp. eremaea Leaves narrowly to broadly hastate or deltoid, thin; seed c. 1 mm diameter ... 3 3. Leaves narrowly hastate; berry ovoid lb. subsp. oxycarpa Leaves variable in shape; berry depressed globular (if ovoid then leaves deltoid-hastate) la. subsp. nutans la. subsp. nutans R. nutans var. parvifolia Moq. in DC., Prod. 13/2:54(1849). Lectotype: Ex herb. Fee (G-DC, photo seen) lecto. nov. Chenopodium triangulare subsp. convolvulinum Murr, Allg. Bot. Z. Syst. 16:56 (1910).— C. triangulare var. convolvulinum (Murr) Maiden, Census N.S.W. Plants 66 (1916). Syntypes: Stonehenge, Dec. 1899, J. H. Maiden (NSW 33761); Jenolan Caves, Dec. 1899, W. F. Blakely (NSW 142922). Lectotype: Stonehenge, Dec. 1899, J. H. Maiden (NSW 33761) Lecto. nov. ? R. nutans var. fallacina Domin, Biblioth. Bot. 89: 616 (1921). Type: Cheltenham, Victoria, Domin IV. 1910, n.v. R. chenopodioides Moq., op. cit. (1840). Leaves chartaceous, hastate to deltoid and shortly lobed at base. Inflorescence slender or branched near base; fruits sessile or pedicellate. Berry depressed globular; seed variably reticulate, c. 1 mm diameter. QUEENSLAND: Tballon, R. Roe 26 (CANB); 10 mi [16.1 km] N of Wondoan, N. H. Speck 1961 (NSW). NEW SOUTH WALES: 5 mi [8 km] S of Cobar, C. W.E. Moore 4480 (CANB); Brush Island, 5 Sept. 1937, F. A. Rodway (NSW). VICTORIA: 11.6 km from Dargo, E. M. Canning 3937 (CBG); Nandaly, W. W. Watts 465 (NSW). TASMANIA: Lewisham, 3 Mar. 1952, W. M. Curtis (HO); Carlton Bluff, 19 Feb. 1959, W. D. Jackson (HO). SOUTH AUSTRALIA: Vihlpena Pound, A. E. Orchard 2594 (AD); Oraparinna Nat. Park, J. Z. Weber 2578 (AD). NORTHERN TERRITORY: Mt Le Hunte, P. K. Latz 913 (NT); 30 km S of Alice Springs, R. Pullen 10556 (CANB). Distribution. Central and south-eastern Australia including Tasmania. Map 23. Habitat. Occupying a wide range of habitats but usually in loam or heavy soil. This polymorphic subspecies comprises, apart from the type variant, those vari- ants not included in the other three subspecies. In New South Wales it grades to the north into subsp. linifolia and to the north-west into subsp. eremaea. However, plants with the leaf shape of subsp. nutans are also found in northern New South Paul G. Wilson, Tribe Chenopodieae 203 Wales and southern Queensland where plants intermediate between these and subsp. linifolia also occur. A variant of subspecies nutans very similar in appearance to the east coast plant also occurs in the southern portion of the Northern Territory and in adjacent regions of South Australia; it is here sympatric with subsp. eremaea although probably ecologically isolated being an inhabitant of the ranges rather than the sandplains. No putative hybrids between the two subspecies in this area have been seen. A variant of Einadia nutans found in south-east Queensland appears to differ significantly from the other variants referred to subsp. nutans. It is a more delicate plant than typical subsp. nutans with very slender branches. The leaves are predomi- nantly opposite, thin, with the lamina bluntly deltoid-hastate, and c. 10 mm long in the south-east becoming larger (c. 15 mm long) farther north. The inflorescence is slender with fruits on long slender pedicels c. 5 mm long. The tepals enlarge so as sometimes to almost cover the young fruit but eventually are reflexed. The berry in some specimens appears to be ovoid in shape as in subsp. oxycarpa but mature fruit is not usually present and no collector has commented on its appearance. The follow- ing specimens are examples of this ‘delicate’ variant in eastern Queensland: 2 mi [3.2 km] S of Inverai School, R. W. Johnson 432, 433 (BRI); Bybera, C. T. White 10387, 10405 (BRI); Mungallala, C. E. Hubbard 6025 (BRI); Hannaford, L. Pedley (BRI, CANB); Chinchilla, A. Beasley 34 (BRI); Woodlands, T. J. McDonald 196 (BRI); Coppersmith Rock, G. Tandy (BRI). lb. subsp. oxycarpa (Gauba) Paul G. Wilson stat. et comb. nov. — Rhagodia nutans var. oxycarpa Gauba, Viet. Nat. 65:167 (1948). Type: Near Loveday, South Australia, 28 May 1943, E. Gauba (holo: MEL 80685). Scrambling sub-sbrub to 1 m high. Leaves somewhat leathery, narrowly hastate acute, c. 15 mm long, lateral lobes narrow, acute. Inflorescence branched; flowers and fruits sessile. Fruiting perianth fleshy and reflexed, red; berry ovoid, c. 4 mm long, pericarp colliculate; seed horizontal to erect c. 1 mm diameter, testa moderately re- ticulate. NEW SOUTH WALES: 56 km N of Wentworth, 29 Aug. 1969, A. Rodd (NSW); 10 km ESE of Mungindi, A. Rodd 1070 (NSW). SOUTH AUSTRALIA: 2 mi [3.2 km] NW of Lake Bonney, B. Copley 2577 (AD); Koonamore, M. Crisp 565 (AD). Distribution. New South Wales and South Australia. Map 24. Habitat. Often on sandhills but also recorded in other situations. This subspecies cannot be readily distinguished when in flower from some variants of the polymorphic subspecies nutans; it usually has thicker and narrower leaves which have narrow divaricate basal lobes, however, since leaf characters are notoriously plastic in this genus it is really only by the ovoid fruit that subsp. oxycarpa may be positively identified. According to M. D. Crisp (pers. comm.) subsp. oxycarpa is edaphically segregated from subsp. nutans, the former being found on the top of sand dunes and the latter on the flats. While this is evidently the case in some areas, over its total range subsp. oxycarpa occupies a variety of habitats. 204 Nuytsia Vol. 4, No. 2 (1983) The wide and scattered distribution of subsp. oxycarpa suggests that it may rep- resent a fruit-form which appears sporadically in populations of subsp. nutans, but extensive field and experimental work is required to establish the biological situ- ation. Ic. subsp. linifolia (R. Br.) Paul G. Wilson, Comb, et stat. nov. — Rhagodia linifolia R.Br., Prod. 408 (1810). — Einadia linifolia (R.Br.) Raf. l.c. (1838); Ulbrich in Engler et Prantl. Nat. Pflanzenfam. ed. 2, 16c; 558 (1934) pro syn. sub Suaeda linifolia Pallas. — E. nutans var. linifolia (R.Br.) A. J. Scott, Feddes Repert. 89:4 (1978). Lectotype: see above. Chenopodium australasicum Moq., op. cit. (1840), see note below. Prostrate or scrambling perennial. Leaves thin, linear to narrow-oblong. Inflorescence slender; fruits frequently slender pedicellate. Tepals enlarged and spreading or reflexed in fruit; berry depressed globular; seed finely reticulate, 1 mm diameter. QUEENSLAND; Noondoo Stn, S. L. Everist 829 (BRI); East of Condamine, W. T. Jones 3733 (CANB). NEW SOUTH WALES; Warrah, Mar. 1920, W. W. Froggatt (BRI); 5 mi [8 km] from Aberdeen, R. Story 1092 (CANB). VICTORIA; Echuca, H. King 4 (MEL). Distribution. Eastern Queensland and New South Wales, northern Victoria. Map 24. Habitat. Predominantly found on heavy soils. The typical variant of this subspecies is found in north east Queensland and east- ern New South Wales. In New South Wales it grades into subsp. nutans and to the west into subsp. eremaea. Some inland variants have rather thick leaves and com- pact inflorescences, possibly as a result of soil salinity, but this is not evident from the label data on the herbarium specimens. I have not seen the type of Chenopodium australasicum, however, Moquin’s de- scription corresponds to that of subsp. linifolia and it was with this taxon that he later (1849) synonymized the name. A variant of Einadia nutans with very narrow leaves is found in N.W. Victoria and south central New South Wales but the material I have seen is too incomplete to de- termine its precise affinities. I have included this variant within subsp. linifolia on the basis of its leaf shape. The synonymizing of E. linifolia under Suaeda linifolia Pallas by Ulbrich (1934) was presumably made on the assumption that they were nomenclatural synonyms. This is not the case and I do not know why Ulbrich came to this conclusion. Id. subsp. eremaea Paul G. Wilson, subsp. nov. Herba perennis leviter scandens. Rami graciles sulcati. Folia opposita vel alterna, cinerea; lamina aliquantum coriacea, anguste triangularis ad basim sagittata, 15- 40 mm longa. Flores fructusque sessiles. Baccus depresso globularis. Semen c. 1.5 mm diam.; testa valde alveolata. Paul G. Wilson, Tribe Chenopodieae 205 Typus: Close to Ayers Rock, Northern Territory, sandy soil, 25 Aug. 1973, A. A. Munir 5132 (holo: AD 97342352; iso: DNA, NT). Scrambling perennial herb to 1 m high with slender sulcate branches. Leaves op- posite or alternate, grey; lamina somewhat coriaceous, narrowly triangular, sagittate at base, 15-40 mm long. Inflorescence paniculate, branched near base. Flowers and fruit sessile. Berry depressed globular; seed c. 1.5 mm diameter, testa strongly alveo- late. QUEENSLAND: Earlstoun Stn., S. T. Blake 5480 (BRI); Boorara, Vk. J. Bisset 253 (BRI). NEW SOUTH WALES: Tundulya, C. W. E. Moore 4559 (CANB). SOUTH AUSTRALIA: 35 km W of Mabel Creek, T. R. N. Lothian 2783 (AD); 48 km W of Todmorden Stn., T. Reichstein 24 (AD). WESTERN AUSTRALIA: 8 km S of Giles, R. H. Kuchel 161 (AD); Giles Settle- ment, R. Hill 1369 (AD). NORTHERN TERRITORY: Tanami Sanctuary, T. S. Henshall 1256 (NT); Petermann Creek, N. M. Henry 576 (NT). Distribution. Central and Western Queensland, north western New South Wales, northern South Australia, east-central Western Australia, the Northern Territory. Map 25. Habitat. Generally in sand. This subspecies is sympatric with subspecies nutans in parts of the Northern Ter- ritory and in the Musgrave Ranges area of South Australia, where the two evidently occupy different ecological niches (subsp. eremaea in sand and subsp. nutans on hilly country) and I have seen no suggestion of intergradation between the two in that re- gion. Subspecies eremaea is a more robust plant than any of the other subspecies and is bigger in all its parts; it grades to the east into subsp. linifolia and to the south and south-east into subsp. nutans. 2. Einadia triandra (G. Forster) A. J. Scott, Feddes Repert. 89:5 (1978). — Chenopodium triandrum G. Forster, FI. Ins. Austr. 21 (1786); Moq., Chen. Mon. Enum. 25(1840) p. p.; J. D. Hook., FI. Nov.-Zel. 1:212 (1857) p.p.; J. D. Hook., Handb. N. Zeal. FI. 230 (1864) p.p. — Rhagodia triandra (G. Forster) Aellen, Candollea 8:5 (1939); Allan, FI. N. Zeal. 1:231 (1961). Type: Queen Charlotte Sound, New Zealand, J. R. and J. G. A. Forster (holo: BM; iso: K n.v. fide Allan 1961). Fig- ure 5B. [Rhagodia nutans auct. non R.Br.: Cheeseman, Man. N. Zeal. FI. ed. 2, 404 (1925)] Spreading or scrambing perennial with woody (sometimes rhizomatous?) rootstock and numerous slender matted stems. Branches striate, mealy when young. Leaves opposite or alternate, mealy or glabrescent; lamina thin, ovate, 5-10 mm long, entire, apex obtuse and minutely apiculate, base truncate to shortly sagittate; petiole slen- der c. 4 mm long. Inflorescence of glomerules in terminal or axillary slender racemose panicles c. 5 cm long in fruit. Flowers globular 1-2 mm diam.; tepals ovate, flat or cu- cullate, enlarging and becoming fleshy in fruit, mealy outside; stamens 0-3, spreading 206 Nuytsia Vol. 4, No. 2 (1983) at anthesis; filament thickened at base; stigmas 2 sessile, slender c. 0.5 mm long. Fruit a berry; pericarp red, globular, succulent c. 4 mm diam. Seed depressed sphe- roid, c. 1.7 mm diam, testa prominently reticulate. NEW ZEALAND: N of Kaikoura, R. Melville 5833 (CHR); Near Cape Campbell, W. R. Sykes 500/70 (CHR). Distribution. New Zealand; ‘from near north Cape to a little south of lat. 44°’ fide Allan 1961. Habitat. ‘Rocky and gravelly coastal places’ fide Allan 1961. I have not seen the type of Einadia triandra but the description of the leaves given by J. G. Forster agrees with current concepts and with the plant described under the name Rhagodia triandra by Aellen (1939). The transfer of Chenopodium triandrum to Rhagodia was intimated by F. Mueller in 1886 and 1887 when he wrote ‘The Chenopodium triandrum of New Zealand is transferable to the genus Rhagodia', but I cannot find any publication in which he made the combination. The name Chenopodium triandrum G. Forster was considered by Murr (1910) to include C. microphyllum F. Muell., and his synonymy was accepted by Maiden and Betche (1916). The two species are, in fact, quite distinct and differ, among other characters, in stamen number and fruit type, although it is probable that Murr had specimens of C. allanii ( = Einadia allanii) in mind rather than C. triandrum when he suggested the synonymy. Einadia triandra is very similar to some variants of E. nutans and if it were not for the fact that the one is endemic to New Zealand and the other to Australia a subspecific status for E. nutans would probably be more suitable. The leaves of E. triandra have consistently an ovate shape obtuse at apex and truncate at base usually with short downwardly directed lobes (sagittate) whereas in E. nutans the leaves tend to be triangular, acute at apex and hastate at base (i.e. with divaricate lobes). In E. nutans the seed is lenticular with rounded margin and the surface rugulose to deeply reticulate; in E. triandra the seed is depressed-spheroid (i.e. thicker than in E. nutans) and consistently deeply reticulate. Prior to the paper by Aellen (1939) material of Einadia triandra had usually been referred to Rhagodia nutans and the name Chenopodium triandrum incorrectly ap- plied to the species described by Aellen as C. allanii. The last species may be dis- tinguished by its thin dry pericarp; it also differs from all related species found in New Zealand by its entire, elliptic to orbicular leaves. The recording by Sykes (1977) of E. triandra (as Rhagodia triandra) in the Kermadec Islands was based on several collections some of which are of a variant of E. trigonos, q.v. while others are of an undescribed species. 3. Einadia trigonos (Roemer et Schultes) Paul G. Wilson, comb. nov.— Chenopodium trigonon Roemer et Schultes, Syst. Veg. 6:275 (1820) based on following.— C. triangulare R.Br., Prod. 407 (1810) nom. illeg. non Forsskal (1775). Type: Nepean and Hawkesbury Rivers, R. Brown (holo: BM). C. stellulatum Aellen in Probst, Mitt. Naturf. Ges. Solotb. 8:57 (1928). Type: Wollkompost der Kammgarnfabrik Derendingen, 1916, R. Probst, n.v. Paul G. Wilson, Tribe Chenopodieae 207 Weak prostrate to erect straggling perennial herb branching from the base. Branches slender, faintly striate, glabrous or sparsely mealy when young. Leaves alternate or subopposite, thin, lamina broadly ovate to deltoid (or hastate with short spreading lobes), 8-15 mm long, glabrescent above, sparsely to moderately mealy below with white spherical vesicles; petiole slender, short or equal in length to lamina. Inflorescence a congested to slender terminal and axillary panicle with numerous sessile flowers aggregated into glomerules. Flowers, if terminal to a partial cluster, bisexual; the others female. Perianth depressed globular 0.5-1 mm diam.; tepals broadly ovate to obovate, spathulate or linear, often cucullate, sparsely mealy or glabrous. Bisexual flower, stamen solitary, divaricate at an thesis, filament thickened (pulvinoid) at base; ovary depressed, papillose; stigmas slender, spreading, ± sessile. Female flower: stamens and staminodes absent, otherwise as in bisexual flower. Fruit: tepals narrow to broad oblong or obovate, herbaceous or indurated, smooth, if indurated then with a slender claw and cucullate limb enclosing and per- sisting around fruit; pericarp dry (?slightly succulent in subsp. leiocarpa), thin, often black and usually prominently papillose. Seed lenticular with rounded margin, c. 1 mm diam.; testa slightly granular (or striate in subsp. leiocarpa). In habit and leaf shape E. trigonos is similar to some variants of E. nutans but in the latter species the leaves are usually more prominently hastate. The tepals of E. trigonos vary from being broad-oblong to linear. The variant with broad tepals (in subsp. trigonos) when in flower is difficult to distinguish from some variants of E. nutans and even fruiting material can be troublesome since, although the latter species usually has baccate fruit, the succulence does not always develop or is not ap- parent in dried material. A further distinction is found in the seed which in E. trigonos is granular and the pericarp smooth to strongly papillate (but see also subsp. leiocarpa); in E. nutans the seed is alveolate to rugulose and the pericarp smooth or colliculate (i.e. with low convex markings). It appears probable, on the basis of herbarium material, that the two species do hybridize. To separate Einadia trigonos from E. hastata is also sometimes difficult in the ab- sence of fruit, however, the leaves of E. hastata are usually darker green, thicker, and they are nearly always opposite, whereas in E. trigonos they are predominantly alter- nate. Key to subspecies 1. Tepals spathulate to linear; pericarp strongly papillose, frequently black 3b. subsp. stellulata Tepals ovate or elliptic to obovate; pericarp almost smooth 2 2. Leaves broadly hastate or deltoid with rounded apex 3a. subsp. trigonos Leaves hastate to narrow hastate, acute 3c. subsp. leiocarpa 3a. subsp. trigonos Plant prostrate to decumbent. Leaves broadly hastate or deltoid with rounded apex. Tepals ovate to obovate, herbaceous in fruit. Pericarp almost smooth. NEW SOUTH WALES: Moona Plains, Feb 1899, S. R. Crawford (NSW); Conjala, May 1899, W. Heron (NSW). VICTORIA; Mt Tingaringy, A. C. Beauglehole 35726 (MEL); Melbourne, 1886, J. Minchin (MEL). 208 Nuytsia Vol. 4, No. 2 (1983) NEW ZEALAND: Coppermine Is., Mauraki Gulf, 25 Oct. 1968, M. A. and I. M. Ritchie (CHR); Little Sister Is., 25 Oct. 1973, B. D. Bell (CHR); Curtis Island, Kermadec Group, W. R. Sykes 923/K (CHR). Distribution. Southern Victoria and extreme eastern New South Wales to just north of Walcha. New Zealand (an atypical variant). Map 26. Habitat. Often near water but also in open country or forested areas. The typical variant of this subspecies has thin, deltoid to broadly hastate leaves and broadly ovate tepals, it is found in the extreme south east of Victoria north to an area just north of Sydney. Farther north still the tepals become narrower and the plant grades into subsp. stellulata which has linear tepals. The presence of Einadia trigonos subsp. trigonos in New Zealand is probably due to its relatively recent introduction. The earliest collection seen by me is dated 1926 although only the specimens in herb. CHR have been studied. In New Zealand it is variable in leaf form and whereas some of these variants are found in Australia others are distinct and approach in aspect, and in flowers and fruit, E. allanii, a species endemic to New Zealand, suggesting that hybridization may be taking place. Sykes (1977) recorded Rhagodia triandra from several islands in the Kermadec Is. group but the specimens I have seen from these islands which have been so anno- tated either belong to the New Zealand variant of E. trigonos subsp. trigonos or to an undescribed species. Sykes also recorded Einadia allanii (as Chenopodium allanii), based on the presence of its seed in the crop of a parakeet; since, however, the seed of E. allanii is virtually indistinguishable from that of E. trigonos it seems likely that they came from plants of the latter species which on some islands was recorded as abundant. The New Zealand specimens of subsp. trigonos have flowers with broad tepals which enlarge with the fruit and almost envelop it at maturity; many also have pur- plish stems and leaf margins, characters not observed (or not noted) on Australian specimens. 3b. subsp. stellulata (Benth.) Paul G. Wilson, comb, et stat. nov. — Chenopodium triangulare var. stellulatum Benth., FI. Austral. 5:161 (1870).— C. stellulatum (Benth.) Aellen, Verh. Naturf. Ges. Basel 41:93 (1931) nom. illeg. non Aellen (1928). Type citation: “New England, C. Stuart.” Lectotype: River Severn. C. Stuart 44 (MEL 80677) lecto. nov. C. stellulatum Aellen in Probst, op. cit. Prostrate to spreading herb with stout taproot. Inflorescence congested when in flower, elongating in fruit to a slender racemose panicle with disjunct glomerules of flowers. Tepals spathulate to linear, frequently indurated in fruit. Pericarp strongly papillose, frequently black. QUEENSLAND: Waterford, W. Jones 3345 (CANB); Cooranga North, C. T. White 2492 (BRI). NEW SOUTH WALES: Grafton, H. Henholz (NSW); Swan Bay, 8 June 1957, L. A. S. Johnson (NSW). NEW ZEALAND: Tai Tapu, Canterbury, A. J. Healy 54/123 (CHR); Between Halswell and Tai Tapu, 24 Aug. 1949, S. E. Archer (CHR 68003). Paul G. Wilson, Tribe Chenopodieae 209 Distribution. South-east Queensland and eastern New South Wales. A recent introduction in New Zealand. Map 27. Habitat. In partial shade frequently in moist situations in heavy soil. A ready colon- iser of disturbed situations. The vernacular names of Tar vine and Fish weed applied to this plant are a reflection of the strong fish smell possessed by some variants; it has been reported to cause cows to produce tainted milk (C. T. White 1921 as Chenopodium triangulare). As a recent introduction it is stated to be well established along a roadside near Halswell in the South Island of New Zealand (A. J. Healy 1954 in sched.) 3c. subsp. leiocarpa Paul G. Wilson, subsp. nov. Herba perennis erecta, ad basim ramificans. Folia alterna; lamina hastata vel anguste hastata acuta, c. 15 mm longa. Pericarpium laeve, translucens (non denigratum). Semen c. 1.2 mm diam., testa striata. Typus\ Pilliga, New South Wales, Nov. 1932, H. M. R. Rupp 5 (holo: NSW 142708). Erect perennial herb branching from base. Leaves alternate; lamina hastate to nar- row hastate (upper ones narrow elliptic), c. 15 mm long, acute, noticeably mealy below; petiole slender, c. half length of lamina. Tepals broadly elliptic to obovate. Pericarp smooth, translucent (not blackened), evidently herbaceous when mature. Seed c. 1.2 mm diam.; testa glossy, radially striate. NEW SOUTH WALES: Gurley, Nov. 1914, E. Breakwell (NSW); Boggabri, R. H. Cambage 3655 (NSW); 6.5 mi [10.5 km] W of Wee Waa, 19 Dec. 1966, L. A. S. Johnson (NSW); Moree, Feb. 1907, J. T. Scott (NSW). Distribution. North-east New South Wales. Map 26. Habitat. Only recorded with one collection as “Common on heavy grey alluvium under Eucalyptus camaldulensis”. This subspecies is distinctive in having small narrowly hastate leaves and in having a fruit with a smooth transluscent pericarp; the striate seed also clearly separ- ates it from the other subspecies. The subspecies leiocarpa is possibly more closely related to E. nutans, with which the seed ornamentation and herbaceous pericarp would have more in common, but it seems preferable to restrict the latter species to those plants with a succulent fruit. A collection from 16 mi [25.7 km] W of Garah (K. L. Soiling 431, NSW) differs from typical subsp. leiocarpa in having a more triangular leaf, a thin pericarp and an almost smooth seed. From the same locality has been collected a plant with leaf shape typical of subsp. leiocarpa but with tepals in fruit somewhat succulent and a pericarp thin but succulent (K. L. Soiling s.n. NSW 142982). It appears to be intermediate in morphology between E. trigonos subsp. leiocarpa and E. nutans. The infraspecific epithet is derived from two Greek words, leios, smooth, and carpos, fruit, referring to the smooth pericarp. 210 Nuytsia Vol. 4, No. 2 (1983) 4. Einadia allanii (Aellen) Paul G. Wilson, comb nov. —Chenopodium allanii Aellen, Candollea 8:7 (1939). Lectotype: Lammermoor Mts, New Zealand, Jan. 1929, H. H. Allan (CHR 1064) lecto. nov. Figure 5A. [Chenopodium triandrum auct. non G. Forster: J. D. Hook., FI. Nov.-Zel. 1:212 (1853) p.p.; J. D. Hook., Handb. N. Zeal. FI. 230 (1864) p.p.; ? T. F. Cheeseman, 111. N. Zeal. FI. 2:t.l64 (1914).] Spreading perennial woody towards base. Branches weak, slender, glabrescent. Leaves alternate; lamina thin, broadly elliptic to orbicular, 5-15 mm long, entire, sparsely mealy to glabrous; petiole slender. Inflorescence of axillary and terminal slender or branched panicles (congested when young). Flowers c. 0.5 mm diameter; tepals (4)5, ovate, not cucullate, united towards base, moderately mealy outside, en- larging in fruit but not enclosing nut; stamens 0-3, slightly exceeding tepals; fila- ments thickened at base, glabrous, disc absent; style absent, stigmas papillose c. O. 5 mm long. Fruit depressed spheroid c. 1.5 mm diam.; pericarp thin, papillose, loosely adherent. Seed depressed spheroid c. 1.3 mm diam; testa faintly punctuate, black. NEW ZEALAND: North Island:— Wellington, 19 Feb. 1950, G. B. Cone (CHR); Te Kaminara Bay, 21 Nov. 1947, V. D. Zotov (CHR). South Island: — Ben Lomond, 10 Dec. 1943, A. J. Healy(CHR). Distribution. North and South Islands of New Zealand. Habitat. ‘Coastal, lowland and montane sandy and rocky places throughout, but rather local inland’, fide Allan (1961). Until Paul Aellen’s paper in 1939 Einadia allanii had been confused with E. triandra (while plants of the latter were generally referred to Rhagodia nutans). The two species have superficial resemblances but they differ in leaf-shape and in fruit since the pericarp of E. allanii is thin and dry while that of E. triandra is succulent. Einadia allanii is evidently closely related to E. trigonos for they are similar in habit as well as in flower and fruit character. The two are separable on leaf-shape, that of E. allanii being broadly elliptic (with rounded apex) to orbicular while in E. trigonos it is bluntly hastate. The situation is complicated by the presence in New Zealand of the typical subspecies of E. trigonos, of its subspecies stellulata (both presumably introduced in the past 200 years), and of plants which are intermediate in character between the typical variants of E. trigonos and E. allanii. It seems likely that these ‘intermediate’ collections are of hybrid origin but this suggestion needs to be sub- stantiated by field observations since E. trigonos is extremely variable in Australia, as regards leaf form. This makes its recognition (and of putative hybrids) in New Zealand uncertain. Einadia allanii exhibits considerable variability in leaf shape and texture, and in inflorescence character. I have adopted a rather broad concept of the species since I am unable to find any clear characters by which the complex could be subdivided. One variant does stand out and would appear to warrant recognition as a distinct species, this is a plant found on Little Barrier Island that has an open paniculate in- florescence the branches of which are destitute of bracts. A plant very similar in form is also found on the Kermadec Islands. Until further work is done on possible rela- tives in other islands of the Pacific or in South America it would seem best not to de- scribe it as a distinct taxon. The following specimens from New Zealand represent this variant. Paul G. Wilson, Tribe Chenopodieae 211 LITTLE BARRIER IS.: T. F. Cheeseman (CHR 331646); R. Melville 6572 (CHR 140167). KERMADEC ISLANDS: Curtis Is., W. R. B. Oliver (CHR 289968); Raoul Is., VL. R. Sykes 1423/K (CHR 326878). The variant found on Little Barrier Island probably represents the plant illus- trated in Cheeseman, Illustrations of the New Zealand Flora vol. 2:t 164 (1914), under the name Chenopodium triandrum. The variation in morphology was com- mented on by Allan (1961 p. 228), however, it is not clear what taxa he had in mind since the names Chenopodium allanii and Rhagodia triandra have been applied rather imprecisely. 5. Einada hastata (R.Br.) A. J. Scott, Feddes Repert. 89:4 (1978). — Rhagodia hastata R.Br., Prod. 408(1810). Type: Port Jackson, near Duck River on the Parramatta Road, R. Brown (holo: BM;? iso: K, MEL, NSW). Figure 4F. Spreading perennial herb (rarely erect to 1.5 m), branching at base. Branches quadrangular, when young, with vitreous tubular and vesicular hairs (sometimes pale yellow) soon glabrous. Leaves opposite or subopposite, somewhat coriaceous and resinous when dry, glabrous or sparsely pubescent with vitreous vesicular hairs when young; lamina deltoid to broadly hastate, entire, 10-25 mm long, rounded and often apiculate at apex, lateral lobes short and obtuse, base cuneate to obtuse; petiole slen- der, somewhat shorter than lamina. Inflorescence a condensed or slender racemose panicle with scattered glomerules. Flowers sessile or shortly pedicellate, depressed, 0.5-1 mm diam.; tepals oblong-obovate, somewhat fleshy, thickened and cucullate at apex, remaining incurved over developing fruit, glabrescent, red and somewhat suc- culent in fruit; stamens 1-2, glabrous, filaments thickened at base, spreading at an- thesis; ovary depressed, glabrous, smooth; stigmas sessile, spreading c. 0.5 mm long. Fruit a berry, depressed, clasped by persistent red succulent tepals, c. 3 mm diam.; pericarp succulent red. Seed lenticular with rounded margin, 1.5 mm diam., radially rugulose. QUEENSLAND: Russell Is., C. T. White 3405 (BRI); Harrisons Pocket, S. T. Blake 3203 (BRI). NEW SOUTH WALES: Tallebung, G. M. Cunningham 645 (NSW); 8 km S of Inglewood, L. A. S. Johnson 7216 (NSW). VICTORIA: Mt Wheeler, A. E. Orchard 2662 (AD); Bendigo, A. C. Beauglehole 42128 (MEL). Distribution. Victoria, eastern Queensland and New South Wales. Map 25. Habitat. Shady gullies in rainforest, rocky hill slopes, and open plains. Einadia hastata may be recognised by its relatively thick, green, almost glabrous, broadly hastate leaves with rounded apex. The hairs on the young shoots have a vit- reous appearance unlike the milky colour of those in E. nutans and E. trigonos. Maiden in Agric. Gaz. New South Wales 8:123-125 (1897) states that the crushed leaves have an odour of perfectly fresh fish. It varies considerably in habit and in leaf size and texture but less so in leaf shape and least of all in seed characters. 212 Nuytsia Vol. 4, No. 2 (1983) 6. Einadia polygonoides (Murr) Paul G. Wilson, comb. nov.—Chenopodium triangulare R.Br. var polygonoides Murr, Bull. Herb. Boiss. 4:994 (1904). — C. triangulate subsp. polygonoides (Murr) Murr, Allg. Bot. Zeitschr. 16:56 (1910). — C. polygonoides (Murr) Aellen, Mitteil. Ges. Soloth. 8:213 (1928). Type citation: ‘Port Jackson district NSW, leg. Ernst Betke, 2. 1898, comm. Charles Walter (v. in herb. Turic.).’ Iso (?): Kogarah (Sydney), Feb. 1898, J. H. Camfield (NSW 143135, ? MEL 88743), see comment below. C. triangulare R.Br. var. angustifolium Benth., FI. Austral. 5:161 (1870). Type ci- tation: ‘To this belong the Queensland specimens [of C. triangulare] and some from New England.’ Lectotype: Armadilla, W. Barton 140 (MEL 80674) lecto. nov. Spreading weak grey-green herbaceous perennial with numerous stems arising from woody taproot. Branches slender, striate, very sparsely mealy when young with vesicular hairs. Leaves opposite or alternate; lamina chartaceous, narrowly oblong-el- liptic to narrowly hastate, 0.5-1. 5 cm long; apex acute; base obtuse to truncate; upper surface glabrous or glabrescent; lower surface mealy with white vesicular hairs when young; petiole slender c. one quarter length of lamina. Inflorescence a compact ter- minal racemose panicle of dense glomerules of flowers, elongating in fruit to 15 cm with disjunct glomerules; axis when young with short tubular and globular hairs and with small gland-tipped hairs. Flowers sessile, 0.5-1 mm diameter; tepals broadly ob- ovate with rounded apex; stamens 1(2), glabrous, filament thickened at base and spreading at anthesis; ovary depressed, papillose; style c. 0.1 mm; stigmas slender, di- varicate, c. 0.5 mm long. Fruit: tepals imbricate, closely enveloping nut, coriaceous, with circular limb and short narrow claw, glabrous somewhat resinous when fresh, eventually straw-coloured to black, midrib prominent, pericarp thin, dry, promi- nently papillose above. Seed lenticular with rounded margin, 1 mm diam.; testa finely radially rugulose. QUEENSLAND: Kindon Station, L. S. Smith 514 (BRI); Chinchilla, J. P. Ryan (BRI). NEW SOUTH WALES: Marayong, 3 July 1966, R. Coveny (NSW); Narellan, E. McBarron 13673 (NSW). SOUTH AUSTRALIA: Arno Bay, Jan. 1920, T. G. B. Osborn (AD); Clapham, 2 Mar. 1962, R. Pearce (ADW). Distribution. Central-east Queensland to central-east New South Wales; as an introduction, in Eyre Peninsula and Adelaide, South Australia. Map 28. Habitat. Appears to favour heavy loamy soils. Einadia polygonoides may be readily recognised by its narrow leaves, slender infructescence and free coriaceous tepals with circular limbs. The indumentum of this species is unusual for Einadia since some of the small multicellular hairs are ter- minated by a minute spherical cell which appears to be filled with resin. The type of Chenopodium triangulare var. polygonoides was stated by Murr (1904 l.c.) to have been collected by ‘Betke’ (a misspelling of ‘Betche’). However, a note on a probable isotype in herb. NSW states that the collector was J. H. Camfield. Murr (1910) cites the Camfield collection and states that Betche collected it at the same place and date. Whether there were one or two collections is not clear but it seems certain that they are of the same population. The type specimen probably represents a relatively recent introduction to the Port Jackson district. Paul G. Wilson, Tribe Chenopodieae 213 5. RHAGODIA R.Br. R.Br., Prod. 408 (1810); A. J. Scott, Feddes Repert. 89:5 (1978). Lectotype: R. billardieri R.Bt. (= R. baccata (Labill.) Moq.), see Ulbrich (1934 p. 480). Dioecious shrubs, ± mealy (at least when young) with minute globular or lobed vesicular hairs, these (in some species) collapsing and becoming scurfy, scaly, or fusing to form a continuous silvery sheen. Leaves opposite or alternate, simple, entire or bluntly lobed at base, 2-40 mm long. Flowers small, glomerulate, arranged in pan- icles or reduced to spike-like thyrses; bracts small and linear or absent; perianth ± globular c. 1.5 mm diameter, usually mealy outside and glabrous within; tepals 5, shortly united at base, often spreading and slightly enlarged in fruit, chartaceous or slightly fleshy. Male flowers: stamens, 5, filaments linear, usually united at base into a thickened saucer- or cup-shaped disc woolly or puberulous within (rarely glabrous), anthers exserted; pistillode minute, with or without a pair of minute erect stigmas. Female flower if terminal to glomerules or branches then with staminodes and a disc which is woolly within, if lateral then with small usually glabrous staminodes or these absent; ovary glabrous and globular; stigmas 2, sessile, divaricate, papillose. Fruit a berry, depressed globular, c. 3 mm diameter, yellow or red; pericarp fleshy usually with watery interior, viscous on drying. Seed horizontal, depressed globular or lenticular, 1-1.5 mm diameter; testa crustaceous, black, smooth to rugulose or re- ticulate; embryo annular; perisperm copious central. Eleven species, all endemic to Australia (including Tasmania). Key to species 1 . 2 . 3. 4. 5. 6 . Leaves small c. 3 mm long; intricately branched spinescent shrubs 2 Leaves more than 3 mm long or plant not spinescent 3 Stamens woolly at base; seed radially verruculose (W.A.) 4. R. acicularis Stamens glabrous; seed smooth to minutely granular (N.S.W., S.A., W.A.) 3. R. ulicina Perianth glabrous or almost so 4 Perianth mealy or with minute vesicular hairs 5 Leaves linear to narrowly or broadly obovate somewhat mealy below 5. R. preissii Leaves narrowly elliptic with minute spherical or saucer-shaped vesicular hairs (rarely glabrous) 6. R.baccata Leaves mealy when young, becoming scaly and covered with a silvery sheen; staminodes present in female flower 6 Leaves mealy or of irregular to spherical vesicular hairs or glabrous (not becom- ing scaly or forming a sheen) 7 Leaves papery or slightly fleshy, flat 1. R. candolleana Leaves fusiform or semiterete, fleshy 2. R. crassifolia 7. Indumentum of spherical (or saucer-shaped) vesicular hairs 8 Indumentum of irregularly shaped vesicular hairs 10 8. Leaves oblong to narrowly elliptic, c. 30 mm long, glabrous above, sparsely ves- icular pubescent below (coastal south-west Australia) 6. R. baccata Leaves elliptic to deltoid or circular (inland or eastern States) 9 22493 — ( 6 ) 214 Nuytsia Vol. 4, No. 2 (1983) 9. Leaves narrow elliptic, lamina mostly 7-10 mm long; female flowers lacking both disc and staminodes; seed almost smooth (W.A.) 8. R. drummondii Leaf-shape varied, lamina (3) 5-20 mm long; lateral female flowers with disc and often staminodes; seed prominently reticulate (central and eastern Australia). 9. R. spinescens 10. Leaves broadly obovate to orbicular or bluntly hastate, flat; inflorescence pyramidal 11 Leaves linear to elliptic or oblong-elliptic 12 11. Leaves obovate to orbicular, often conduplicate; indumentum of closely packed glossy, irregularly shaped vesicular hairs (W.A., Shark Bay to Geraldton) 7. R. latifolia Leaves broadly ovate to bluntly hastate, flat; indumentum of very minute ves- icular hairs, dull (S.A., N.T., eastern Australia) 11. R. parabolica 12. Leaves semiterete, somewhat fleshy; panicle narrow, often racemose (W.A., Geraldton area) 7. R. latifolia subsp. recta Leaves elliptic to ovate (inland or coastal south-west) 13 13. Leaves ovate or ovate-oblong, c. 15 mm long, thinly covered below with minute mealy hairs; flowers in dense slender spikes, sometimes paniculately arranged; staminodes usually absent in female flower (S.A., W.A., N.T., inland species) 10. R. eremaea Leaves elliptic, 10-20 mm long, densely covered below with minute vesicular hairs; flowers in dense panicles (W.A., coastal south-west). 6. R. baccata subsp. dioica 1. Rhagodia candolleana Moq., Chenop. Enum. 10 (1840). — R. baccata var. candolleana (Moq.) Moq. in DC., Prod. 13/2:50 (1849). Lectotype: Nouvelle hollande, Labillardiere (holo: P) lecto. nov. (See discussion below.) Figures 1, 2. Cbenopodium furfuraceum Moq. in DC., Prod. 13/2:64 (1849). Type: Terre de Diemen — detroit d’ Entrecasteaux, 1802, herb. Delessert (holo: P). [R. billardieri auct. non R.Br.: Benth., FI Austral. 5:152 (1870) p.p. excluding West- ern Australian specimens cited.] [R. baccata auct. pi. non (Labill.) Moq.: e.g. J. Black, FI. S. Austral, ed. 2, 287 (1948); Beadle et al., FI. Sydney Reg. 192 (1972); J. H. Willis, Handb. FI. Viet. 2:83 (1973).] [R. billardieri var. congesta auct. non (J. D. Hook.) Benth.: Benth., FI. Austral. 5:153 (1870).] [R. baccata var. congesta auct. non (J. D. Hook.) A. J. Scott: A. J. Scott, Feddes Repert. 89:7 (1978).] Weak spreading shrub or erect and up to 5 m high. Branches striate, mealy when young. Leaves opposite to alternate, shortly petiolate; lamina chartaceous to cori- aceous or somewhat fleshy, narrow to broad elliptic to broad ovate or hastate, 10- 25(40) mm long, acute to obtuse; upper surface sparsely mealy to glabrous; lower sur- face mealy developing a continuous grey to silvery sheen with age or glabrescent. In- florescence a terminal pyramidal panicle to 15 cm long. Flowers predominantly di- oecious; perianth fleshy, densely pubescent outside with weak irregularly shaped hairs, tepals ovate and united near base, midrib thickened. Male flowers: stamens 5, filaments linear-lanceolate, united at base into a thick saucer-shaped disc shortly puberulous within; pistillode from minute to normal size with small erect stigmas. Female flower: staminodes with ovate membranous filaments 1/4 to 1/2 length of Paul G. Wilson, Tribe Chenopodieae 215 tepals, sometimes terminating in minute antherodes, united at base into a thickened disc woolly within; ovary glabrous, style arms spreading, c. 0.5 mm long. Berry clasped at base by the fleshy red perianth. Seed depressed globose, 1.5-2.5 mm diam., testa almost smooth to faintly reticulate or rugulose. Key to subspecies Leaves elliptic to broadly ovate or hastate, lower surface grey or glabrescent la. subsp. candolleana Leaves hastate, acute, lower surface with a silvery sheen lb. subsp. argentea la. subsp. candolleana R. baccata var. parvifolia Moq. in DC., Prod. 13/2:50 (1849). Type: Port Jackson, Durville (holo: P). Chenopodium furfuraceam Moq. op. cit. Leaves narrow to broad elliptic, broad ovate, or hastate, obtuse; lower surface covered with a grey sheen or glabrescent. NEW SOUTH WALES: 2 mi [3.2 km] N of Bermagui, H. Salasoo 5107 (NSW); Kogarah, Nov. 1899, J. H. Camfield (NSW). VICTORIA: Lady Julia Percy Is., A. C. Beauglehole 49662 (MEL); Coringle, N. W. Wakefield 4274 (MEL). TASMANIA: Sulphur Creek, Jan. 1954, W. M. Curtis (HO); Coles Bay, 20 Apr. 1930, L. Rodway (HO). SOUTH AUSTRALIA: Henley Beach, 26 Mar. 1921, E. H. Ising (AD); Port Noarlunga, L. D. Williams 5134 (AD). WESTERN AUSTRALIA: Middle Is., Recherche Archipelago, A. S. Weston 8856 (PERTH); Observatory Is., A. S. Weston 9386 (PERTH). Distribution. Tasmania, southern New South Wales, Victoria, South Australia, Recherche Archipelago in Western Australia. Map 29. Habitat. Coastal situations. lb. subsp. argentea Paul G. Wilson, subsp. nov. Frutex omnis argenteo-squamosus. Folia aliquantum hastata 10-25 mm longa, acuta. Typus: South Australia; Maralinga Village, 15 July 1972, T. R. N. Lothian 5547 (holo: AD; iso: NCU n.v.). Shrub closely covered all over with a silvery scaly indumentum. Leaves somewhat hastate, 10-25 mm long, acute. SOUTH AUSTRALIA: Ifould Lake, T. R. N. Lothian 5429 (AD); Maralinga, 7 Feb. 1960, H. Turner (AD). 216 Nuytsia Vol. 4, No. 2 (1983) Distribution. South Australia, North-West Plains. The subspecies argentea has a distinctive leaf shape and a thick silvery sheen. It is clearly distinct in its distribution from the typical subspecies and could be separated (on morphological characters) at the species rank. Near Maralinga and at Ifould Lake subsp. argentea has an acutely hastate leaf; at Ooldea has been collected a plant (Spooner 2185, AD) which is similar but has an el- liptic leaf; also at Ooldea has been collected a plant (J. B. Cleland, 17 Aug. 1939, AD) with leaves similar to the Spooner specimen but with flowers and indumentum more like those of Chenopodium desertorum. Until extensive field investigations take place no satisfactory interpretation of this data is possible. On the Nullarbor Plain 12 km NW of Cocklebiddy Rhagodia crassifolia may be found growing with a plant intermediate in morphology between that species and R. candolleana subsp. argentea. This putative hybrid (P. G. Wilson, 11829, PERTH) suggests that R. candolleana subsp. argentea is also found in that district. Lectotypification of the name Rh^odia candolleana. The correct application of the name Rhagodia baccata, and the reasons for rejecting it for the species occurring in eastern Australia, are discussed under that name. The lectotypification of the alternative name, R. candolleana, also requires an explanation since it was based on mixed material. Moquin (1840) cited the type of R. candolleana as “In nova Hol- landia (v.s. in herb. Mus. Paris. DC., et Vindob.)”. In 1849, under R. baccata var. candolleana (Moq.) Moq., he wrote “v.s. in h. Labil. et Mus. Paris.” I have seen four sheets which appear to be syntypes of this species, their labels are as follows: A. Herb. P Label (1) Rhagodia candolleana Moq./Nouvelle hollande/Labiiiardfere/Herb. Poiret. (2) Rhagodia Candolleana Moq./Nova hollandia/herb. Vindob. (3) Herb. Mus. Paris/Herbier Moquin-Tandon/(Monographies du Prodrome) B. Herb. P Label (1) Rhagodia/Nova Holland/ Drummond No. 209. (2) Rhagodia hastata R.Br./(non)/Nova hollandia/herb. Vindob. (3) Herbarium Moquinianum/Rhagodia baccata Moq./var. Candolleana/ (R. candolleana Moq.) /nova Holland. (La Bill). (4) Herb. Mus. Paris/Herbier Moquin-Tandon/(Monographies du Prodrome) C. Herb. FI Rhagodia Candolleana Moq. [In Moquin’s handwriting] D. Herb. G-DC Label (1) Nouv Hollande cote d’orientale/Museum de Paris 1821 (2) baccata/Rhagodia Billardieri R.Br./var.CandoWeana/Moq. (3) Rhagodia/Nouv Hollande cote Merid./Mus de Paris 1821 Paul G. Wilson, Tribe Chenopodieae 217 holo IVPE KEW NEGATIVE NO, 171 90 ' mj frt'l »*'! o-ll ■’■Jt) 2U 34 4-U ‘L^ oflt n# ii'* 0^* ■y¥ 01 02 »0 I2» !3'0 01 14.0 Figure 1. Lectotype of Rhagodia candolleana Moq. in herb. P. 218 Nuytsia Vol. 4, No. 2 (1983) Sheets A and D contain only specimens of the eastern Australian plant here referred to Rhagodia candoUeana. Sheets B and C each contain a small vegetative specimen of R. candoUeana and a larger specimen of Tetragonia implexicoma (Miq.) J. D. Hook. (Aizoaceae). I have selected sheet A in herb. P. (Figure 1) as the lectotype since it bears Moquin’s handwritten determination and is of unmixed ma- terial. It is certainly one of the specimens studied by Moquin when describing the species. The sheet B in herb. P has, as noted above, a Drummond label no. 209. This is misplaced since James Drummond could not have collected either of the two speci- mens on the sheet; the Drummond no. 209 collection in herb. MEL is of Rhagodia baccata sensu stricto (i.e. the Western Australian plant) from which the Drummond label in herb. P had evidently strayed. Rhagodia candoUeana subsp. candoUeana varies considerably in leaf form, texture and sculpturing of the seed (faintly rugose in Tasmania to faintly reticulate in South Australia); it also varies in stature, being a shrub to 5 m high in Tasmania while rarely exceeding 1 m on the Australian mainland. However, there are insufficient supporting characters with which to establish further infraspecific taxa. In South Australia it is often not possible to clearly distinguish R. candoUeana from the coastal variant of R. crassifolia, however, since elsewhere these two species appear to be clearly separable I have continued to recognise them. Although predominantly coastal in its distribution subsp. candoUeana has been collected at Mt Gambier in South Australia. J. M. Black (1948) recorded it inland along the R. Murray but the relevant specimens are all of the inland variant of R. crassifolia. Black also recorded Rhagodia baccata var. linearis for South Australia, however, material so determined in his herbarium is either Suaeda australis (R.Br.) Moq. or R. crassifolia. Rhagodia candoUeana is nearly always dioecious but populations of apparently hermaphrodite plants do occur. Male plants sometimes have prominent pistillodes and these frequently enlarge to form berry-like structures which are probably galls. The peculiar indumentum of Rhagodia candoUeana (and of R. crassifolia) is made up of extremely thin-walled mealy hairs of irregular shape. These soon collapse and fuse together to form a metallic sheen similar to that found in some species of Atrip lex. The name Chenopodium congestum J. D. Hook, is based on a specimen of C. mu- rale L., q.v. However, as a variety of Rhagodia billardieri or of R. baccata it has been misapplied by Bentham (1870) and Scott (1978a) to variants of R. candoUeana in which the leaves tend to be rhombic or hastate and the inflorescence somewhat con- gested. The type of Chenopodium furfuraceum is a male specimen of Rhagodia candoUeana. It was not seen by Bentham (1870) who thought it might be conspecific with C. auricomum. 2. R. crassifolia R.Br., Prod. 408 (1810). Lectotype: Goose Island Bay, larger island, 15-16 Jan. 1802 R Brown, dist. no. 3038 (holo: BM) lecto. nov. Kparvifolia Moq. in DC., Prod. 13/2:52 (1849). Type: Ad Cygnorum fluvium, 1843, J. Drummond 715 (holo: P). Paul G. Wilson, Tribe Chenopodieae 219 Shrub to 1.5 m high, divaricately branched. Leaves mostly alternate, fleshy, linear to narrow-elliptic in outline, fusiform or semiterete, often conduplicate or channelled above, falcate when thick, 1-2(4) cm long, mealy when young, becoming glabrescent above, hairs collapsing and fusing to form a metallic sheen. Inflorescence pyramidal to 5 cm long or reduced to a short spike. Flowers and fruit similar to that in R. candolleana. Seed depressed globose, 1.5 mm diam., very faintly reticulate or rugulose. VICTORIA: 16 mi [25.7 km] NNW of Underbool, A. C. Beauglehole 40470 (MEL); 5 mi [8 km] E of Murrayville, 29 Aug. 1955, J. H. Willis (MEL). SOUTH AUSTRALIA: Nundroo, A. G. Spooner 2194 (AD); Port Lincoln, 17 Dec. 1941, J. B. Cleland (AD). WESTERN AUSTRALIA: Corbett Is., Recherche Archipelago, R. D. Royce 6189 (PERTH); 12 km E. of Mundrabilla, P. G. Wilson 11832 (PERTH). Distribution. North western Victoria westwards to near Albany on the south coast of Western Australia. Map 30. Habitat. Principally a coastal species but also inland where usually on calcareous soil; often in a mallee community. Rhagodia crassifolia intergrades with R. candolleana in coastal regions of South Australia and many collections cannot be confidently placed in either species. Near Cocklebiddy in the southern Nullarbor Plain of Western Australia R. crassifolia has been found growing with an apparent hybrid (P. G. Wilson 11834, PERTH) between that species and R. candolleana subsp. argentea. The coastal variant of R. crassifolia differs somewhat from that found in the Murray-Mallee region of South Australia and adjacent areas of Victoria; this inland variant has a strongly divaricate habit and longer more slender leaves. It was princi- pally this variant that J. M. Black (1948) incorrectly referred to under the name R. baccata var. linearis. Moquin (1849 loc. cit.) based his concept of R. crassifolia largely on a specimen collected by Alan Cunningham on Dirk Hartog Island. This collection is, in fact, of a distinct species, R. latifolia (Benth.) Paul G. Wilson. The specimen of R. crassifolia collected by James Drummond in Western Australia, being considered by Moquin to be specifically distinct from that of Cunningham, was described as a separate species, R. parvifolia. Although Moquin cited its locality as ‘Ad Cygnorum fluvium’, i.e. from Swan River Western Australia, the specimen was probably collected near the Fitzgerald River mouth since it closely matches other specimens from that area while the species is not found on the west coast of Western Australia. Bentham (1870) and Ewart and Davies (1917) recorded R. crassifolia from North- ern Australia, in both cases citing a specimen from Sturt Creek collected by F. Mueller. This specimen is probably a syntype of Chenopodium nitrariaceum (F. Muell.) Benth. but is not conspecific with the lectotype of that species; it belongs in fact to R. eremaea. 3. Rhagodia ulicina (Gand.) Paul G. Wilson, Nuytsia 4:53 (1982.)— Chenopodium ulicinum Gand., Bull. Soc. Bot. France 66:224(1919). Type: Mt Lyndhurst, April 1900, M. Koch 366 (holo: LY; iso : PERTH). Figure 5F. 220 Nuytsia Vol. 4, No. 2 (1983) Dense intricately branched shrub 0.5-1 m high, dioecious. Branches divaricate, spinescent, mealy when young with close dense covering of small branched vesicular hairs. Leaves alternate, often clustered, fleshy, elliptic, 2-5 mm long, closely vesicular mealy, narrowed at base into an indefinite petiole. Inflorescence a few-flowered nar- row condensed cyme (sometimes spinescent) with 1-4 lateral and a large terminal flower, or reduced to a single flower. Perianth depressed globose 1.5 mm diam., mealy outside with small branched vesicular hairs; tepals broadly ovate, united at base, eventually spreading beneath fruit. Male flower: stamens 5, filaments slender, glabrous, disc inconspicuous, anthers shortly exserted c. 0.5 mm long; pistillode min- ute. Female flower: staminodes represented by 5 scarcely observable antherodes c. 0.5 mm long; ovary glabrous; style absent to short and c. 0.5 mm long, glabrous or sparsely hirtellous; stigmas spreading, slender subulate, pilosulose. Fruit baccate, at first enveloped by perianth; pericarp succulent, appearing inflated on drying, free from seed. Seed lenticular with rounded margin, c. 1.5 mm diam., testa smooth to minutely granular, black. NEW SOUTH WALES: Fowlers Gap, S. Jacobs 2268 (NSW); 20 mi [32.2 km] N of Balranald, W. E. Mulham W518 (NSW). VICTORIA: Yatpool area, J. H. Browne 117 (PERTH). SOUTH AUSTRALIA: Mt Grainger, R. J. Chinnock 1479 (AD); Wilkawillina Gorge, T. R. N. Lothian 5126 (AD). WESTERN AUSTRALIA: 19 km NNW of Mt Glass, K. Newbey 5609 (PERTH); 12 km NW of Cocklebiddy, P. G. Wilson 11827 (PERTH). Distribution. Eastern New South Wales and north-eastern Victoria to Northern Eyre Peninsula in South Australia westwards to the south-east of Western Australia around the Bremer Range. Map 31. Habitat. Generally rocky situations often over limestone, but has been recorded from clay flats. This species has been confused in herbaria with both R. spinescens and Cheno- podium nitrariaceum. It was included by Bentham (1870) under the former name (i.e. the specimens cited as ‘Murray river, F. Mueller’, ‘Gawler Ranges, Sullivan’, and ‘Barrier Range, Victorian Expedition’), however, his description of R. spinescens was based not only on that species and R. ulicina but also on Chenopodium desertorum. Gandoger (1919 loc. cit.) recognised R. ulicina as being a distinct species but his de- scription was very brief and suggested that he was unaware that the plant was di- oecious, shrubby, and had baccate fruits, for these characters establish the species as a member of the genus Rhagodia and not Chenopodium. Scott (1978b) included R. ulicina (as Chenopodium) with C. nitrariaceum under Chenopodium sect. Rhagodioides Benth.; however, Scott’s sectional description is inapplicable to R. ulicina and the two species are not closely related. Rhagodia ulicina may be distinguished from R. spinescens and C. nitrariaceum by its intricately branched habit and usually by its small fleshy leaves. Some specimens of C. nitrariaceum resemble R. ulicina in leaf shape but differ in indumentum: that of C. nitrariaceum is scurfy and forms a silvery sheen with age, while the vesicular hairs of R. ulicina are irregularly lobed and remain distinct. Rhagodia ulicina is most similar to R. acicularis (an endemic of the Wongan Hills, Western Australia); the differences between them are give under the latter species. Paul G. Wilson, Tribe Chenopodieae 221 4. Rhagodia acicularis Paul G. Wilson, Nuytsia 4:51 (1982). Type: Monk’s Well Gully, Wongan Hills, 15 Feb. 1980, P. G. Wilson 11713 (holo: PERTH; iso: CANB). Figure 5E. Compact intricately branched rounded shrub c. 0.5 m high, closely covered with minute vesicular hairs which retain their shape with age, dioecious. Branchlets spinescent. Leaves alternate, small, somewhat fleshy, narrowly elliptic, 2-5 mm long, glabrescent above, closely vesicular puberulous below. Inflorescence spicate (or shortly paniculate) with a spinescent axis c. 10 mm long; flowers few, sessile, sub- tended by a minute bract. Male flowers broadly turbinate, c. 1 mm long; tepals 5, divided to base, narrowly obovate, incurved, closely vesicular hairy outside, glabrous within; stamens 5; filaments slender-terete, lanate towards base, united into a thick cup-shaped disc; anthers shortly exserted, c. 0.5 mm long; pistillode minute. Female flowers spheroidal, c. 1.2 mm diam.; tepals narrowly obovate not enlarging in fruit; staminodes absent; ovary glabrous; style absent; stigmas slender, papillose, c. 0.3 mm long. Terminal flower on female inflorescence sterile, densely filled with woolly hairs arising from staminodes; pistillode minute with 2-3 erect glabrous stigmas. Fruit bac- cate, depressed spherical, clasped around margin by tepals but exposed at apex and towards base in inter-tepalar spaces; pericarp succulent, adherent to seed. Seed len- ticular with rounded margin, 1.5 mm diam., radially verruculose. WESTERN AUSTRALIA: Wongan Hills, K. F. Kenneally 2384 and 6448 (PERTH); ibid., P. G. Wilson 11712 (PERTH). Distribution. Wongan Hills, south-western Western Australia. Habitat. Red soil on gravelly lateritic slope in eucalypt woodland. Rhagodia acicularis closely resembles R. ulicina but differs in the following charac- ters: 1) Terminal flower of female inflorescence sterile (not large and fertile), 2) Seed radially verruculose (not smooth or minutely granular), 3) Male flower with woolly stamens united into a cup-shaped disc (not glabrous and with an inconspicuous disc). It is a typical member of the genus and evidently closely related to R. spinescens R. Br. and R. drummondii Moq. 5. Rhagodia preissii Moq. in DC., Prod. 13/2:49 (1849).— Chenopodium preissii (Moq.) Diels, Bot. Jahrb. 35:181(1904). Type: Near York, Western Australia, Preiss 1252 (iso: K, MEL). R. obovata Moq., Chen. Mon. Enum. 10(1840). Type: Nouv. Holland, cote meridi- onale, 1821 (holo: P). [R. linifolia auct. non R.Br.: Nees in Lehm., PI. Preiss. 1:633(1845).] Shrub to 2 m high. Branches slender, when young costate and sparsely mealy. Leaves chartaceous (slightly fleshy when fresh), to coriaceous (subsp. obovata), lin- ear to broadly obovate or spathulate, 3-4 cm long, gradually tapering at the base into a slender petiole, apex rounded to acute, lower surface sparsely and minutely mealy when young. Inflorescence a pyramidal panicle c. 5 cm long, branches slender, flowers in disjunct glomerules. Flowers spherical, c. 1 mm diam; tepals almost orbicu- lar, glabrous and eciliate (or sparsely mealy in subsp. obovata) enlarging slightly in fruit. Male flowers: filaments united into a saucer-shaped disc, somewhat pilose at base; pistillode minute (c. 0.5 mm high) with a pair of short erect subulate stigmas. Female flowers: terminal flowers usually sterile with prominent linear staminodes 222 Nuytsia Vol. 4, No. 2 (1983) densely pilose at base; pistillode ovoid, 0.3-1 mm high: non-terminal flowers fertile; staminodes absent; ovary with a pair of subulate stigmas. Berry depressed spherical, c. 3 mm diam. Seed c. 1.5 diam.; testa glossy with faint radial ribbing. Key to subspecies Leaves linear to narrow obovate, usually chartaceous 5a. subsp. preissii Leaves broadly obovate, coriaceous 5b. subsp. obovata 5a. subsp.preissii Leaves linear to narrow obovate or spatbulate chartaceous (to coriaceous), flat or cymbiform. Perianth glabrous. SOUTH AUSTRALIA: Barton, N.T. Burbidge 1866 (CANB); Dublin, J. C. Noble 84 (AD). WESTERN AUSTRALIA: Merredin, M. Koch 2882 (NSW); 32 km NNE of Stokes Inlet, A. E. Orchard 1645 (CANB). Distribution. West coastal and south-west Western Australia, southern South Aus- tralia. Map 32. Habitat. Often in deep sand. 5b. subsp. obovata (Moq.) Paul G. Wilson, comb, et stat. nov. — R. obovata Moq., Chen. Mon. Enum. 10 (1840). Type: Nouv. holland. cote meridionale, 1821 (holo: P). Leaves broadly obovate, gradually to abruptly narrowed at base into a slender petiole c. 1/3 length of leaf, in all 1.5-3 cm long, coriaceous, glabrescent to thinly mealy, apex rounded to truncate, apiculate. Perianth glabrous to sparsely mealy. WESTERN AUSTRALIA: Dorre Is., A.S. Weston 10496 (PERTH); Kalbarri to Red Bluff, R.J. Hnatiuk 760494 (PERTH). Distribution. Shark Bay-Geraldton region of Western Australia. Map 32. Habitat. Frequently growing over limestone. The application of the names Rhagodia dioica and R. obovata is difficult to deter- mine due to intergradation of the typical variants of these species with R. baccata and R. preissii. The type of R. obovata comes from Dirk Hartog Island where the plant has leathery, broadly obovate leaves and glabrous flowers. On the mainland around Shark Bay the leaves are somewhat narrower and the flowers slightly mealy, while further south the mealiness increases and the leaf shape varies from obovate (near Geraldton) to elliptic (near Perth). Near Perth the flowers and leaves are densely mealy, and here the plant intergrades with R. baccata; this variant of intermediate aspect has been described as the species R. dioica. In addition to this apparent intergradation, R. obovata also passes, to the north and east of Shark Bay, into typical R. preissii with thin narrow leaves and glabrous flowers. I have, in this paper, recognised R. baccata as a distinct species because of its normally clearly dis- tinguishable morphological characters while R. obovata I am including as a Paul G. Wilson, Tribe Chenopodieae 223 subspecies of R. preissii whose typical subspecies is also clear cut in its characters. Since the syntypes of R. dioica come near to the typical variant of R. baccata I am treating them as conspecific. 6. Rhagodia baccata (Labill.) Moq. in DC., Prod. 13/2:50 (1849). — Chenopodium baccatum Labill., Nov. Holl. PI. Sp. 1: 71, t. 96 (1805). — R. billardieri R. Br., Prod. 408 (1810), nom. illeg. Type “Habitat in capite Van-Diemen.” Lectotype: FI (see notes below) lecto. nov. Figures 2, 3. Rhagodia dioica Nees in Lehm., PI. Preiss. 1:636 (1845). Lectotype: Fremantle, Preiss 1253 (holo: K; iso: MEL) lecto. nov. Rhagodia radiata Nees in Lehm., PL Preiss. 1:637 (1845). — R. billardieri var. linearis Benth., FI. Austral. 1:153 (1870). — R. baccata var. linearis (Benth.) J. Black, FI. S. Austral. 683 (1929); Ewart, FI. Viet. 449 (1931). Type citation: ‘In elevatis arenosis juxta urhiculam Freemantle, Decembri a 1838. Herb. Preiss. No. 1250’. Isotype: MEL 80680, 80681, 80682. Rhagodia cygnorum Gandoger, Bull. Soc. Bot. France 66:224 (1919). Type: Fremantle, 15 March 1899, A. Morrison (holo: LY). Spreading shrub to 2 m high. Branches slender, costate, mealy when young. Leaves opposite or alternate shortly petiolate; lamina chartaceous (somewhat fleshy in subsp. dioica), narrowly oblong-elliptic to elliptic, 1-4 cm long, 3-8 mm wide, apex rounded to obtuse and minutely apiculate, margin often recurved, upper surface smooth and glabrous or sparsely mealy, lower surface with prominent midvein, sparsely to densely mealy with saucer-shaped collapsed vesicular hairs, rarely glabrous (minutely mealy in subsp. dioica). Inflorescence an open terminal panicle; flowers clustered in sessile lateral dichasial glomerules. Flowers predominantly di- oecious, globular, 1.5 mm diam.; perianth densely covered with irregularly shaped vesicular hairs (rarely glabrous); lobes broadly ovate to orbicular, fleshy in centre. Female plant: terminal flower of main and lateral branches of panicle with 5 mem- branous lanceolate staminodes 1 mm long united into a fleshy saucer-shaped disc densely lanate within, pistil (or pistillode) glabrous with a pair of short erect styles; lateral flowers (which predominate) glabrous within, lacking staminodes and disc, ovary globular, glabrous with a pair of short spreading subulate styles. Male plant: staminal filaments narrowly lanceolate to linear, glabrous; disc saucer-shaped, fleshy; pistillode ellipsoidal, minute (c. 0.5 mm high) with a pair of short stylar processes. Fruiting perianth spreading to retlexed with somewhat hardened midrib, pale fawn in colour, persistent; berry depressed globular, red, pericarp fleshy. Seed lenticular, c. 1.8 mm diam.; testa glossy, minutely and irregularly tuberculate. Key to subspecies Leaves chartaceous with recurved margin (on drying), mealy beneath with discrete saucer-shaped hairs 6a. subsp. baccata Leaves fleshy, margin not recurved, mealy beneath with very small hairs 6b. subsp. dioica 6a. subsp. baccata Rhagodia radiata Nees in Lehm., op. cit. (1845). 224 Nuytsia Vol. 4, No. 2 (1983) Rhagodia baccata var. angustifolia Moq. in DC., Prod. 13/2:50 (1849). Type citation: ‘v.s. in h. Poir. Mus. Vindob. et DC., comm, a cl. Labill. Cunningh. et F. Bauer.’ Lectotype: Nova Hollandia. Ex herb. Labill. (P; iso: FI) lecto. nov. (see notes below). R. cygnorum Gandoger, op. cit. (1919). Leaves narrowly oblong-elliptic to oblong-elliptic, 2-4 cm long, 4-8 mm wide, char- taceous, margin recurved, lower surface sparsely to densely mealy with saucer- shaped collapsed vesicular hairs, rarely glabrous. WESTERN AUSTRALIA: Nornalup Inlet, 15 Jan. 1947, M. B. Johnson (PERTH); Recherche Archipelago, R. D. Royce 6280 (PERTH); 31 mi [49.9 km] S of Perth, R. A. Saffrey 877 (PERTH). Distribution. Western Australia from Guilderton (80 km N of Perth) south and east- wards around the coast to Cape Arid. Map 31. Habitat. Coastal or near coastal situations. This subspecies (under various names) has frequently been considered to be conspecific with Rhagodia candolleana. It may be readily distinguished by its thin, narrow leaves with recurved margins and by the absence of staminodes in the female flowers (when lateral in position). A further distinction lies in the indumentum of the leaves which consists of collapsed vesicular hairs circular in outline. In R. candolleana the hairs are irregular in shape, very thin walled, and when collapsed fuse to form a silvery sheen. The species are both found in the Recherche Archipel- ago where they retain their distinctive characters; their distribution is otherwise quite distinct. Almost glabrous plants of R. baccata subsp. baccata are found at some localities but this variant grades into the typical one with a mealy indumentum and there is no reason to segregate it as a distinct taxon. Lectotypification of Chenopodium baccatum. Labillardiere (1805 loc. cit.) gave as the provenance of Chenopodium baccatum ‘Habitat in capite Van-Diemen’, i.e. from the south coast of Tasmania. Material collected by Labillardiere in herb. FI, and so determined by him, is mounted on four sheets; these contain some specimens which correspond to the Tasmanian plant and others which could only have been collected on the south-west coast of Western Australia. The plants from the two areas I con- sider to be specifically distinct (they have been generally considered by earlier workers to be distinct at either the level of species or variety); therefore it is import- ant that C. baccatum be lectotypified. Three of the four sheets mentioned above were re-identified by Moquin, presum- ably prior to 1849 when he cited the specimens in his treatment of the Chenopodiaceae in De Candolle’s Prodromus vol. 13/2. The sheet containing the largest number of specimens was determined by Moquin as R. baccata, a second sheet as R. baccata var. candolleana, and a third as R. baccata var. angustifolia. Moquin had therefore selected one of the four sheets as being representative of the typical variety. Labillardiere refers to the leaves as being ‘lanceolatis’ and the inflorescence as ‘racemis compositis’; the fruit was also described. The specimens which match the Western Australian species have narrowly elliptic leaves and bear large inflor- escences; they correspond well with Labillardi§re’s description and are a good match Paul G. Wilson, Tribe Chenopodieae with his illustration (tab. 96). The specimens which match the Tasmanian species have short elliptic to obovate leaves and are either vegetative or bear short racernes of young flowers; they do not agree with either Labillardiere’s illustration or descrip- tion. The lectotype of Chenopodium baccatum is, therefore, to be chosen from among those specimens collected by Labillardiere in Western Australia and preferably from those selected by Moquin as being typical of the species. The sheet determined as Rbagodia baccaia by Moquin contains Labillardiere’s manuscript description, four pieces of the Tasmanian plant, none of which corresponds to the illustration, and two pieces of the Western Australian, one of which is in flower and the other in fruit (Figure 2). As this fruiting specimen is an excellent match with tab. 96 (it was prob- ably the one illustrated) and with the description, I am selecting it as the lectotype of Chenopodium baccatum. The Tasmanian material I am referring to Rbagodia candolleana Moq. (q.v.) a species which also requires lectotypification. I have seeri a photograph of a specimen collected by Labillardiere in herb. PH named by him Chenopodium baccatum. This is of the Western Australian plant and therefore of Rbagodia baccata sensu lectotypica. Lectotypification of Rbagodia baccata var. angustifolia. In the Decandolle Her- barium at Geneva (G— DC) there are two specimens which appear to be syntypes of this variety. They are as follows: (1) Lewis-Land, Feid. Bauer (2) King George Sound, A. Cunningham In the Paris Herbarium (P) there is one specimen ‘Ex herb. LabilL’ which came from Moquin’s herbarium and which was seen by him during the writing of the Chenopodiaceae for Decandolle’s Prodromus vol. 13/2. In the Florence Herbarium (FI) there is a specimen collected by Labillardiere which has been determined by Moquin as Rbagodia baccata var. angustifolia. Each of the four collections cited above is of the variant of R. baccata found on the south coast of Western Australia between the Recherche Archipelago and Albany. As lectotype of var. angustifolia I have selected the Labillardiere specimen in herb. P, (Figure 3) of which the specimen in herb FI is probably a duplicate. 6b. subsp. dioica (Nees) Paul G. Wilson, comb, et stat. nov.—R. dioica Nees in Lehm., PI. Preiss. 1:636 (1845). Lectotype: see above. Leaves elliptic, 10 x 3 - 20 x 7 mm, obtuse, fleshy, coriaceous on drying, upper sur- face glabrescent, lower surface densely white mealy with very small hairs. WESTERN AUSTRALIA: Garden Island, May 1946, G. R. Meadly (PERTH); Fremantle, P. G. Wilson 8693 (PERTH). Distribution. Western Australia, Geographe Bay northwards to Fremantle. Map 30. Habitat. Coastal. This subspecies is sympatric over part of its range with the typical subspecies and plants intermediate in form between the two taxa may be found. It has thicker leaves 226 Nuytsia Vol. 4, No. 2 (1983) Figure 2. Lectotype of Rhagodia baccata (Labill.) Moq. Mixed collection made by Labillardiere of R. baccata and R. candolleana. The left-hand, the upper central, and the lower specimen in the right- hand cluster are of R. candolleana. The lower central specimen and the upper specimen of the right- hand cluster are of R. baccata. The lectotype of R. baccata is indicated by an arrow. Photo of sheet in herb. FI. Paul G. Wilson, Tribe Chenopodieae 227 H0L01 YPh. K£W NEGATIVE NO, 1TJ93 ■ - 7 i ■ i GP' o.i .ja^§!8 Oi ya ii) %i> ^ 70 , ! o m ' ■ ' j,- . 95 OV . (iiS Q'i SiO 90 SGO tiD i/f) n-0 -<■ HKRIi. MUS. PARIS. llEBilAiSiry WloHM.TNi | H»rbi« MOyl-iN-TANm>S Figure 3. Lectotype of Rhagodia baccata var. angustifolia Moq. in herb. P. 228 Nuytsia Vol. 4, No. 2 (1983) than subsp. bsccata and a more dense indumentum which is made up of smaller and less well-defined bladder hairs. The type is of a variant of subsp. dioica rather close morphologically to the typical subspecies; that is the leaves are thinner and lep furfuraceous. At the other extreme are found plants with very fleshy leaves similar in shape and size to those found in R. crassifolia. 7. Rhagodia latifolia (Benth.) Paul G. Wilson, comb, et stat. nov.— R. crassifolia var. latifolia Benth., FI. Austral. 5:155 (1870). Type: Dirk Hartog Is., Jan 1822, A. Cunningham 321 (holo: K; iso: MEL). Shrub to 1.5 m high, dioecious. Branches firm, densely covered when young with compact spherical or irregularly shaped vesicular hairs. Leaves opposite or alternate, densely covered all over with compact vesicular hairs; lamina coriaceous to fleshy, linear to broad elliptic or orbicular, c. 1 cm long; petiole slender c. 5 mm long. Inflor- escence terminal, pyramidal or slender, glomerules scattered along its branches. Flowers depressed globular c. 1.5 mm diam; tepals obovate, cucullate, densely vesicu- lar outside, not enlarging in fruit. Male flower: filaments united into a cup-shaped disc woolly within; pistillode minute c. 0.2 mm long without stigmas. Female flower when terminal in position with short membranous staminodes and densely woolly disc, when lateral in position without staminodes and glabrous within; stigmas divaricate. Berry depressed globular c. 3.5 mm diam., exceeding the spreading peri- anth which is pale red within. Seed lenticular, 1.3 mm diam., with faint radial rugulosity. Key to subspecies Leaves elliptic to orbicular; inflorescence pyramidal 7a. subsp. latifolia Leaves semiterete; inflorescence slender 7b. subsp. recta 7a. subsp. latifolia Divaricately branched shrub. Leaves coriaceous, elliptic to orbicular, often con- duplicate and then lunate to semicircular, densely covered all over with compact spherical vesicular hairs. Inflorescence pyramidal. WESTERN AUSTRALIA: Dirk Hartog Is., A. S. George 11580 (PERTH); Dorre Is., A. S. Weston 10482 (PERTH). Distribution. Shark Bay south to Murchison River. Map 29. Habitat. Sand dunes, frequently associated with limestone. 7b. subsp. recta Paul G. Wilson, subsp. nov. Rami ascendentes, graciles. Folia crassiuscula, semiteretia, c. 10 mm longa, 1 mm lata, supra plana glabra, infra dense farinosa cum pilis minutis vesicularibus, irregulariter formatis. Inflorescentia in paniculam gracilem spiciformen disposita. Typus: Victoria District, Geraldton, Jan 1901, Diels et Pritzel 212 (holo: PERTH). Branches ascending, slender. Leaves fleshy, semiterete, c. 10 mm long, 1 mm wide, ± flat and glabrous above, densely mealy below with very small irregularly shaped vesicular hairs. Inflorescence a slender, spike-like panicle. Paul G. Wilson, Tribe Chenopodieae 229 WESTERN AUSTRALIA: Mt Hill, Oldfield (MEL 87904); Geraldton, C. H. Ostenfeld343 (PERTH). Distribution. Western Australia; near Geraldton north to Murchison River. Map 29. Habitat. Sand dunes. The typical variant of Rhagodia latifolia is very distinct in appearance since the orbicular leaves become conduplicate (at least on drying) and are then. lunate in shape. It evidently grades to the south into subsp. recta (in which the leaves are semiterete), since plants intermediate in form have been collected. Rhagodia latifolia is quite distinct from R. crassifolia, under which name it was placed by Bentham (1870), for in the latter species the hairs of the leaves soon col- lapse and coalesce to form a silvery sheen while the female flowers possess prominent staminodes. Material of subsp. recta was considered by Bentham to represent typical R. crassifolia, a view evidently based solely on the leaf shape. The infraspecific epithet is taken from the Latin word rectus, straight, referring to the straight branches of this plant. 8. Rhagodia drummondii Moq. in DC., Prod. 13/2:52 (1849). Type: Drummond 133 (P). Intricately branched, loose, dioecious shrub up to 1.5 m high. Leaves opposite or alternate, somewhat coriaceous (slightly fleshy when fresh), narrowly elliptic to ellip- tic, shortly petiolate, 7-10(15) mm long, 2-3(4) mm wide, apex acute, base cuneate, upper surface glabrescent, lower surface densely covered with pale grey silvery spherical to saucer-shaped vesicular hairs. Influorescence a short terminal panicle to 2.5 cm long, frequently reduced to a short spike-like thyrse. Flowers sessile, de- pressed globular to broadly turbinate, solitary or in small glomerulate dischasia; tepals broadly ovate, shortly united at base, bluntly keeled towards apex, densely covered with vesicular hairs. Male flowers: staminal filaments united into a fleshy disc which is woolly within, pistillode minute without trace of stigmas. Female flowers: terminal flowers with minute staminodes and disc, slightly woolly within; lateral Rowers glabrous within and without staminodes or disc, stigmas slender. Fruiting perianth at first clasping fruit hut eventually spreading, c. 4 mm diameter, pale fawn or pale red within; berry depressed globular; seed lenticular, c. 1.8 mm diam., faintly radially rugulose, glossy. SOUTH AUSTRALIA: 12 km E of the Serpentine Lakes, 28°30'S, 129°04'E, D. E. Symon 12540 (ADW); 9 km N of Connie Sue Crossing on S.A.-W.A. border, 28°25'S, 129°00'E, D. E. Symon 12621 (ADW). WESTERN AUSTRALIA: Dowerin, A. M. Ashby 2849 (AD); 14.7 mi [23.7 km] N. of Winchester, M. D. Tindale 1289 (NSW). Distribution. Inland southern Western Australia eastwards to the extreme west of South Australia. Map 33. Habitat. Somewhat saline inland areas, also in Eucalyptus salmonophloia (Salmon Gum) Woodland. 22493 — ( 7 ) 230 Nuytsia Vol. 4, No. 2 (1983) Rhagodia drummondii is similar to R. spinescens but differs most obviously in the leaf shape (always elliptic) and in the absence of staminodes in the lateral female flowers. It was considered by Bentham (1870) and by Scott (1978b) to be a form of R. crassifolia, however, the two are not closely related; they differ very greatly in in- dumentum (the hairs of R. crassifolia collapsing to form a continuous metallic sheen) and in flower morphology, particularly in the presence of prominent staminodes in the female flowers of R. crassifolia. 9. Rhagodia spinescens R.Br., Prod. 408 (1810). Type: Inlet XIV [St. Vincent Gulf], R. Brown (holo: BM; iso: K). Figure 5D. R. spinescens var. deltophylla F. Muell., Rep. PI. Babbage’s Exped. 19 (1859).— R. deltophylla (F. Muell.) A. J. Scott, Feddes Repert. 89:10 (1978). Type citation: Lake Campbell. Lectotype: N.E. of Lake Gairdner, Hersch [Herschel] Babbage.’ (MEL 87960) lecto. nov. (see note below). Intricately branched shrub to 1-5 m high. Branches divaricate, frequently spinescent, densely mealy when young. Leaves opposite or subopposite, chartaceous to coriaceous, mealy all over with discrete spherical or saucer-shaped hairs, lamina ovate to deltoid or hastiform 0.5-2 cm long, apex obtuse to rounded, petiole slender half length of lamina. Inflorescence a rigid, straight, terminal panicle sometimes reduced to a single axis. Flowers sessile, densely mealy outside, perianth 5-partite. Male flowers: perianth globular, 0.5-1 mm high, filaments united at base into a thick broad saucer-shaped disc, glabrous; pistillode hemispherical to obturbinate c. 0.5 cm high with a pair of minute erect subulate stigmas (rarely the pistillode is fertile). Fe- male flowers: perianth at first soft and globular to broadly turbinate c. 1 mm high, eventually spreading under the fruit (never enveloping it) and 2.5-3 mm diameter, becoming thick and hard and often pale red; staminodes lanceolate, c. 0.3 mm long, with minute antherodes, united at base into a broad glabrous or sparsely hairy disc; ovary with a pair of spreading stigmas. Berry depressed globular, pericarp fleshy, pink or red. Seed lenticular c. 1 mm diameter; testa black, prominently reticulate with a radially elongated matrix. QUEENSLAND: Noondoo, S. T. Blake 10543 (BRI); 20 mi [32.2 km] W of Tara, T. J. McDonald 71 (BRI). NEW SOUTH WALES: Woorandara Stn., E. M. Canning 3808 (CBG); 16 mi [25.7 km] W of Garah, K. L. Soiling 332 (NSW). VICTORIA: 5 mi [8 km] NNW of Kerang, H. J. Aston 559 (MEL); 10 mi [16.1km] W of Cohuna, J. Vickery 2021 (NSW). SOUTH AUSTRALIA: 6 km W of Hawker, B. Copley 215 (AD): Tallaringa Well, T. R. N. Lothian 2689 (AD). WESTERN AUSTRALIA: 3 mi [4.8 km] S of Reid, T. E. H. Aplin 1677 (PERTH); Nunna, A. W. H. No. E 17 (PERTH). NORTHERN TERRITORY: Mt Brassey, A. C. Beauglehole 44701 (NT); Hale River Plot, T. S. Henshall 930 (NT). Distribution. Central and eastern Australia. Map 34. Habitat. Extremely variable due to the numerous ecotypes involved. Paul G. Wilson, Tribe Chenopodieae 231 Lectotypification of the name Rhagodia spinescens var. deltophylla. This name has been applied in herbaria by F. Mueller and by later botanists to broad leaved vari- ants of several species, principally R. spinescens, Chenopodium desertorum and C. curvispicatum. When searching for a suitable lectotype a number of specimens were found in herbaria K and MEL which had been collected prior to the publication of R. spinescens var. deltophylla and which Mueller had so annotated (or as R. deltophylla), and had evidently referred to when drawing up the protologue; how- ever, none were labelled as having been collected at Lake Campbell, the principal lo- cality mentioned therein. I have therefore selected as lectotype a specimen (MEL 87960) which agrees with the original description and which is the only collection I could find of this taxon that was gathered on the expedition by Babbage to the “North-western interior of South Australia” in 1858. The specimen is labelled “Scrub N. E. of Lake Gairdner”; since Lake Campbell is about 65 km NE of Lake Gairdner it is possible that this specimen is the one referred to by Mueller when he cited that locality. The name “Rhagodia deltophylla” was used by Mueller in his Second General Report (1854) but without a description. A specimen of R. spinescens which bears the name R. deltophylla was collected by him along the River Murray in January 1854 (herb. K); it presumably represents the plant referred to in the 1854 Report and is also evidently a syntype of R. spinescens var. deltophylla. Scott (1978a) gave the type citation of that variety as “South Australia, Lake Campbell; F. Mueller; Jan 1854”; he evidently had in mind the locality cited by Mueller for the specimen collected on the Babbage Expedition in 1858 and confused it with the data on the Mueller collection from the Murray River. Those specimens cited by Scott under R. deltophylla, that I have been able to check, are of R. spinescens, Chenopodium desertorum and C. curvispicatum. A similar mixture of species was included by J. M Black in his private herbarium under R. spinescens var. deltophylla. Rhagodia spinescens, as delimited in this paper, exhibits considerable variability, more, in fact, than has been accepted for any other species of Rhagodia. The leaves vary in shape from narrow elliptic c. 5 mm long to deltoid and 20 mm long, and in texture from papery to fleshy. I have not been able to discriminate clearly distinct infraspecific taxa since the dif- ferent variants appear to grade into each other and the leaves vary in shape and tex- ture on the same plant. Some variants are rather distinctive; these appear to be very limited in their distribution and since they also exhibit variability it is not clear as to whether they should be treated separately. The more easily recognisable of these variants are as follows; (1) Glossy variant. Leaves narrow elliptic to deltate, 4-20 mm long, thin or fleshy; indumentum thin below of irregularly shaped glossy thin-walled bladder hairs, almost glabrous above. Staminodes glabrous, disc almost absent. North west Victoria. Representative collections. VICTORIA: Hattah Lakes N.P., 13 Mar 1969, G. Anderson (MEL 88273); 10 mi [16.1 km] NNW of Underbool, A. C. Beauglehole 40379 (MEL); Wyperfeld N.P., A. C. Beauglehole 29535 (MEL). This variant appears to grade into the local small-leaved variant of typical R. spinescens which is found in north western Victoria and neighbouring areas of South Australia and New South Wales. (2) Bisexual variant. Leaves ovate, small, c. 5 mm long. Flowers bisexual, disc puberulous. Testa faintly rugose. Indumentum more finely furfuraceous than in typical variant. Northern South Australia. 232 Nuytsia Vol. 4, No. 2 (1983) Representative collections. SOUTH AUSTRALIA: 36 km W of Abminga siding, Lazarides 8276 (AD); Lindsay Ck., B. Major 65 (AD). (3) Central Australian foetid variant. Leaves deltate to hastiform c. 10 mm long, closely furfuraceous with very small round bladder hairs. Flowers sessile or pedi- cellate. Testa almost smooth. Simpson Desert. Representative collections. NORTHERN TERRITORY: Umbeara Stn., T. S. Henshall 1545 (NT); Beddome Range, T. S. Henshall 1519, 1533 (AD). Rather variable and possibly intergrading with the typical variant which is found in the same area. The foetid plant appears to occur on rocky hillsides whereas the typical variant is found (in this area) in sand. (4) Thin leaved variant. Leaves broadly elliptic, thin, c. 10 mm long, finely furfuraceous with very small irregular bladder hairs. Queensland and Northern Territory. Representative collections. QUEENSLAND: Laglan Stn., L. S. Smith 10315 (BRI); Taroom, R. W. Johnson 696 (BRI); 10 mi [16.1 km] N of Wandoon, Speck 1959 (CANB). NORTHERN TERRITORY: 28.7 mi [46.2 km] SW of Ooratippra H.S., G. C. Chippendale (NT 3528). Recorded as having a nauseous, fishy odour attractive to flies. Speck states that the creeping stems root at frequent intervals. The intergradation between the variants listed and those more typical of R. spinescens is such as to make any formal classification difficult to apply. Although Rhagodia spinescens as I delineate it is a very variable species, it is not so variable as has been considered by some authors. A. J. Scott (1978a), for instance, perceived it more broadly, including under this name specimens which I recognise as R. drummondii (Wilson 6088b), R. ulicina (Symon 4659), R. eremaea (Latz 114), and Chenopodium desertorum (Cunningham 384). The name R. prostrata Moq. was also cited by him as a synonym of R. spinescens whereas it is a synonym of C. des- ertorum. Rhagodia spinescens is evidently most closely related to R. drummondii and R. eremaea. Apart from differences in leaf shape the latter two species may be dis- tinguished by the absence of disc or staminodes in the female flowers, the inflor- escence shape and, in R. eremaea, by the minute irregularly shaped vesicular hairs. 10. Rhagodia eremaea Paul G. Wilson, sp. nov. Figure 5C. Rhagodia sp. B, Paul G. Wilson in Jessop, FI. Central Australia 53 (1981). Frutex dioecius ad 2 m altus. Folia alterna ovata vel elliptica 0.5-2 cm longa (petiolo 0.3-1 cm longo incluso) furfuracea cum pilis minutis vesicularibus irregulariter formatis ornatis vel glabrescentia. Flores congest! in paniculam ter- minalem pyramidalem vel paniculam gracilem spiciformen continuam dispositi. Flos masculus disco patelliformis intra pubescent!. Flos femineus staminodiis plerumque absentibus. Typus: Rudall River, 22°33'S, 122°11'E; In rocky loam by creek-line. Much- branched shrub to 1 m, fruit deep red-pink, 19 May 1971, A. S. George 10676 (holo: PERTH; iso: CANB, K, MEL, NSW). Rounded dioecious shrub to 2 m high becoming a scrambler in shade. Branches slender, closely grey-mealy when young with short branched vesicular hairs. Leaves alternate, ovate to elliptic, 0.5-2 cm long including petiole c. 1/2 length of lamina. 233 Paul G. Wilson, Tribe Chenopodieae grey mealy with minute irregularly shaped hairs, glabrescent above. Inflorescence a terminal pyramidal to slender spicoid panicle, flowers congested so as to appear con- tinuous along slender branches of panicle or (in northern Australia) in slender spicoid inflorescences with disjunct glomerules. Perianth 5-partite, densely mealy with short branched vesicular hairs. Male flower caducous; perianth globular, c. 1.5 mm diam.; filaments united into a thick saucer-shaped disc pubescent within; pistillode minute (c. 0.2 mm high). Female flower: perianth at first soft and globular c. 1 mm high, eventually hard and spreading beneath fruit becoming red within; staminodes usually absent (present in terminal flowers); ovary glabrous with a pair of slender stigmas. Berry depressed globular, c. 3 mm diameter; pericarpfleshy, red. Seed depressed globular, c. 1.3 mm diam.; testa black with reticulate ornamentation. SOUTH AUSTRALIA: Ernabella, 9 Aug. 1933, J. B. Cleland (AD); 96 km W of Mabel Creek H.S. T. R. N. Lothian 3778 (AD). WESTERN AUSTRALIA: Finucane Is., 5 Feb. 1981, G. Craig (PERTH); Lorna Glen Stn., S. Hill 25 (PERTH). NORTHERN TERRITORY: Ayers Rock,G. Chippendale (NT 663); 20 mi [32.2 km] NW of Alice Spr., D. J. Nelson 1703 (NT). Distribution. Central Australia westwards. Map 31. Habitat. Recorded in clay-loam and on rocky hillsides with Acacia kempeana, also on sand with spinifex. The flowers are arranged in glomerules each of which is subtended by a caducous linear bract. These glomerules may consist of a single terminal flower or this flower may be surrounded by others which may or may not be bracteate. The terminal flower of a glomerule in a female plant has staminodes while in the lateral flowers these are usually absent. This species exhibits the following geographic variation. In central Australia the leaf is oblong-elliptic with rounded apex and somewhat leathery when dry. In the north-west of Western Australia (Hamersley Range area) the leaf is narrow-elliptic with acute apex and is thinner in texture. The plant from the north-west and from the far north of both Western Australia and the Northern Territory usually has a very slender, spicate type of inflorescence with discontinuous glomerules of flowers, unlike the continuous, dense spikes of the central Australian plant. Another variarit found in Western Australia between Geraldton and Wittenoom has thin, broadly el- liptic to obovate leaves with rounded apices; the leaves of those specimens growing in shade are thinner and broader than those in the open. In herbaria female plants of R. eremaea have generally been identified as R. spinescens (or, more rarely, as R. crassifolia) and male plants as Chenopodium nitrariaceum. From both R. spinescens and R. crassifolia this species may be dis- tinguished by the very small irregularly shaped vesicular hairs of the leaf indumen- tum, the slender, compact inflorescence, and by the lack of staminodes in the remale flowers. Male plants may be distinguished from C. nitrariaceum by the indumentum, the scattered leaf arrangement, and the absence of spinescent branchlets. A specimen of R. eremaea collected by F. Mueller in 1856 along Sturt Creek (north-east Western Australia) is evidently one of several syntypes of Chenopodium nitrariaceum (F. Muell.) Benth. This specimen was later re-identified by Bentham (1870) as R. crassifolia which name was also applied by Ewart and Davies (1917). Nuytsia Vol. 4, No. 2 (1983) The specific epithet is taken from the Greek word eremia, a desert, alluding to the typical habitat of this plant. 11. Rhagodia parabolica R. Br., Prod. 408 (1810). Type: Inlet XII [Spencers Gulf], March 1802, R. Brown (holo: BM). R. parabolica var. typica Domin, Bibl. Bot. 89:62 (1921) nom. illeg. R. reclinata A. Cunn. ex Moq. in DC., Prod 13/2:51 (1849).--R. parabolica var reclinata (Moq.) Domin, Bibl. Bot. 89:62 (1921). Type: Liverpool Plains, May 1825, A. Cunningham 7 (holo: K). Rounded dense dioecious shrub 1-2 m high. Branches striate, somewhat flattened when young, mealy grey with minute irregularly shaped vesicular hairs. Leaves pre- dominantly opposite or subopposite, chartaceous, broadly or very broadly elliptic to broadly or very broadly ovate (rarely bluntly hastate) 1.5-4 cm long, apex acute to rounded, apiculate, base truncate to cuneate passing into the slender petiole half the length of or equal in length to lamina, minutely mealy becoming reddish brown with age. Inflorescence a broad pyramidal terminal panicle 5-8 cm long, the flowers in dis- crete glomerules along its branches. Flowers spherical c. 1 mm diameter; perianth 5- partite, closely mealy, eventually spreading in the female flower to 2.5 mm diam. but scarcely indurated. Male flowers: staminal filaments united at base into a thick cu- pular disc which is densely wooly at junction with filaments; pistillode minute. Fe- male flowers: staminodes in terminal flower of glomerule prominent, linear with min- ute antherodes and united into a deep cupular disc densely wooly within, in lateral flowers reduced or absent; stigmas subulate and spreading. Berry depressed globular c. 2.5 mm diameter. Seed depressed globular to lenticular, c. 1.8 mm diam. testa glossy with faint radial ribbing. QUEENSLAND: Online, S. T. Blake 10930 (BRI); Isle Gorge, P. Sharpe 00646 WALES: Mt Hope, G. M. Cunningham and P. Milthorpe 2511 (NSW); 6 mi [9.7 km] W by S of Jerrys Plains, R. Story 7131 (CANB). VICTORIA: Werribee Gorge, A. C. Beauglehole 39927 (MEL); Head of Coimadai Creek, 7 May 1944, J. H. Willis (MEL). Koondoolka, J. Z. Weber N()RTHERN TERRITORY; Ilpapa Swamp, T. S. Henshall 1196 (NT); [0.8 km] W of Ooraminna Rockhole, D. J. Nelson 1964 (MEL). 0.5 mi Distribution. South-east Queensland and north east New South Wales, southern Vic- toria, southern South Australia and Central Australia. Map 33. 2nd banks^'''’'''*''^ hillsides with skeletal soil, along creek banks and on This species appears to occupy four disjunct areas in Australia but the plants from each area are sufficiently alike not to warrant infraspecific recognition. In the Northern Territory occasional plants are hermaphrodite and some have tical (not horizontal) seeds, but it is not possible to establish from herbarium tenal now common these variants are. ver- ma- Paul G. Wilson, Tribe Chenopodieae 235 In Yorke and Eyre Peninsulas and north of Adelaide, plants are found with elliptic leaves and a hair type intermediate between those found in typical R. parabolica (with small discrete vesicular hairs) and that found in R. candolleana (in which the hairs collapse to form a scurfy or silvery sheen). The typical, broad-leaved R. para- bolica has been collected in the same areas so that it is possible that the narrow- leaved plant has arisen through hybridization with another species such as R. preissii or R. candolleana. Doubtful or incorrect records Blituw virgatum L. (= Chenopodium foliosum (Moench) Asch.) Recorded by J. D. Hooker, FI. Tasman. XCVI (1959) as being found in Australia. I have seen no specimens which support this record. Chenopodium bonus-henricus L. In the Victorian Naturalist 22:79 (1905) it is recorded that H. B. Williamson exhibited a specimen of this species collected from ‘Hopkins mouth’, Victoria. This record is referred to by Ewart and Tovey (1909) and by Ewart (1931) who state that it was naturalized in south-west Victoria in 1905. 1 have been unable to locate the Hopkins mouth collection but a specimen collected by Williamson nearby at Hawksdale in 1900 (MEL 88604 and 88605), and identified by Ewart as C. bonas-henricus, is of a species of Atriplex. Chenopodium iicifolium Smith. Recorded as a weed in Brisbane by F. M. Bailey, Queensland Agric. J. 31: 321 (1913). I have seen no specimens to validate this record. Chenopodium rubrum L. Recorded by Audas and Morris (1925) as widely spread in Victoria and by Ewart (1931) as naturalized at North Melbourne in 1910 and widely spread in Victoria. I have seen no Aus- tralian collection of this species. The following two specimens in herb. MEL which had been deter- mined as C. rubrum are, in fact, both Chenopodium glaucum: Railway Reserve, N. Melbourne, May 1910, C. French and R. J. Tovey (MEL 88793); Coode Island, 23 March 1912, R. J. Tovey (MEL 88794). Chenopodium viride L. Recorded by J. D. Hooker, FI. Tasman. CIX (1859) as being found in Australia. I have seen no specimen to support this record. Excluded names Chenopodium insulare J. Black, Trans. Roy. Soc. S. Austral. 69: 309 (1945) = Suaeda australis (R. Br.) Moq. See Hj. Eichler, Supplement to J. M. Black’s Flora of South Australia (1965). Chenopodium australe R. Br,, Prod. 407 (1810) = Suaeda australis (R. Br.) Moq. Rhagodia eschscholtziana Fenzl, Denkschr. Akad, Wien (Math. -Nat.) 1: 257 (1850). Described from Oahu in the Hawaiian Islands; the description agrees with that for Chenopodium oahuense (Meyen) Aellen (Atriplex oahuensis Meyen 1835). Acknowledgements I am grateful to the Directors of the State, Commonwealth, and University herbaria in Australia and to the herbarium of the DSIR in Christchurch, for the loan of material. I am also grateful to the Directors of the herbaria in Paris (P), Florence (FI), Lund (LD), London (BM), and Kew (K) for the loan of type specimens. The National Herbarium of New South Wales (NSW) kindly sent manuscript herbarium notes on many of the species treated in this paper. Mr W. R. Sykes (CHR) sent com- ments on several of the species found in New Zealand. The present study was commenced in 1978 while a guest of the Division of Plant Industry, CSIRO, Canberra. I am grateful to its Chief, Dr W. J. Peacock, for ex- tending an invitation and to Dr Hj. Eichler, then Curator of the Herbarium Aus- traliense, for making space and facilities available and for his personal support. The SEM photographs, figures 4 and 5, were prepared by Mr Stuart Craig of the Division of Plant Industry, Canberra and figures 6 and 7 by Mr Terry Robertson of the Queen Elizabeth II Medical Centre, Perth, to both of whom I wish to express my appreci- ation for their careful work. 236 Nuytsia Vol. 4, No. 2 (1983) References Aellen, P. (1929). Systematik der Chenopodium- Arten Amerikas, vorwiegend auf Grand der Sammlung des United States National Herbarium in Washington, D. C. Feddes Repert. 26: 31-64, 119-160. Aellen, P. (1929). Die wolladventiven Chenopodien Europas. Verb. Naturf. Ges. Basel 41: 77-104. Aellen, P. (1930). Die systematische Stellung und Gliederung der R. Brownschen Gattung Dysphania. Bot. Jahrb. Syst. 63:483-492. Aellen, P. (1933). Die Arten der Sect. Orthosporutn R.Br. der Gattung Chenopodium L. Verb. Naturf. Ges. Basel 44: 308-318. Aellen, P. (1937-1938). Revision de Australischen und neuseelandischen Chenopodiaceen 1. Bot. Jahrb. Syst. 68: 345-434. Aellen, P. (1939). Klarstellung von Chenopodium triandrum Forster und einigen Australischen Chenopodien. Candollea 8: 5-11. Aellen, (1960-1961). Chenopodiaceae. In Hegi, ‘Illustrierte Flora von Mitteleuropa’ ed. 2, 3: 533-747. (Carl Hanser Verlag: Mlinchen.) Aellen, P. (1961). Dysphaniaceae. In Hegi, ‘Illustrierte Flora von Mitteleuropa’ ed. 2, 3: 748. (Carl Hanser Verlag; Miinchen.) Aellen, P. and Just, T. (1943). Key and synopsis of the American species of the genus Chenopodium L. Amer. Midi. Naturalist 30: 47-76. Allan, H. H. (1940). ‘A handbook of the naturalized flora of New Zealand’. (Government Printer: Wellington.) Allan, H. H. (1961). ‘Flora of New Zealand’ vol. 1. (Government Printer: Wellington.) Ascherson, P. and Graebner, P., (1913). ‘Synopsis der Mitteleuropaischen Flora’ vol. 5. (Verlag von Gebriider Borntraeger: Leipzig.) Audas, J. W. and Morris, P. F. (1925). ‘Supplement to Professor Ewart’s “Weeds, poison plants and natu- ralized aliens of Victoria” ’. (Melbourne University Press: Melbourne.) Battandier, J.-A. (1888-1890). In J.-A. Battandier, and L. Trabut, ‘Flore de L’Algerie’ vol. 1. Dicotyledones. (Adolphe Jourdan: Alger.) Beauge, A. (1974). ‘Chenopodium album et especes affines.’ (Societe d’Edition d’Enseignement Superieur: Paris.) Bentham, G. (1870). ‘Flora Australiensis’ vol. 5. (Reeve: London.) Black, J. M. (1912). Additions to the flora of South Australia. Trans. & Proc. Roy. Soc. S. Austral. 36: 171- 172. Black, J. M. (1924). ‘Flora of South Australia’ pt. 2. (Government Printer: Adelaide.) Black, J. M. (1948). ‘Flora of South Australia’ ed. 2, pt. 2. (Government Printer: Adelaide.) Black, R. F. (1954). The leaf anatomy of Australian members of the genus Atriplex. 1. Atriplex vesicaria Heward and A. nummularia Lindl. Austral. J. Bot. 2: 269-286. Brenan, J. P. M. (1954). Chenopodiaceae. In W. B. Turrill and E. Milne-Redhead, ‘Flora of tropical East Africa’. (Crown Agents: London.) Brenan, J. P. M. (1956). A new section of the genus Chenopodium from Africa. Kew Bull. 11: 165-167. Brian, R. C. and Cattlin, N. D. (1968). The surface structure of leaves of Chenopodium album L. Ann. Bot. 32: 609-612. Britton, N. L. and Brown, A. (1913). ‘An illustrated flora of the northern United States, Canada and the British possessions’ ed. 2, vol. 2. (Charles Scribner’s Sons; New York.) Brown, R. (1810). ‘Prodromus florae Novae Hollandiae et Insulae Van-Diemen’. (Richard Taylor: London.) Cheeseman, T. F. (1884). Notice of the discovery of the genus Rhagodia in New Zealand. Trans, and Proc. New Zealand Instit. 16: 408-409. Cheeseman, T. F. (1914). ‘Illustrations of the New Zealand flora’ vol. 2. (Government Printer: Wellington.) Cheeseman, T. F. (1925). ‘Manual of the New Zealand Flora.’ (Government Printer; Wellington.) Christensen, C. (1922). Index to Peter Forsskal: Flora aegyptiaco-arabica 1775 with a revision of Her- barium Forskalii contained in The Botanical Museum of the University of Copenhagen. Dansk Bot. Arkiv. 4/3: 1-54. Cole, M. J. (1961). Interspecific relationships and intraspecific variation of Chenopodium album L. in Britain. 1. The taxonomic delimination of the species. Watsonia 5: 47-58. Cole, M. J. (1962). Interspecific relationships and intraspecific variation of Chenopodium album L. in Britain. 2. The chromosome numbers of Chenopodium album and other species. Watsonia 5: 117-122. Curtis, W. M. (1967). ‘The student’s flora of Tasmania’ pt. 3. (Government Printer: Hobart.) Paul G. Wilson, Tribe Chenopodieae 237 Domin, K. (1925). Dysphania. Biblio. Bot. 89. 102-104. Eckardt, T. (1967). Bliitenbau and Blutenentwicklung von Dysphania myriocephala Benth. Bot. Jahrb. Syst. 86: 20-37. Eckardt, T. (1967). Vergleich von Dysphania mit Chenopodium und mit Illecebraceae. Bauhinia 3: 327-344. Ecka^rdt, T. (1968). Zur Blutenmorphologie von Dysphania plantaginella F.v.M. Phytomorphology 17: 165- Eckardt, T. (1969). Morphology and Systematic position of Dysphania. 11th International Botanical Con- gress, Abstracts. (Seattle.) Engstraad, L. and Gustafsson, M. (1973). Drawings of Scandinavian plants: Chenopodium murale L. Bot. Not. 127: 275-276. Esau, K. (1977). Anatomy of seed plants. Ed. 2. (John Wiley and Sons: New York.) Ewart, A. J. (1908). Contributions to the flora of Australia, no. 8. Victorian Nat. 24: 190-194. Ewart, A. J. (1931). ‘Flora of Victoria.’ (Melbourne University Press: Melbourne.) Ewart, A. J. and Davies, O. B. (1917). ‘The flora of the Northern Territory.’ (McCarron, Bird and Co.: Melbourne.) Ewart, A. J. and Tovey, J. R. (1909). ‘The weeds, poison plants, and naturalized aliens of Victoria.’ (Government Printer: Melbourne.) Giusti, L. (1964). Chenopodiaceae. In A. Cabrera, ‘Flore de la Provincia de Buenos Aires’ vol. 4/3: 113-127. (Coleccian Cientifica del I.N.T.A.: Buenos Aires.) Giusti, L. (1970). El genero Chenopodium en Argentina. 1. Numeros de cromosomas. Darwiniana 16: 98- 105. Hellmuth, E. O. (1968). Eco-physiological studies on plants in arid and semi-arid regions in Western Aus- tralia. 1. Autecology of Rhagodia baccata (Labill.) Moq. J. Ecol. 56: 319-344. Hooker, J. D. (1880). Chenopodiaceae. In G. Bentham and J. D. Hooker, ‘Genera plantarum’ vol. 3. (Reeve: London.) Jorgensen, P. M. (1973). The genus Chenopodium in Norway. Norw. J. Bot. 20: 303-319. Kjellmark, S. (1934). Einige neue Chromosomenzahlen in der Familie Chenopodiaceae. Bot. Not. 136-149. Maiden, J. H. and Betche, E. (1916). ‘A census of New South Wales Plants.’ (Government Printer: Sydney.) Moquin-Tandon, C.H.B.A. (1840). “Chenopodearum monographica enumeratio.” (P. J. Loss: Paris.) Moquin-Tandon, C.H.B.A. (1849). Salsolaceae. In A. P. de Candolle, “Prodromus systematis naturalis regni vegetabilis” vol. 13/2. (Teuttel et Wiirtz: Paris.) Mueller, F. (1859). “Report on the plants collected during Mr. Babbage’s expedition into the north-west interior of South Australia in 1858.” (Government Printer: Melbourne.) Mueller, F. (1869-1871). “Fragments Phytographiae Australiae” vol. 7. (Government Printer: Melbourne.) Mueller, F. (1886). New Australian plants. Australas. J. Pharm. 1:429-431. Mueller, F. (1887). New Australian Plants. Bot. Centralbl. 29:114-115. Mueller, F. (1889-1891). “Iconography of Australian salsolaceous plants.” (Government Printer: Melbourne.) Murr, J. (1910). Australische Chenopodien. Allg. Bot. Zeit. 16:55-58. Murr, J. (1926). Meine Phanerogamen-Bastarde. Vierteliahrsschr. Gesch. Landesk. Voralbergs 10:185-194. 194. Murr, J. and Thellung, H. (1915). Chenopodium auricomiforme. Vierteliahrsschr. Naturf. Ges. Zurich 60:432. Osmond, C. B. et al. (1969). Ion absorption in Atriplex leaf tissue. II. Secretion of ions to epidermal blad- ders. J. Biol. Sci. 22:797-814. Pax, F. A. (1889). Dysphania. In A. Engler and K. Prantl, “Die natiirlichen Pflanzen-familien” 3, lb:92. (Verlag von Wilhelm Englemann:Leipzig.) Pax, F. A. (1927). Zur Phylogenie der Caryophyllaceae. Bot. Jahrb. Syst. 61:223-241. Pax, F. A. and Hoffmann, K. (1934). Dysphaniaceae. In A. Engler and K. Prantl, “Die naturlichen pflanzenfamilien” ed. 2, 16c: 272-274. (Duncker and Humblot: Berlin.) Planchuelo, A. M. (1975). Estudio de los frutos y semillas del genero Chenopodium en la Argentina. Darwiniana 19:528-565, Probst, R. (1949). “Wolladventiflora Mitteleuropas.” (Vogt-Schild Ag.: Solothurn.) Rodway, L. (1903). “The Tasmanian flora.” (Government Printer: Hobart.) Scott, A. J. (1978a). Rhagodiinae: A new subtribe in the Chenopodiaceae. Feddes Repert. 89:1-11. 238 Nuytsia Vol. 4, No. 2 (1983) Scott, A. J. (1978b). A review of the classification of Chenopodium L. and related genera (Chenopodiaceae). Bot. Jahrb. Syst. 100:205-220. Shapter, R. E. (1941). A preliminary investigation of the yield and composition of the oil distilled from Chenopodium ambrosioides (Linn.) var. anthelminticum (Gray). J. Council Sci. Ind. Rea. 14:201-208. Standley, P. C. (1931). The Chenopodiaceae of northwestern South America. Field Mus. Nat. Hist. Bot 11:115-126. Sykes, W. R. 11977). "Kermadec Islands Flora.” Bulletin no. 219. (New Zealand D.S.I.R.: Wellington.) Ulbrich, E. (1934). Chenopodiaceae. In Engler et Prantl, “Die natiirlichen Pflanzenfamilien” ed. 2, 16c:379- 584. (Duncker and Humblot: Berlin.) Uotila, P. (1978). Variation, distribution and taxonomy of Chenopodium suecicum and C. album in N. Europe. Acta Bot. Fennica 108:1-35. Voroschilov, V. N. (1942). Revision des especes de Chenopodium de la section Ambrina (Spach) Hook.f. J. Bot. URSS 27 No. 3/4:33-47. White, C. T. (1921). Illustrated notes on the weeds of Queensland. No. 21. Goosefoot (Chenopodium triangulare, R.Br.). Queensld. Agric. J. n.s. 15:171-172. Willis, J. H. (1973). “A handbook to plants in Victoria.” vol. 2 (Melbourne Univ. Press: Victoria.) Wilson, H. D. (1980). Artificial hybridization among species of Chenopodium sect. Chenopodium Svst Bot. 5:253-263. Wood, J. G. (1923). On transpiration in the field of some plants from the arid portions of South Australia with notes on their physiological anatomy. Trans. Roy. Soc. S. Austral. 47:259-278. Paul G. Wilson, Tribe Chenopodieae 239 Figure 4. SEM photos of seeds: A — Chenopodium erosum (upper surface with pericarp attached), from A. C. Beauglehole 37201 (PERTH); B — Chenopodium murale (lower surface), from CANB 31776; C — Chenopodium auricomiforme (lower surface), from CANB 36430; D — Chenopodium curvispicatum (lower surface), from H. Schodde 1082 (CANB); E — Einadia nutans subsp. nutans (lower surface), from AD95826008; V- Einadia hastala (lower surface), from CBG04660t. 240 Nuytsia Vol. 4, No. 2 (1983) Figure 5. SEM photos of seeds: A — Einadia allanii (upper surface), from P. Waidle (CHR125728); B — Einadia triandra (upper surface), from iV. J. A. Simpson 7160 (CHR); C — Rhagodia eremaea (lower surface), from P. K. Latz 139 (PERTH); D — Rhagodia spinescens (upper surface), from ADW34053; E — Rhagodia acicularis (upper surface), from P. G. Wilson 11713 (PERTH); F — Rhagodia ulicina (upper surface), from AD96503078. Paul G, Wilson, Tribe Chenopodieae 241 Figure 6. SEM photos of seeds: A—Dysphania glomulifera subsp. glomulifera (side view), from C. Andrews, April 1902 (PERTH); B—Dysphania glomulifera subsp. eremaea (side view showing em- bryo-groove), from R. D. Royce 1493 (PERTH); C^Dysphania glandulosa (side view showing embryo- groove), from A. S. George 717 (PERTH); D—Dysphania platycarpa (side view), from A. S. George 8927 (PERTH); E-F—Dysphania sphaerosperma (side view showing vertical embryo-groove). 242 Nuytsia Vol. 4, No. 2 (1983) Figure 7. SEM photos of seeds; A — Dysphania kalpari (side view showing raphe), from D. J. Nekon 830 (NT); B — Dysphania littoralis (obliquely lateral view), from P. G. Wilson, 24 Feb. 1977 (PERTH); C-D — Dysphania rhadinostachya (C — obliquely lateral view and D — upper surface, showing horizontal embryo-ridge), from R. J. Ch/niioclf 385,1 (PERTH); E — Dysphania plan tagfne/la (side view), from A. C. Beauglehole 48362 (PERTH); F — Chenopodium gaudichaudianum (upper surface), from P. G. Wilson 9911 (PERTH). Paul G. Wilson, Tribe Chenopodieae 243 244 Nuytsia Vol. 4, No. 2 (1983) Paul G. Wilson, Tribe Chenopodieae 245 22493 — ( 8 ) 246 Nuytsia Vol. 4, No. 2 (1983) Paul G. Wilson, Tribe Chenopodieae 248 Nuytsia Vol. 4, No. 2 (1983) Paul G. Wilson, Tribe Chenopodieae 250 Nuytsia Vol. 4, No. 2 (1983) Paul G. Wilson, Tribe Chenopodieae 251 252 Nuytsia Vol. 4, No. 2 (1983) Paul G. Wilson, Tribe Chenopodieae 253 254 Nuytsia Vol. 4, No. 2 (1983) Paul G. Wilson, Tribe Chenopodieae 255 256 Nuytsia Vol. 4, No. 2 (1983) Paul G. Wilson, Tribe Chenopodieae 257 258 Nuytsia Vol. 4, No. 2 (1983) Paul G. Wilson, Tribe Chenopodieae 259 260 Nuytsia Vol. 4, No. 2 (1983) Index to scientific names Numerals in bold type represent main entries of accepted names. New names are given in bold type. Other accepted names are in Roman. Synonyms are in italics. Amaranthus L. 197 enervis (F. Muell.) F. Muell. 197 tenuis Benth. 197 Ambrina Spach 166 sect. Adenois Moq. 166 sect. Botryois Moq. 180 carinata (R. Br.) Moq. 173 var. parvifolia Moq. 174 pinnatisecta Spach 168 punailio (R. Br.) Moq. 172 Atriplex L. 235 glomuHfera Nees 183 oahuensis Meyen 235 vesicaria Benth. Blitum L. sect. Orthosporum (R. Br.) C. Meyer 169 Atriplicinum F. Muell. 197 caiinatum (R. Br.) Steudel 173 cristatum F. Muell. 176 glandulosum Moq. 172 var. parvi folium (Moq.) Moq. 174 pumilio (R. Br.) Steudel 172 virgatum L. 235 Chenopodium L. 139 sect. Ambrina J. D. Hook. 169 sect. Ambrina (Spach) J. D. Hook. 166 subg. Ambrosia A. J. Scott 166 sect. Atriplicina Aellen 197 sect. Auricoma Aellen 161 sect. Botryoides C. Meyer 180 sect. Boti^'ois (Moq.) Moq. 180 sect. Carinata Standley 169 sect. Chenopodiastrum Moq. 144 sect. Chenopodium 134 subg. Chenopodium 139 sect. Degenia Aellen 143 sect. Desertorum Paul G. Wilson 151 sect. Dysphania (R. Br.) Aellen 181 sect. Euchenopodium Ulbrich 144 subsect. Lejosperma Aellen et Iljin 144 sect. Leprophyllum Dumort. 144 sect. Margaritaria Brenan 180 sect. Orthosporum R. Br. 139, 169 sect. Polygonoidea Aellen 199 sect. Pseudoblitum J. D. Hook. 141 sect. Rhagodioides Benth. 164 sect. Roubieva (Moq.) Asch. et Graebner 166 sect. Tetrasepala Aellen 181 sect. Thullingia Aellen 197 acuminatum Willd. 164 album L. 144 subsp. opulifolium (Koch et Ziz) Maire 145 var. reticulatum (Aellen) Uotila 145 aWanij Aellen 210 ambiguum R. Br. 141 var. majus Moq. 141 var. minus Moq. 141 ambrosioides L. 167 var. anthelminticum (L.) A. Gray 167 f. spathulatum (Moq.) Aellen 167 anidiophyllum Aellen 152, 155 antarcticum (J. D. Hook.) J. D. Hook. 198 anthelminticum L. 167 atriplicinum (F. Muell.) F. Muell. 197 auricomiforme Murr et Thellung 162 auricomum Lindl. 163 f. subglabrum Aellen 163 australasicum Moq. 200 australe R. Br. 235 baccatum Labill. 223 berlandieri Moq. 146 bi forme Nees 148 blackianum Aellen 188 xbontei Aellen 171 n-var. cristatiforme Aellen 171 n-var. submelanocarpum Aellen 171 botrys L. 180 bonus-henricus L. 235 browneanum Roemer et Schultes 144 carinatum R. Br. 173 f. foliosum Domin 174 var. holopterum (Thellung) Aellen 174 f. leucocarpum Aellen 176 var. melanocarpum J. Black 175 f. rubellum Aellen 172 carnosulum Moq. 148 xchristii Aellen 171 n-var. intermedium Aellen 171, 172 n-var. semiconnatum Aellen 171, 172 cochlearifolium Aellen 152 congestum J. D. Hook. 148 congolanum (Haum.) Brenan 180 cristatum (F. Muell.) F. Muell. 176 var. holopterum Thellung 174 curvispicatum Paul G. Wilson 159 desertorum (J. Black) J. Black 152 subsp. anidiophyllum (Aellen) Paul G. Wilson 155 subsp. desertorum 153 subsp. microphyllura Paul G. Wilson 154 subsp. rectum Paul G. Wilson 158 subsp. virosum Paul G. Wilson 157 Paul G. Wilson, Tribe Chenopodieae detestans J. Kirk 147 erosum R. Br. ISO ficifolium Smith 235 foliosum (Moench) Asch. 235 furfuraceum Moq. 214 gaudichaudianum (Moq.) Paul G. Wilson 160 giganteum D. Don 145 glandulosum (Moq.) F. Muell. 172 glaucum L. 139 var. ambiguum (R.Br.) J. D. Hook. 141 subsp. ambiguum (R.Br.) Thell. 141 var. littorale Rodway 141 f. minus (Moq.) Aellen 141 var. paschale Fuentes 141 holocarpum J. Black 175 hoJopterum (Thell.) Thell. et Aellen 174 hubbardii Aellen 151 inflatum Aellen 193, 194 insulare J. Black 235 integrifolium Vorosch. 168 lanceolatum R.Br. 144 littorale (L.) Thunb. 141, 188 littorale Moq. 141 lycioides Gand. 164 macrospermum J. D. Hook. 143 subsp. halophilum (Philippi) Aellen 143 f. subviride Aellen 143 melanocarpum (J. Black) J. Black 175 f. leucocarpura (Aellen) Paul G. Wilson 176 f. melanocarpum 175 microphyllum F. Muell. 152, 154 var. desertorum J. Black 152 microphyllum Thunb. 152 multifidum L. 168 f. intermedium Aellen 168 f. spathulatum Aellen 168 murale 148 var. bi forme (Nees) Moq. 148 var. sinuosifrons Murr 148 myriocephalum (Benth.) Aellen 183 nitrariaceum (F. Muell.) Benth. 164 oahuense (Meyen) Aellen 235 olidum Curtis 146 opulifolium Koch et Ziz 145 osbornianum Aellen 192 plantnginellum (F. Muell.) Aellen 188 polygonoides (Murr) Aellen 212 preissii (Moq.) Diels 221 probstii Aellen 144 f. lanceolatum Aellen 144 f. parvo-angustifolium Aellen 144 f. subintegrifolium Probst 144 prostratum (Pallas) Roemer et Schultes 139 pseudomicrophyllum Aellen 152, 154 pumilio R.Br. 172 f. glandulosum (Moq.) Aellen 172 var. oblongifolium J. Black 172 f. rubellum (Aellen) Aellen pussillum J. D. Hook. 179 22493 — ( 9 ) 261 reticulatum Aellen 144 rhadinostachyum F. Muell. 193 var. paniculatum F. Muell. et Tate 193 rubrum L. 142, 235 saxatile Paul G. Wilson 179 scabricaule Speg. 148 serotinum L. 145 simulans (Tate) F. Muell. 192 spathulatum Sieberex Moq. 167, 168 stellulatum Aellen 206, 208 stellulatum (Benth.) Aellen 208 suecicum Murr 145, 150 suffruticosum Willd. 168 subsp. remotum Vorosch. 168 tenue Colla 139 triandrum G. Forster 205 var. lanuginosum F. E. Haviland 152, 154 triangulare Forsskal 145 triangulare R.Br. 206 var. angustifolium Benth. 212 subsp. convohulinum Murr 202 var. convolvulinum (Murr) Maiden 202 subsp. polygonoides (Murr) Murr 212 var. polygonoides Murr 212 var. stellulatum Benth. 208 trigonocarpum Aellen 171 trigonon Roemer et Schultes 206 truncatum Paul G. Wilson 177 ulicinum Gand. 219 viride L. 235 vulvaria L. 146 Dysphania R.Br. 180 sect. Caudatae A. J. Scott 181 sect. Tetrasepalae (Aellen) A. J. Scott 181 benthamiana Domin 183, 184 glandulosa Paul G. Wilson 186 glomulifera (Nees) Paul G. Wilson 183 subsp. glomulifera 184 subsp. eremaea Paul G. Wilson 184 inf lata (Aellen) A. J. Scott 193 kalpari Paul G. Wilson 195 littoralis R.Br. 188 myriocephala Benth. 183 plantaginella F. Muell. 190 platycarpa Paul G. Wilson 189 rhadinostachya (F. Muell.) A. J. Scott 193 subsp. inflata (Aellen) Paul G. Wilson 194 subsp. rhadinostachya 194 simulans Tate 192 sphaerosperma Paul G. Wilson 191 valida Paul G. Wilson 187 Einadia Raf. 199 allanii (Aellen) Paul G. Wilson 210 hastata (R.Br.) A. J. Scott 211 linifolia (R.Br.) Raf. 200, 204 nutans (R.Br.) A. J. Scott 200 subsp. eremaea Paul G. Wilson 204 subsp. linifolia (R.Br.) Paul G. Wilson 204 262 Nuytsia Vol. 4, No. 2 (1983) var. lini folia (R.Br.) A. J. Scott 204 subsp. nutans 202 subsp. oxycarpa (Gauba) Paul G. Wilson 203 polygonoides (Murr) Paul G. Wilson 212 triandra (G. Forster) A. J. Scott 205 trigones (Roemer et Schultes) Paul G. Wilson 206 subsp. leiocarpa Paul G. Wilson 209 subsp. stellulata (Benth.) Paul G. Wilson 208 subsp. trigonos 209 Euxolus Raf. enervis F. Muell. 197 Monolepis Schrader 181, 197, 199 Orthospermum (R.Br.) Opiz 169 Orthosporum (R.Br.) T. Nees 169 bonus -henriciis (L.) T. Nees rubrum (L.) T. Nees Rhagodia R.Br. 213 acicularis Paul G. Wilson 221 baccata (Labill.) Moq. 223 var. angustifolia Moq. 224 subsp. baccata 223. var. candolleana (Moq.) Moq. 214 var. congesta (J. D. Hook.) J. D. Hook. 148 subsp. dioica (Nees) Paul G. Wilson 225 var. linearis (Benth.) J. Black 223 var. parx'ifolia Moq. 215 billardieri R.Br. 223 var. congesta (J. D. Hook) Benth. 148 var. linearis Benth. 223 candolleana Moq. 214 subsp. argentea Paul G. Wilson 215 subsp. candolleana 215 cbenopodioides Moq. 200, 202 congesta (J. D. Hook.) Moq. 148 coralliocarpa S. Moore 160 crassifolia R.Br. 218 var. latifolia Benth. 228 cygnoruni Gand. 223 deltophylla (F. Muell.) A. J. Scott deltophylla F. Muell. dioica Nees 223, 225 drummondii Moq. 229 eremaea Paul G. Wilson 232 eschscholtziana Fenzl 235 gaudichaudiana Moq. 160 hastata R.Br. 211 latifolia (Benth.) Paul G. Wilson 228 subsp. latifolia 228 subsp. recta Paul G. Wilson 228 linifolia R.Br. 200, 204 nitrariacea F. Muell. 164 nutans R.Br. 200 var. fallacina Domin 202 var. oxycarpa Gauba 202 var. parvifolia Moq. 202 obovata Moq. 221, 222 parabolica R.Br. 234 var. reclinata (Moq.) Domin 234 var. typica Domin 234 parvifolia Moq. 218 preissii Moq. 221 subsp. obovata (Moq.) Paul G. Wilson 222 subsp. preissii 222 prostrata Moq. 152 radiata Nees 223 reclinata Nees 234 spinescens R.Br. 230 var. deltophylla F. Muell. 230 triandra (G. Forster) Aellen 205 ulicina (Gand.) Paul G. Wilson 219 Roubieva Moq. 166 multifida (L.) Moq. 168 Salsola L. carinata (R.Br.) Sprengel 173 Scleroblitum Ulbrich 197 atriplicinum (F. Muell.) Ulbrich 197 Suaeda Forsskal ex Scop. 199 australis (R.Br) Moq. 235 linifolia Pallas 200 Tetragonia L. iraplexicoma (Miq.) J. D. Hook. 218 Publication date of Nuytsia Volume 4 Number 1: 14 Oct. 1982. 22493/1 1 /82— 1 M— L/3286 WILLIAM C BROWN. Government Printer. Western Australia Notes for Authors Nuytsia publishes papers relating to the flora of Western Australia. All papers are refereed outside the Western Australian Herbarium. The Herbarium reserves the right to reject papers. Manuscripts must be submitted in duplicate. They should be typewritten, double spaced, with 2.5 cm margins and no underlining. Pages should be numbered. The de- sired position of text figures may be pencilled in the margin. An abstract should be provided. For style and layout, recent numbers of Nuytsia should be followed. Galley proofs will be forwarded to authors for checking. Reprints may be ordered at the author’s expense. CONTENTS A taxonomic revision of the tribe Chenopodieae (Chenopodiaceae) in Australia. By Paul G. Wilson 135 Publication date of Volume 4 Number 1 262