ISSN 0085-4417 NUYTSIA Bulletin of the Western Australian Herbarium Vol. 2 No. 1 1975 Department of Agriculture of Western Australia r* ISSN 0085—4417 NUYTSIA Bulletin of the Western Australian Herbarium Edited by A. S. George Volume 2 1975-1979 Department of Agriculture of Western Australia NUYTSIA VOLUME 2 Dates of publication Number I Number 2 Number 3 Number 4 Number 5 Number 6 pages 1-84 pages 85-124 pages 125-166 pages 167-264 pages 265-334 pages 335-378 27 June 1975 25 May 1976 4 April 1977 7 February 1978 11 December 1978 21 December 1979 CONTENTS Page BENL, G. — gfl/Y/ne/'i Beni. sp. nov. (Amaranthaceae) 93 Ptilotiis tetrandms sp, nov. (Amaranthaceae) 232 Three new species of Ptilotiis (Amaranthaceae) from Western Australia 341 BENNETT, Eleanor M. — New taxa and new combinations in Australian Pittosporaceae 184 BROOKER, M. I. H. — Six new taxa of Eucalyptus from Western Aus- tralia 103 BROOKER, M. I. H., and BLAXELL, D. E. — Five new species of Eucalyptus from Western Australia 220 COURT, A. B. — Three new species of Acacia (Mimosaceae) from Western Australia 168 GEORGE, A. S.^Hakea acuieata (Proteaceae) a rare and endangered new species from Western Australia 375 GREEN, J. W. — Corynanthera, a new genus of Myrtaceae (Subfamily Leptospermoideae, Tribe Chamelaucieae) 368 HOPPER, S. D. — A new species of Anigozanthos Labill. from the Murchison River, Western Australia 181 Nomenclatural notes and a new taxa in the Conostylis acuieata group (Haemodoraceae) 254 KENNEALLY, K. F., WILSON, P. G. and SEMENIUK, V.— A new character to distinguish vegetative material of the mangrove genera Bruguiera and Rhizopliora (Rh\zophoraceae) 178 KUCHEL, R. H. — A taxonomic revision of the genus Chamaexeros Benth. (Xanthorrhoeaceae) 118 MASLIN, B. R.— Studies in the genus Acacia (Mimosaceae) — 5. Mis- cellaneous new phyllodinoLis species 96 Studies in the genus ylrac/n (Mimosaceae)— 6. Miscellany 145 Studies in the genus Acacia (Mimosaceae) — 7. The taxonomy of some diaphyllodinoLis species 200 Studies in the genus Acacia ( Mimosaceae)— 8. A revision of the Uninerves-Triangulares, in part (the tetramerous species) 266 Studies in the genus Acacia (Mimosoideae)— 9. Additional notes on the Series Pulchellae Benth 354 MASLIN, B. R. and WHIBLEY, D. J. E. — The identity of Acacia /c/o/^Z/jV/a Benth. (Mimosaceae) 162 MITCHELL, Andrew S. — A new Hihiscus (Malvaceae) from Central Australia 336 ORCHARD, A. E. — Taxonomic Revisions in the family Haloragaceae 11. Further notes on Haloragis, Haloragoclendron and Gonocarpus. 126 STONE, B. C.— Notes on the genus Pandanus (Pandanaceae) in Western Australia 236 TINDALE, Mary D. and MASLIN, B. R.— Two new species of Acacia from Western Australia 86 WILSON, P. G. — A taxonomic Revision of the genus Maireana (Cheno- podiaceae) 2 91669/1/80— IM WILLIAM C. BROWN, Government Printer, Western Australia ISSN 0085 4417 NUYTSIA Bulletin of the Western Australian Herbarium Vol. 2 No. 1 1975 Department of Agriculture of Western Australia ( 1)44205 A Taxonomic Revision of the genus Maireana (Chenopodiaceae) By P. G. Wilson Abstract A taxonomic revision is provided for the genus Maireana (Chenopodiaceae) which as circumscribed includes the Australian species previously placed in the genus Kochia. Nine species and one subspecies are described as new; 48 new combinations are made. The Australian genera within the subtribe Kochiinae are briefly described and their relationships discussed. Introduction The genus Maireana, as here delimited, encompasses those species of Australian Chenopodiaceae which have commonly been placed in the genus Kochia. There appear to the author to be strong reasons, based on mor- phological data, for considering the Australian species to be distinct from Kochia s.str. (a predominantly northern hemisphere genus) and the acceptance of this distinction appears on strictly taxonomic grounds to be necessary. The alternative, that of continuing to consider the Australian species as con- generic with the Kochia species of the northern hemisphere, raises problems of nomenclature with unsettling consequences. It has always been recognised that Kochia and Basxia (both in the strict sense) are very closely allied and were considered by most authors in the 19th century, and in several recent European floras, to be congeneric. This synonymy is taxonomically sound, and thus, if it is considered that the Australian taxa here referred to Maireana are in fact true species of Kochia. and the latter genus is considered by European botanists to be congeneric with Bassia. then it follows that the Australian taxa in turn will have to be transferred to the last genus, it being the earliest. In addition to the complication outlined above it is here considered that the Australian species commonly placed in Bassia are in fact generically dis- tinct and should, for the most part, be transferred to Sclerolaena. A situation could thus arise in which the Australian species of 'Kochia' were placed in Bassia and most of the 80-1- Australian species Bassia' were transferred to Sclerolaena. Although the author considers that the acceptance of the genus Sclerolaena is both necessary and inevitable, he does not consider it either taxonomically sound, or from the practical viewpoint, desirable, to transfer the Australian Kochia species to Bassia. Historical Survey of the Use of the Names Kochia and Bassia The genus Bassia was described by Allioni in 1766. It was based on the species later known as Bassia nniricata (L.) Aschers., a plant in which short spines develop on the hack of the perianth segments when in fruit. The genus Kochia was described by Roth in 1801. It was based on the one species K. arenaria (Maerklin) Roth ( K. laniflora (Gmel.) Borbas). In this species the perianth segments give rise to short, horizontal, chartaceous, wing-like outgrowths. Later in 1801 (or 1802?) (and in the same volume of the same periodical) Maerklin described the genus IVillemetia: he based it on two species, IV. arenaria (Maerklin) Maerklin ( - K. laniflora (Gmel.) Borbas) and II'. lanala Maerklin which was based on "Salsola laniflora Linn.” ( K. laniflora) but provided a description for the latter which correctly applies to Ba.ssia hyssopifolia (Pall.) Volk. Robert Brown (1810) discussed the taxonomy of the Kochia group and suggested (but did not adopt the suggestion) that it could be divided into two genera, the genus Kochia containing those species with spiny appendages 2 and the genus WiUemetia containing those with membranous appendages. All the species he dealt with (which were members of the Australian Hora) possessed membranous appendages. He did not recognise any spiny append- aged Kochia species as being present in Australia. C. F. Ledebour (1829) followed Brown in using the name Kochia to co\'er the Bassiu-Koebia group but divided it into two sections, the section Kochiae containing those species with scale-likc appendages and the section Vl'illeineliae those with spiny appendages. He thus reversed Brown's application of these names. This nomenclature was also followed by Blutf cl al. (1836 -38). Moquin-Tandon (1834a) adopted Robert Brown's suggestion that these two groups of species should be placed in separate genera but again reversed Brown's application of the names to accord better with the intentions of their original authors (he spell the second name as ' Villeinciia'). The name Bassia Allioni (1766) was not taken up by early or mid 19th century botanists because of the then current use of its homonym Bassia Koenig ex Linnaeus (1771). a genus in the Sapolaceae. For this reason Moquin (1834a) believed it necessary to adopt the later name U'ilicnu’lia. However, shortly after the paper by Moquin it was pointed out by Soyer-Willcmet ( 1834) that the name W'illcmciia Maerklin was a later homonym of that of Necker (1790). Moquin (1834a) therefore, in a footnote to the article by Soyer- Willcmet, published the name Eciiinopsilon as a nomcn novum for WiUemetia Maerklin. Necker's name can be regarded as a unitary designation of a species and is therefore to be rejected from the point of view of priority (Intern. Code Bot. Nomenclature (1972) article 20), but there is also the name ll'il/cmcia Cothenius (1790). currently considered to be a synonym of Kocircuicria Laxman ( 1 772). If Willciiieria Maerkl. is regarded as a homonym of Willemcia Coth. the former would be illegitimate and the name Ecliinopsiloii would then be a legitimate nom. nov. The application of the name Willeniciia Maerkl. depends on its Icclo- typification and on the inlcrprelalion of the International Code of Botanical Nomenclature. As noted above, two species were cited by Maerklin, both of which (as to the names used) are now considered to belong to the genus Kochia. The description given by Maerklin of the second species, •|f'. laiiata'. obviously applies to a species of Bassia All. and it was the species covered by this description (but which did not apply to the basic name cited) that was selected by Moquin (18.34) as the leclolypc of WiUcmcHu (by excluding the other species). Article 10 of the I.C.B.N. slates that "The type of a name of a genus ... is a species". It is not clear whether it is intended that the type should be that species referred to by the name, or that intended by the des- cription (when there is one). If the former, then the names IVil/cniclia and Ecliinopsiloii become congeneric with Kochia: if the latter, then by Moquin's lectotypilication they become congeneric with Bassia All. (For a discussion on a similar nomenclatural question see Regnum Vegetabilc 81 ; 14 15. 98-100 (1972).) When considered to be distinct genera the names Ecliinopsiloii and Kochia were used by most 19th century bolanists in the sense adopted by Moquin; possibly two of the last authors to do so were M.M. ll'in in flora U.S.S.R. 6:124 (1936) and Abrams, lllust. FI. Pacif. States 2:90 (1944). Bentham (1870 and 1880) preferred to consider Ecliinopsiloii (and therefore also Bassia All.) as a .section of Chenolea Thunb. (1781) but he retained the genus Kochia as distinct. However, most writers of European tloras have not (until recently) accepted the suggestion that Bassia (or Ecliinopsiloii) and Kochia arc generically distinct and have usually used the name Kochia to include all the species in this complex. Examples arc Boissier. Elora Orienlalis (1879); Wagner, lllust. Deutsche Flora (1882): Post, Flora Syria. Palestine, and Sinai (1896); Bonnier, Flore Complete France, Suisse et Belgique (1927). The reason these 3 writers adopted the name Kochia was, as is pointed out above, the confusion as to the legitimacy of the earlier name Bassia All. It appears that F. Mueller (1882a and 1882b) was the first to recognise the priority of the latter name over Bassia Koenig. He was followed by Baillon (1887), O. Kuntze (1891), Volkens (1893). and by many later botani.sts. who have, however, circumscribed the genus in various ways. Hermann (1956), for instance, made Kochia a section of Bassia while Allen (1961) retained it as a separate genus. The Australian species which are customarily referred to the genus Kochia have generally had an uneventful nomenclatural history. Robert Brown (1810) described the first two Australian species (K. hrevifolia and K. aphyUa), and suggested that they could belong to the genus WiUemetia. These were transferred to Salsola by Sprengel (1825) (who did not recognise the genus Kochia as being distinct), but. except for one species in Maireana, they, and most of the other species which have been described later, have otherwise been accepted as belonging to the genus Kochia (apart from the re-arrangement of a few species with controversial affinities). The genus Maireana Moquin (1840) was based on the species M. tomentosa Moq. The generic name was accepted by Mueller (1859) who established in it the section Asterocarpiis with the one species M. stelligera F. Muell. This species was subsequently (1869) transferred by him to Echinopsilon while in the same paper he made Maireana a section of Kochia and included in it those species of the latter genus with a simple wing. Volkens (1893) made Maireana a section of Bassia and included in it M. stelligera, in which decision he was obviously following Mueller in his paper of 1859. Thus apart from the one species described by Mueller, which was based on a plant soon removed to another genus, the name Maireana has not been accepted at the generic level subsequent to the two publications by Moquin. The Australian species which are currently referred by Australian authors to the genus Bassia have been placed in several different genera by different workers. Robert Brown (1810), who described the first Australian species, placed them in his new genera Anisacantha and Sclerolaena, and these were treated by botanists as being distinct until F. Mueller (1882) synonymized them both under Bassia. Volkens (1893) also took this position and under Bassia created the sect. Anisacantha in which he placed Sclerolaena in syn- onymy. Domin (1921) and Ulbrich (1934) retained the genus Sclerolaena and placed under it Anisacantha. Bentham (1870 and 1880^ who recognised the genera Anisacantha and Sclerolaena, included under Chenolea four species which w'ere later placed by Anderson (1923) and Ising (1964) in Bassia. Six other genera based on Australian types were also included by Ising in the genus Bassia. These genera are Kentropsis Moq. (1840), Sclero- chlaniys F. Muell. (1858), Dissocarpus F. Muell. (1858), Coilocarpns Domin (1921), Austrohassia Ulbrich (1934) and Sclerohassia Ulbrich (1934). None of those that were published before 1880 had been accepted by Bentham (who included Sclerochlamys under Kochia, and Dissocarpus and Kentropsis under Sclerolaena), or by Volkens (1893). All, except Sclerochlamys, were recognised as distinct genera by Ulbrich (1934). Australian botanists have without exception followed Mueller (1882a, b) and have synonymized the segregate genera (including those described since 1882) under Bas.sia, while ^Aellen (1961), a Swiss botanist who had a comprehensive knowledge of Eurasian Chenopodiaceae, kept up the genera Sclerolaena and Austrohassia. The Eurasian-Australian Generic Disjunction The genera of those members of the subtribe Kochiinae whose type species are from Australia are all endemic to that country. These taxa exhibit in their fruiting perianths a multiplicity of form which causes considerable difficulty in delimitation. The genera may, however, be segregated on the basis of several 4 characters from the “Eurasian” members of the subtribe (which also iiicludes a few African and North American species), and in particular from the genera Kochia and Bcissia. In Bassia and Kochia the species are annual (with one exception) and have herbaceous stems. In Maireana and Scierolaena the species are perennials, sometimes shrubby, and have woody stems. In Bassia and Kochia the flowers are usually aggregated into condensed axillary cymes and may or may not be subtended by secondary bracts. In Maireana and Scierolaena the flowers are axillary and solitary or paired. In Bassia and Kochia the perianth remains chartaceous and weak, and is not modified opposite the radicle. The spines or wings when formed are on the back of the perianth lobes. The embryo is always horizontal and the radicle is variously positioned. In Maireana and Scierolaena the fruiting perianth is variously enlarged and thickened and is modified (by a slit, rib, canal, tubercle or spines) opposite the radicle which is always in the same intertepaline position. This character I have referred to as the radicular anomaly. The spines in Scierolaena are intertepaline in position while the embryo is erect to horizontal. (Figs. lA-D, 2E) The combinations of distinguishing characters listed above appear to be sufficient to consider the Australian species which have at various times been included in the genera Bassia (or Echinopsilon), Kochia, and Chenolea, to be generically distinct from these and to constitute, as does the “Eurasian” group, a complex of closely related taxa without sharp intergeneric delimitations. Generic Delimitation within the Suhtrihc Kochiinae The genera under discussion were placed by Bentham (1880) m the tribe Chenoleae and by Ulbrich (1934) in the tribe Camphorosmeac subtribe Kochi- inae. Ulbrich included 15 genera within this subtribe of which two (Londesia and Chenolea) are not recognised by recent authors as being represented in Australia and need not be further discussed. Of the other 13. four have generally been considered as representing distinct genera. These are Enchylaena, Threlkeldia, Didynumthus and Babhagia, while the remaining 9. as regards recognition or delimitation, have been variously treated by different workers; these are Bassia, Malacocera, Atistrohassia, Scierolaena, Dissocarpus, Kochia, Durialu, Coilocarpus, and Sclerohassia. To this list should be added two genera which have been described since 1934; they are Roycea C. A. Gardn. (1948) and Cyrilwhitea Ising (1964). The characters used by various authors for distinguishing the Australian genera have not been satisfactory, and in fact the multiplicity of form shown in the fruiting perianths of the Australian species usually included in the genera Kochia and Bassia has not made it possible to use key characters which provide a completely natural separation between these and the other genera within the subtribe. Furthermore, characters which appear not to be homologous have been used to classify as being congeneric, species with obviously different affinities. To appreciate the position it is necessary to understand the basic mor- phology of the flower in the Australian representatives of these genera. The perianth is normally 5-merous (occasionally 3-4-merous) and consists of a saucer- to cup-shaped basal portion and a (3) 5-lobed upper portion. The perianth bears two series of veins, 5 tepaline and 5 intertepaline in position (opposite the radicle there may be a pair of inter-tepaline veins). As the fruit develops the basal portion of the perianth usually enlarges and becomes variously thickened; outgrowths often develop from one or other series of veins, or just below the lobes a wing-like structure may form which encircles the perianth. The embryo may be horizontal and almost circular (to horse- shoe shaped), or the radicle may be erect, or the whole embryo slanting to erect. 5 Whatever form the embryo takes it is always positioned in such a manner that the radicle is opposite a particular point of the perianth; at this point the perianth normally develops a slit, bulge, ridge, canal, or one or a pair of tuber- cles or spines (the radicular anomaly). These various radicular modilications of the perianth are associated with the emergence of the radicle at germination; the greater the thickness of the perianth, the more definite the modification. In a few species where the radicle is erect and the perianth open at the summit, there is no prominent radicular modification since the radicle emerges through the open apex of the perianth. It is evident that the characters used to delimit the genera which are here recognised do not always create a natural grouping of species. If an attempt were made to do this it would be found that several small ‘genera of convenience’ would be absorbed into others, with an appreciable loss of value to workers. The genus Bahhagia (1858). for instance, is at present considered to contain four species which are obviously closely related and apparently linked by variants intermediate in form. The perianth structure and plant habit in these species is very similar to that found in Tbrclkeklia salsiiginosa — the type of Osleocarpuin (1858)— which lacks, however, the vertical perianth wings of the Bahhagia species. Threlkehlia salsuginasa is itself very similar to Bassia urceolata (which has only small intertepaline tubercles) and to B. aniscanlhoides — the type of Coilocarpiis (1921). A study of herbarium material suggests that these species are either connected by a series of intermediate forms, or else that hybridization occurs between them in the field. In any case they are obviously closely related. In addition, through the two species of Bassia mentioned, the group shows strong connections to the monotypic genus Cyrihvhilea (1964). Intergeneric relationships may also be recognised in the following groups, although hybridization does not necessarily occur between their members. (1) Threlkeldia proceriflora- Bassia astrocarpa — Dissocarpus spp. (2) Enchylaena tomentosa-Maireana georgei, and Enchylaena tomentosa- Maireana turbinala. (3) Maireana amoena-M. luehmanii-Bassia symoniana -Kochia fimbriolata (Bassia longifolia W. V. Fitzg.). (4) Maireana oppositifolia- Roycea spp. The intergeneric relationship involved in the above cases are discussed more fully elsewhere in the text. Where a species is obviously misplaced its transfer to another genus is warranted, but a mass rearrangement of species in an attempt to provide a more natural grouping of taxa appears at this stage to be unwise, especially as the characters then required to circumscribe the genera w'ould be very difficult to delineate. Key to Australian Genera of the siibtribe Kochiinae 1. Fniiling periamh succulent. 2. Fruiting periantli sub-globular with a marked vertical radicular slit , , Ench.vlaciia 2. Fruiting perianth without a vertical slit Threlkeldia I. Fruiting perianth not succulent. 3. Fruiting perianth with wing-like outgrowths, or surrounded by a single horizontal wing. 4. Flowers in pairs and tused by their bases; embrvo erect; perianth with S horizontal wings; leaves opposite ' Didymanthus 4. Flowers solitary or II In pairs then not fused by their bases or without wing-like growths. 5. Wings 1-5 intertepaline and vertical, situated on top of perianth .... Bahhagia 5. Wing (or wings) horizontal (or both horizontal and vertical) Maireana 3. Fruiting perianth without wing-like outgrowths. 6. Perianth urceolatc, regularly, deeply lobed, not enclosing utricle and without any outgrowths 6 6. Perianth enlarging in fruit and enclosing utricle, usually bearing outgrowths. 7. Flowers in axillary clusters of 2-16 rigidly fused together in fruit Dissocarpus 7, Flowers solitary or if in pairs then not fused together in fruit. 8. Fruiting perianth without outgrowths — sec Threlkeldia and Maireana. 8. Fruiting perianth with spines. 9. Fruiting perianth with spine-like outgrowths which alternate with perianth lobes (sometimes with additional accessory lobes) or with ca. 10 short radiating teeth Sclerolaena 9. Fruiting perianth with 3-5 outgrowths opposite the perianth lobes. 10. Embryo and outgrowths horizontal. II. Outgrowths solid and hard ... Maireana 11. Outgrowths cylindrical, soft Malacoccra 10. Embryo vertical, spines erect or somewhat spreading at apex of cylindrical perianth Threlkeldia proceriflora and Bassia astrocarpa Babbagia F. Muell. (1858). Type: B. diptcrocarpa F. Muell. Fruiting perianth .subglobular, crustaceous or woody, S-merous with 2-5 erect wings at apex which are intertepaline in position. Seed horizontal with an ascending radicle. Four species are known in this genus. Bahhagia is closely related to Sclerolaena (from which it differs principally in the inter-tepaline processes being wing-like and not spiny) and to Threlkeldia sahuginosa, q.v.. Collections have been made of plants which are presumably hybrids between Bahhagia &p. and Threlkeldia .salsiigino.sa, and between Bahhagia and Sclerolaena sp. (Bassia iirceolata Ising). Cyrihvhitca Ising (1964) Type: C. walkeri (White) Ising In this monolypic genus the fruiting perianth has five main spines which are erect and intertepaline in position with five much shorter spines between them. The lube has 5 narrow vertical wings (also inter-tepaline) with 5 promin- ent ribs betw'cen them. The lower half of the 'tube' consists of a 5-chambered base. The centrifugal radicle emerges through a canal in line with oneof the main spines. The perianth structure is thus basically similar to that found in Bassia hrachyptera and B. niicrocarpa. for the chambered hollow base is only a more prominent example of a character frequently found in Australian 'Bassia' species (including B. niicrocarpa). There appears to be no good reason for segregating this species in a genus distinct from Scleroleana. Didymanthus Endi. (1839) Type: D. roei Endl. Flowers in pairs. Fruiting perianths cylindrical, fused by their bases and strongly divaricate: apex of perianth with live horizontal chartaceous wings which are tepaline in position. Embryo erect. This genus is represented by only one species. It differs from Maireana principally in the position of the embryo and the virtually actinomorphic fruiting perianth which lacks the radicular anomaly. Dissocarpus F. Muell. (1858) Type: D. hijioriis F. Muell. Flowers clustered in leaf axils and fused in condensed dichasia (ca. 2-7 in D. bifiorus and 8-16 in D. parado.xus). Fruiting perianth hardened, 5-lobed; at the base of each lobe a spine or protuberance develops (not always present in D. bifiorus). Seed horizontal; radicle ascending to erect. 7 Two species, each of which exhibits considerable variation. This genus was distinguished by Mueller and recognised by Ulbrich (1934) on the basis of the flowers being united in clu.sters. The union of the flowers does not by itself warrant generic delimitation and for this reason both Anderson (1924) and Ising (1964) placed Dissocarpus in synonymy under Bassia. However, Dissocarpus differs markedly from other Australian "Bassia' species (i.e. from Sclerolaena) in having spines developing from the base of the perianth lobes (and not from between them) and in not having any radicular protuberance or slit. Anderson described spines as being occasionally present in D. hifiorus, but the protuberances that he observed were the remains of sterile flowers forming part of the dichasia at the base of the fruiting perianth. These sterile structures range in size from minute excrescences to miniature but recognisable flowers, one or two of which may be present at the base of the same mature perianth. True spines (i.e. structures homologous to those found in D. paradoxus) are also found in D. biflorus; these arise at the base of the perianth lobes (which they resemble) and grow upwards to overtop and obscure them. A superficial examination may thus lead one to assume that the spines are the perianth lobes. (Fig. 2C-D). Under Bassia sect. Dissocarpus a third species (B. georgei Ising) was included by E. H. Ising (1964). This species has fruits with irregularly lobed wings while the fruiting perianths themselves are united into clusters and appear to be sterile. It is not related to the other two members of the genus Disso- carpus and is quite possibly an intergeneric hybrid involving Maireana and Sclerolaena. For a discussion on the nature of the axillary inflorescence in Dissocarpus see Bisalputra (I960). The two species Bassia astrocarpa and Threlkeldia proceriflora have a similar perianth shape and spine arrangement to that found in Dissocarpus. This may be of phylogenetic significance. Enchylaena R. Br. (1810) Lectotype: E. tomentosa R. Br. (The only other species described by R. Brown was E. paradoxa; this, according to Bentham, is a monstrous state of E. tomen- tosa, and the name is therefore illegitimate.) Perianth pentamerous, succulent in fruit with an inner cartilaginous layer, while towards the apex often a ring-like succulent outgrowth grows upwards and ovei the perianth lobes (corresponding to the wing of Maireana); opposite the radicle a vertical slit is present, the margins of which overlap. ’ Embryo horizontal. (Fig. lE-F). ^ This genus is represented by several species only one of which has been described. Other species which have been referred to the genus Enchvlaena are either synonyms of E. tomentosa or correctly belong to other genera. Hybridisation between E. tomentosa and Mairaena georgei has been observed in the field at several localities. Maireana Moq. (1840) Type: M. tomentosa Moq. Flowers 5-merous. Fruiting perianth patelliform to hemispherical with a d: horizontal wing-like outgrowth developing from beneath the perianth lobes or a separate process developing from beneath each lobe; opposite the radicle a sht, bulge or canal is present in the perianth tube. Seed + horizontal- the radicle centrifugal or enclosed. ’ The genus consists of 58 species. Malacocera Anders. (1926) Type: M. tricornis (Benth.) Anders. Fruiting perianth globose, 5-Iobed, not hardened, with a prominent vertical slit opposite the radicle; from below the perianth lobes arise three soft cylindrical horizontal processes. Seed horizontal; radicle centrifugal. The genus consists of two species. Malacocera differs from Sclerolaena in having a horizontal embryo and in the tepaline position of the processes. From the genus Bassia s.slr. it differs in having a prominent radicular slit in the perianth, and from Maircana in the shape and texture ot the processes. In the reduced number of processes compared to the perianth lobes, and in the shape and texture of the perianth, it corresponds closely to some Eurasian species of Bassia, but no close relation- ship is suggested. Roycea C. A. Gardn. (1948) Lectotype: R. pycHoplivlloides C. A. Gardn. Perianth deeply 5-lobed, without any radicular anomaly and scarcely enlarged in fruit. Utricle ovoid when ripe and considerably overtopping the perianth; pericarp crustaceous. Embryo vertical, circular. (Fig. IG-H). Three species (one undcseribed). The fruit of the two species when first described was unknown and the characters used by C. A. Gardner to distinguish the genus were not substantial since they could equally have applied to several species of Maireana. Fruiting material of all three species has recently been collected and it can be shown that the characters apparent therein clearly delimit Roycea from related genera. The two species originally described {R. pycnophylloides and R. spinescens) are predominantly dioecious, but a third species (undescribed) is hermaphro- dite. The leaves, indumentum, and flowers of the three species in this genus correspond closely to that found in Maireana opposilifolia. The fruiting perianth is, however, quite different. Sclerolaena R. Br. (1810) Lectotype: S. nniflora R. Br. (fide Ulbrich, 1934). Fruiting perianth usually globular to urceolate, 3-4-5-merous, variously hardened, normally bearing 2-6 spines at apex of perianth-tube alternate to perianth lobes (when 4 to 6 spines then usually with a pair opposite the radicle). Seed ascending to erect. 80-90 species. As delimited here the genus Sclerolaena includes the genera Anisacantha, Coilocarpiis, Auslrohassia, Sclerochlaniys, Sclerohassia Cyrilwhirea, and Kentro- psis. It differs from Bassia s.str. in usually having a woody fruiting perianth, in the seed position, in the spines being intertepaline in position, and in the radicular canal of the perianth. The plants are always woody perennials (annuals in Bassia). The arrangement of the spines is fairly constant and is related to the position of the intertepaline veins of the perianth tube. Opposite the radicle the intertepaline rib consists of a contiguous pair, of which either or both members may give ri.se to a spine. Thus in species such as B. euroiioides, B. parvifiora, and B. microcarpa (which have 5-merou.s flowers) there are six spines, two of which arise close together opposite the radicle. In B. densiflora (which has 4-merous flowers) there are 5 spines, two of which ari.se opposite the radicle. In many of the species of Sclerolaena one or more of the inter- tepaline veins does not develop into a spine, or one or both of the pair of radicular veins forms only a small tubercle. Thus in B. eriacantha there are 9 Iwo spines of which one is next to the radicle and is associated with a tubercle which has developed from the other of the pair of veins. In B. crenata there are usually two tubercles opposite the radicle. At the position of the pair of radicular spines or tubercles the division between the adjacent tepals develops more deeply than between the other tepals (or perianth lobes) and this division passes between the spines as a radicular slit, through which in many cases the radicle eventually emerges. In Bassia astrocarpa the 5 spines are tepaline in position, and in flower structure and habit this species is very similar to Threlkeldia proceriflora q.v. Thrdkeldia R. Br. (1810) Type species: T. diffusa R. Br. Perianth subglobular to urceolate, 3 or 5-merous, fleshy in fruit with a hard inner layer which on drying may form several apical knobs. Embryo horizontal to erect with an ascending radicle. Represented by 2 4 species depending on the circumscription of the genus. T. diffusa has a 3-merous perianth and on drying the woody inner perianth develops inter-tepaline knobs (of which a contiguous pair are opposite the radicle). This condition is comparable to the situation found in the majority of Sclerolaena species. In T. inchoata the flower is 5-merous and the fruiting perianth is raised opposite the radicle. In T. proceriflora a shallow cup-shaped outgrowth forms at the apex of the perianth which gives rise to 5 teeth opposite the perianth lobes. The perianth appears to be dry (not succulent). This then is comparable to the situation in Bassia astrocarpa. It is obvious from general morphological characters that T. proceriflora and B. astrocarpa are closely related and possibly should both be segregated as a distinct genus. T. salgusiiiosa is very similar in both vegetative and fruit characters to the species in the genus Babhagia. The fruiting perianth normally has a rounded protuberance at the position of the radicle and this protuberance is in some forms enlarged and slightly bifld while 4 small additional erect intersepaline processes may also develop. The perianth in this state is then little different from that in some species of Sclerolaena (e.g. Bassia urceolata), or that in Babhagia scleroptera in which the protuberances extend into erect wing-like structures. Morphological Notes on Maircana INFLORESCENCE The flowers are normally solitary and axillary to a leaf Sometimes they are clustered into terminal spikes and are then subtended by modified leaves or bracts. In a few species the flowers bear a pair of small bracteoles (e.g. in M. diffu.>ia, M. marginata, M. suaedifolia, and M. enchylaenoides) but most are ebracteolate. A number of species have their flowers consistently in pairs, e.g. M. amoena, M. luelimannii and M. scleroptera.. and in these the flowers of each pair are equal in size and normally both develop to the fruitina stage. In M. plani folia and M. .sedifolia the flowers are also usually in pairs but normally only one of each pair matures. According to Bisalputra (1960) the .solitary axillary flower in Sclerolaena represents a dichasium in which the lateral buds have been suppressed. If this is correct then the solitary flower of Maireana has probably a similar origin and it may be assumed that paired flowers repre- sent a dichasium in which the terminal bud has been suppressed. (See also notes under Orientation of Flower and Fruit.) In most species the flowers are basically hermaphrodite (although stamens may not always mature), but others are predominantly or strictly dioecious. The plants for example of M. oppositifolia, M. atkinsiana, and M. sedifolia 10 are normally strictly dioecious while M. aphylla and M. prosthecochaeta are polygamodioecioLis. It is interesting to note that the male flowers in those species which are strictly dioecious also possess a radicular slit to the perianth in the same position as it is found in the female Hower. PERIANTH The perianth consists of a united portion (the tube) on which are borne 5 lobes, or the lobes may themselves be partially united above the apex of the tube proper; this united portion of the lobes is distinct both in texture and in indumentum from the tube. The portion of the perianth above the tube (and therefore above the wing in the fruiting perianth) is here referred to as the upper perianth; this corresponds to the 'limb' in Scifrolaeiui. In the position opposite the future radicle the lobes are usually distinct down to the tube, or this division between the two lobes (on either side of the future radicle) may continue for some distance into the tube. This division is here referred to as the radicular slit. The margins of the slit may overlap or may be raised or thickened. THE WING A horizontal wing de\'elops as an outgrowth at the junction of the tube with the upper perianth, or when the perianth lobes arc distinct down to the tube a separate w'ing may form opposite each lobe. In most cases the wing arises from below the level of the united perianth lobes and it is therefore single. Normally (as mentioned above) the radicular slit continues a short distance down the perianth tube and this causes the wing at that point to have a radial (radicular) slit. In a few species (e.g. M. toiuenlosa and N. iiiiegra) the radicu- lar slit does not continue into the perianth tube and the wing in these cases is entire, while in a very few' species (e.g. M. atkinsiana) the presence of the radial slit is variable in perianths on the same plant. ACCESSORY LOBES OR WINGS Some Maircana species bear, as well as the normal wing (or wings) between the tube and the upper perianth, additional structures which may be wing-like or spinose in appearance. Vertical wings are borne on the perianth tube in Af. crioclatla. M. pcniciptcni. and M. tripiera. while in A/, polyptcrygia an additional horizontal wing forms at the base of the tube. The vertical wings, whether on the tube or above the wing (as in A/, dichoptcni). are always alternate to the perianth lobes, and when opposite the radicle often arise as a contiguous pair (compare the contigtious pair of spines opposite the radicle in many species of Sclerohicna). In a number of species accessory processes are pre.sent on the upper perianth (i.e. above the wing). These lobes are sometimes intertepaline in position (as in M. lanosa, M. lohiflora, and A/, prosthecochcicta) and then presumably correspond to the spines found in Sclcrolaeiui: or they may be tepaline in position (;is in M. scicrolueiioicles). In A/, gloiucrifolia and M. alkinsiana there are live processes, a pair of which is present on each of two perianth lobes with a single process on a third lobe while the other two perianth lobes are without processes. In A/. nwUuwconui numerotis trichome-like processes form and these are not associated with the perianth venation. The nature of these trichomes is quite different from that of the processes otherwise found in this genus or in Sclerolueiui and are probably in the nature of emergences. RADICULAR CANAL When the wall of the fruiting perianth tube is relatively thin, then at that position opposite the radicle it may be marked by a slit or weak point which readily breaks open. In those species where the wall is woody or otherwise indurated there is a ‘tunnel’ leading to the outside of the perianth. This 11 'tunnel' is here referred to as the radicular canal. It is not strictly a tunnel for, although enclosed, along its upper side runs the radicular slit of the upper perianth. The outer end of the radicular canal may be open, covered with indumentum, or closed over by a membrane. STYLE The style is normally a weak structure included within the perianth and divided into two or three stigmatic branches. In M . platycarpa it is very firm and prominently exserted, while in M. georgei, M. turhinala and M. convexa the style is broadly cone-shaped and forms a hard areolate cap to the utricle. EMBRYO The embryo is more or less circular with the radicle horizontal and either contained within the circle or slightly exserted (centrifugal). The cotyledons are normally incumbent and narrow but in those species with flattened fruits they are broad and twist to present their edges to the radicle (accumbent). Examples of the latter condition are found in the seeds of A/, platycarpa, M. glomerifoUa and M . atkinsiana. Orientation of Flower and Fruit As has been mentioned previously, the flowers and fruits of the majority of the species belonging to the Australian genera of the subtribe Kochiinae are asymmetrical. They are arranged on the branch in a regular manner according to the phyllotaxy of that branch. The asymmetry of the flower arises basically from the position of the embryo and the eventual emergence of the radicle. Attendant on this emergence is the radicular slit in the perianth tube and overlap of its margins; in addition there is associated the position of the spines (in Sclerolaena) or accessory processes (in several species of Maireana). When the phyllotaxy is an ascending clockwise spiral (here referred to as dextrorse) the radicular slit is on the right hand side (from the point of view of the observer), and when anticlockwise (sinistrorse) on the left-hand side. Normally at each successive branching the direction of phyllo- taxy changes. The radicle position is constant in relation to the subtending leaf or bract. Thus in a 4- or 5-merous flower the radicle emerges between the second and third perianth lobes away from the bract, either on the left or right-hand side depending on the phyllotaxy. The radicle lies on the abaxial (anterior) side of the fruit. In a dextrorse flower it is directed to the right and in a sinistrorse to the left. In Maireana the subtending bract is excentric (placed to the left of the stem axis in a dextrorse phyllotaxy and to the right in a sinistrorse) which causes the radicular slit to occupy an apparent lower left or lower right-hand position. In Sclerolaena the bract is normally centrally placed and the radicu- lar slit occupies an upper left or upper right-hand position. It follows from the above that there are in each species of Maireana, Sclerolaena. and Enchylaena, etc., two flower and fruit forms, and that these forms are mirror images of each other. This is most obvious in those species with a markedly asymmetric fruiting perianth. (Figs. lA-D.) In the groups of species with consistently paired flowers (e.g. the M. amoena and M. atkinsiana groups) the flowers in each pair are identical, i.e they are not mirror images of each other, and it depends on the phyllotaxy as to which of the two forms is present. This is also the case where, as in M. planifolia, either one or two flowers are formed in the axis of the’ bract, but when two, they are of difiTerent sizes and usually only one of them develops to the fruiting stage. The presence of only one type of flower at a node supports the suggestion made previously, that paired flowers represent a reduced dichas- 12 ium in which the terminal flower has failed to form, for if the two flowers consisted of one terminal and one lateral to a reduced raceme, then it could be expected that the one would be sinistrorse and the other dextrorse in morphology. Infraspecific Variation and Hybridization Most species of Maireana, even if widely distributed, are relatively constant in their morphology. In M. georgei, however, there is considerable regional variation both in fruit and foliage characters, while M. oppositifolia exhibits practically no variability in its fruit but a considerable amount in its habit and foliage, especially between the form lound in inland Western Australia and that found along the south coast. When, as in the latter case, the vari- ability is confined to the habit and foliage it is not possible without transplant experiments to distinguish between phenotypic and genotypic variation. In M. amoena the range of variation both in leaf and fruit characters is very great and some forms come close to M. scleroptera with which it may once have had a clinal relationship. The two species are now separated geographically by a large area of the eremean desert in which neither plant has been collected. Interspecific hybridization within the genus Maireana has not been con- firmed in the field" although it is probable that it does occur, for instance, between M. villosci and AT planifolia. These two species are sympatric over much of their range (although they differ in their ecological preferences), and frequently a plant intermediate in morphology between them is lound. This intermedi'ate form is also present in areas where M. planifolia has not been recorded (such as western New South Wales). More field work is required before any clear idea can be formed as to the actual situation involved. Hybridization has been observed at several localities in Western Australia between M. georgei and Enchylacna tonientosa. The hybrid is intermediate between the parents both in leaf and fruit characters although the fruit falls readily even when immature. It is probable that hybridization also takes place between M. turhinata and E. tomentosa since plants intermediate in form between these species have been collected in South Australia and New South Wales in areas where both of the putative parent species are found. It may be noted that the fruiting perianth of E. frequently develops an outgrowth corresponding to the wing of M. georgei but in the former species this ‘wing' is succulent and is incurved over the perianth lobes. Relationships with the genus Sclerolaena The characters which distinguish Maireana from other genera are some- times difficult to determine since species which exhibit intermediate characters do occur. Among the Australian species currently included within the genus Bassia there are two which have sometimes been placed in Kocliia. These are B. slelligera (F. Muell.) F. Muell. and B. hrachyptera (F. Muell.) Anders. They have a woody perianth with usually 5 or 10 short horizontal spines which may be united at their base into a short horizontal wing. The seed is horizontal and the embryo circular. Apart from the scarcely developed wing the fruiting perianths are very similar to those found in some species o\' Maireana. The closest affinity to these two species appears, however, to lie with some 'Bassia' species, e.g. B. niierocarpa Anders, and B. cosiata Anders, which are typical members of the genus Sclerolaena. A further approach to Maireana is found in the species B. symoniana Ising. This species, with B. lueinnannii F. Muell., was placed by Ising (1964) in his sect. Spinosissimae, a section characterised by having its flowers in pairs. In B. luehinannii \ht perianth is woody with 5 flattened horizontal spines oppo- site the perianth lobes. In B. symoniana the perianth is again woody but there are two series of appendages; in one series are five erect ‘spines' which 13 arise opposite the perianth lobes, alternating with 4 -i (2) horizontal 'spines’ which are inter-tepaline in position, the odd pair being placed at the radicular slit. This arrangement of the inter-tepaline spines is identical to that found in most Australian 'Basshi' species (Sclcrolacna). svhile the erect spines pre- sumably correspond to the horizontal spines found in B. hielunaniiii and to the wings found in some Maireaiui species. Almost identical in fruit mor- phology to B. synwniami is Kochici Jimlvinlciia F. Muell. (in which the flowers are also in pairs). Closely related to B. lueliiuaniiii are M. anioena and M. scleropicrcu which in each case have paired flowers and distinct Icpalinc wings to the fruit. Although this group of 5 species has much in common it has been decided to place K. fimhriolata and B. synwniuna in the genus Sclerolaena (in which genus there arc some species with similar perianths) and the others in the genus Maircana. The similarity (and apparent homology) between the spines found in Sclerolaena and the accessory lobes in Maireana lanosa and M. lohiflora, is discussed elsewhere. Distribufion The genus Maircana is endemic to Australia (excluding Tasmania) w'here it is typical of the eremean areas, w'ith a few species extending into the agri- cultural lands. In the latter case the plants are usually salt tolerant species and generally occur in coastal situations or inland in slightly saline soils. Several of the eremean species are also salt or gypsum tolerant and in fact are only found in association with inland ‘salt’ lakes. Other eremean species appear to be restricted to well-drained situations (e.g. M. planifalia. M. inelanocoma, and M. nnirrayana) and a few appear to favoLir non-saline clay soils (e.g. M. aphytla). A number of the species, particularly those that are succulent and almost glabrous, are heavily grazed by slock and for this reason are infrequent in agricultural or pastoral areas. The species M . suacdifoUa and M. thesioides, for instance, are usually only found growing in the protection of a dense bush and their natural growth form has not been observed. MAIREANA Maireana Moqiiin-Tandon, Clienop. Enum. 95 (1 840), et in Ann. Sci. Nat. Ser. 2. 15:97. t.i3 (1841). Type: M. lonicnlosa Moq. Kochiii ^ect. Muircentu (Moq.) F. Muell., Fragm. 7:12 (1869). Bassiu sect. Maircana (Moq.) Volkens in Engler et PrantI, Nat. Plfanzenfam. III. la: 70 (1893), not as to description. Enchylaena sect. Hcierochlamys F. Muell., Trans, Phil. Inst. Viet. 2:76 (1858). Type. E. vilioia F. Muell. Kocliia seel. Diiiiala Anders., Proc. Linn. Soc. N.S. Wales 51 :383 (1926) iiom. illeg. (not validly published and a supernuous name). Type: Encitylueiui viHosa F. Muell. Duriaiu IJIbrich in Engler et PrantI, Nat. Pflanzenfam. ed. 2. 16c:537 (1934), based on above. Bassia seel. ErwchiUm Anders., Proc. Linn. Soc. N.S. Wales 48:320 (1923) Type: Echino- psiion scicrolaenoulcs F. Muell. Kochia sect. Aiisirokocliia Ulbricb in Engler et PrantI, Nat. Plfanzenfam, ed. 2. 16c:535 (1934), without citation of type. Bassia sect. Spinosissimae Ising, Trans, Rov. Soc. S. Austral. 88:75 (1964). Type- B liiehmaimii F. Muell. [Kochia auct. plur. non Roth : e.g. R.Br., Prod. 409 (1810); Bentham, FI. Austral. 5:183 (1870): J. M. Black, FI. S. Austral, ed. 2. 309 (1948); J. H, Willis, Handb PI Victoria 2:102 (1972).] Herbaceous to woody perennials or small shrubs, glabrous or with simple (often woolly) or branched hairs. Leaves alternate or opposite, globular to terete or narrowly-oblong. frequently fleshy or succulent. Flowers axillary, solitary or in pairs, sessile, rarely minutely bibracteolate, hermaphrodite’ dioecious or polygamodioecious; perianth 5-lobed, with a flat to cup-shaped 14 tube, the lobes usually more deeply separated in the position opposite the future radicle; stamens 5, opposite the perianth lobes; ovary sub-globular, style short or long with 2-3 linear stigmas, ovule solitary. Fruiting perianth spongy, leathery, cruslaeeous or woody; tube patelliform to globular, sometimes with membranous extensions (accessory wings); wing arising at the base of perianth lobes, usually horizontal, simple and conlinuous (or with a radial slit opposite the radicle) or divided into 5 separate wings (rarely spines) each opposite a perianth lobe, chartaceous or rarely woody; perianth lobes usually horizontal and obscuring utricle, rarely erect, occasionally giving rise to erect appendages either tepalinc or intertepaline in position, or to emergeneies. Utricle discoid, turbinate or globular; pericarp membranous all over, or crusta- ceous above, or entirely crustaceous; style weak or firm, sometimes massive and hemispherical to cone-shaped; seed horizontal; endospermous; embryo circular or horse-shoe shaped, the radicle enclosed or centrifugal. 58 species endemic to mainland Australia. Named after the French naturalist Mairc. Material has been studied from the major herbaria in Australia but for the sake of economy of time and of publication space only a small selection of this has been cited. The maps record the provenance of all specimens seen except where the localities could not be traced. Key to .Species Note 1. This key is for specimens with fruit and, although several species are dioecious, it is not practicable to prepare a separate key to male plants, or to non-fruiting female or hermaphrodite plants. 2. The term ‘fruit’ as used in the key refers to the fruiting perianth. 3. The wing (or wings) of the fruiting perianth may sometimes fail to develop. This apterous condition is particularly common in M. microphylla; it is also found in M. hrevifoUa. 1. Fruit with 5 • horizontal wings surrounding the upper perianth. 2. Fruit with erect processes arising from upper perianth. 3. Wings Hat, ± truncate or rounded at apex 11. M. lobiflora 3. Wings spine-like (fruit woolly) 1. M. sclcrolaenoides 2. Fruit without erect processes. 4. Leaves slender, glabrous, spurred at base; wings pubescent with curled hairs 3. M. chcelii 4. Leaves various; wings hairy or glabrous. 5. Leaves sericeous: fruit densely silky lanate, ca. 10 mm diameter 2. M. eriantha 5. Leaves and llowers not as above. 6. Flowers in pairs (tube of fruit i hat or very shallow). 7. Wings ■ glabrous. 8. Wings each deeply divided, pungent, woody 4. M. luchmannii 8. Wings neither deeply divided nor pungent 5. M. amoena 7. Wings densely woolly 6. M. selcroptera 6. Flowers solitary (wings glabrous). 9. Wings thin, obovate, to fan-shaped. 10. Wings equal and horizontal; leaves alternate; (lowers hermaphrodite. 1 1 . Leaves slender 2^ (10) mm long, not attenuate at base, often hairy (wing simple if present) 47. M. microphylla 1 1. Leaves attenuate at base, glabrous. 12. Leaves obovoid (to terete), I 5 long: llowers without bracteoles 7. M. brevifolia 12. Leaves on older branches slender-fusiform, to 15 mm long; flowers minutely bracteolate 8. M. diffusa 10. Two of the five wings smaller and sub-erect; leaves often opposite; plants usua ly dioecious 9. M. uppositifulia 9. Wings small, imbricate, lunate, incurved or inflated on margins; slender perennial to 0-3 m high 10. M. encbylacnoides 1. Fruit with a single wing surrounding the upper perianth (accessory wings on tube or upper perianth may also be present). 15 13. Fruit small (under 5 mm diam.), ! densely silky pilose; tube almost fiat and continuous with narrow wing (small perennials). 14. Upper surface of fruit with an erect cup-shaped outgrowth .... 15. M. coronata 14. Fruit without such an outgrowth. 15. Fruit 2-5-4 mm diam., upper surface with a pentagonal shaped ridge J4. M. pentagona 15. Fruit ca. 4 mm diam., upper surface with 5 narrow radial ridges e.xtending from near the centre to the edge of the wing 13. M. ciliata 13. Fruit (including wing) mostly over 5 mm diam. or if smaller not silky pilose. 16. Fruit enveloped in long dense wool. 17. Fruit shortly pedicellate, tube turbinate; (lowers in vertical rows in dense spike 17. M. eriosphaera 17. Fruit sessile, tube broadly hemispherical; flowers spirally arranged in dense spike 18. M. carnosa 16. Fruit not enveloped in long wool. 18. Fruit glabrous with large rounded soft spongy tube; upper perianth ± flat. 19. Branches with a pale fawn indumentum of dendritic hairs; leaves very slender glaucous; horizontal wing not decurrent on tube .. 30. M. campanulata 19. Branches with a white woolly indumentum of simple hairs; leaves fieshy, narrowly terete or fusiform; horizontal wing attached by a short decurrent wing to tube 29. M. spongiocarpa 18. Fruit not with above characters combined. 20. Tube of fruit with an expanded lleshy base; wing a narrow rim 19. M. marginata 20. Tube not expanded at base or the wing obvious. 21. (I) 3-5 vertical wings on tube or above horizontal wing. 22. Fruit small, thin walled, with 5 vertical radial wings above horizontal wing 16. M. dichoptera 22. Fruit thick walled, the vertical wing or wings on tube. 23. Plant glabrous (or almost so) .... .... 32. M. triptera 23. Plant hairy, at least on branches. 24. Fruit with a basal horizontal wing (in addition to normal wing); leaves pubes- cent . . 31. M. polypterygia 24. Fruit without a basal wing; leaves glabrous or pubescent. 25. Leaves villous; perianth lobes erect and longer than tube 35. M. schistocarpa 25. Leaves glabrous; perianth lobes not erect. 26. Vertical wings running length of tube and united at apex with horizontal wing; perianth lobes prominently woolly ciliate ... 33. M. erioclada 26. Vertical wings often present only towards base of tube; perianth lobes arched and prominently tomentose . .34. M. pentatropis 21. No vertical wings on tube. 27. Horizontal wing present at base of tube similar in size and additional to normal wing 31. M. polypterygia 27. No horizontal wing at base of tube (which may however be expanded and hollow). 28. Vertical processes present on upper perianth. 29. Processes of numerous fine needle-like emergences . 37. M. melanocoma 29. Processes 4-6, thick or subulate. 30. Leaves very small and condensed into glomerules along branches 40. M. glomerifolia 30. Leaves well developed. 31. Fruit woolly all over; leaves acute, silky 12. M. lanosa 31. Fruit glabrous or almost so; leaves not silky. 32. Leaves sparsely pubescent, succulent, usuallv rounded at apex (divaricately branched sub-.shrub) 41 . m. atkinsiana 32. Leaves glabrous, narrow, acute : branches erect 43. M. prosthecochaeta 28. No vertical processes above wing. 33. Plant glabrous (apart from axillary tufts of wool); slender lax plants. 34. Upper perianth convex; wing of mature fruit 8-15 mm diam. 35. Leaves narrowly fusiform, basally attached; fruit to 15 mm diam dark coloured when dry 43 . m. thesioides 35. Leaves semi-terete, shortly spurred at base; fruit ca. 8 mm diam , straw- coloured when dry ... 57. M. aphylla 34. Upper perianth i flat; wing 10 mm or less in diameter. 16 36. Leaves slender-fusiform on flowering branches; flowers minutely bracteo- late 44. M. suaedifolia 36. Leaves terete; flowers ebracteolate. 37. Leaves not lleshy; stem sparsely hairy with short straight appressed hairs 47. M. micropliylla 37. Leaves fleshy; stem sparsely hairy with short woolly hairs 45. M.decalvans 33. Plant variously hairy. 38. Leaves opposite; tube patelliform or flat. 34. Leaves narrowed at base; wing (10) 15-23 mm diam. 39. M. platycarpa 39. Leaves sessile by a broad base, apex recurved; wing ca. 6 mm diam. 38. M. cannonii 38. Leaves all or mostly scattered. 40. Style hard and prominently protruding in fruit; tube patelliform or flat 39. M. platycarpa 40. Not as above (usually only stigma arms exerted). 41 . Wing of fruit glabrous. 42. Upper perianth flat or slightly concave or convex. 43. Tube of fruit abruptly narrowed at base into a prominent terete stipe ( I ) 2-3 mm long. 44. Upper perianth convex, open in centre to expose utricle 57. M. apliylla 44. Upper perianth flat or slightly sunken, completely covering utricle. 45. Hairs on leaves dendritic (i.e. with short lateral branches); stipe of fruit papillose, solid 58. M. stipitata 45. Hairs on leaves simple; stipe smooth, hollow .. . 56. M. appressa 43. Tube not as above. 46. Tube expanded into a hollow spongy base (perennial herb) 20. M. excavata 46. Tube not (or scarcely) expanded at base. 47. Wing of fruit with a radial (radicular) slit. 48. Leaves, branches, and flowers closely lomentose with branched (dendritic) hairs; leaves obovoid; tube and convex upper perianth of fruit pubescent ... 27. M. astrotricha 48. Leaves and branches with ± simple hairs or glabrous. 49. Fruit large (wing ca. 15 mm diam. or more); tube thick walled, tur- binate. 50. Upper perianth flat, glabrous except for woolly margin to lobe. 23. M. gcorgei 50. Upper perianth convex, pubescent. 51. Wing to 15 mm diam., leaves linear to narrow-terete, acute, 10-20 mm long ... 24. M. convexa 51. Wing to 25 mm diam.; leaves oblong-obcuneate, very thick, ca. 20 mm long 26. M. niurrayana 49. Fruit with wing 5-14 mm diam.; tube not thick walled. 52. Tube of fruit pubescent; upper perianth ± flat; leaves obovoid, fleshy 28. M. sedifolia 52. Tube of fruit glabrous: leaves various. 53. Leaves flattened, linear to obovale. 54. Leaves linear, appressed villous; upper perianth glabrous; wing with radial anastomosing veins when dry .... 52. M. villosa 54. Leaves narrowly to broadly obovate, pubescent with curled hairs; upper perianth pubescent; wing without obvious nevation 53. M. planifolia 53. Leaves terete, semilerete, or obovoid. 55. Fruit produced into a short terete hollow stipe at base, straw- coloured when dry. 56. Open divaricately branched shrub: branches striate and often spinescent; upper perianth convex and open in centre 57. M. aphylla 56. Branches ± ascending, neither spinescent nor striate; upper perianth flat or sunken and completely covering utricle 56. M. appressa 55. Fruit without a stipe-like base. 57. Upper perianth with a convex disc, open in centre; wing not prominently crenulate. 58. Stem striate; leaves sessile, woolly to glabrescent 57 M. aphylla 17 58. Stem not obviously striate, leaves narrowed at base, glabrous 46. M. rohrlachii 57. Upper perianth t flat, concave, or if convex then closed in in centre. 59. Upper perianth toinentose; wing with fine but obvious radiating nerves 55. M. radiata 59. Fruit glabrous or sparsely villous. 60. Branches J lomentose at least on young parts (rarely glab- rous); flowers not in dense spikes. 61. Leaves toinentose. 62. Fruit straw-coloured when dry and with a short hollow stipe 56. M. appressa 62. Fruit almost black when dry, no stipe pre.sent 54. M. niclanocarpa 61. Leaves glabrous or sparsely villous; fruit brown when dry. 63. Upper perianth closed in centre; tube firm 48. M. microcarpa 63. Upper perianth open in centre (exposing utricle); tube weak. 64. Wing 1 1-16 mm diam., often undulate 46. M. rohrlachii 64. Wing ca. 8 mm diam., flat 45. .M. decalvans 60. Branches sparsely strigose or sparsely villous (or somewhat woolly when very young); leaves slender 2-4 mm long; wing of fruit crenulate, flowers in dense spikes 47. M. mierophylla 47. Wing of fruit continuous (i.e. without a radial slit); leaves semitcrete. 65. Fruit large, wing ca. 15 mm diam.; tube turbinate, smooth 25. M. (urhinata 65. Fruit small, wing II mm or less in diameter; tube hemispherical to cupular. 66. Leaves 2-5 mm long, appressed on younger branches; wing 4-6 mm diam. .... .... .... .... ... .. 49. M. ovata 66. Leaves over 5 mm long, spreading; wing usually 8-10 mm diam. 67. Upper perianth pubescent 51. M. Integra 67. Upper perianth glabrous (or almost so) apart from ciliate lobes 50. M. tomentose 42. Upper perianth columnar in form with large erect lobes. 68. Leaves 2-6 mm long, shortly pubescent; no vertical wings on tube 36. M. pyramidata 68. Leaves 5-12 mm long, appressed villous; tube with a narrow vertical "■og 35. M. schistocarpa 41. Wing of fruit pubescent above. 69. Upper perianth erect and columnar .35. M. schistocarpa 69. Upper perianth not erect. 70. Wing up to 10 mm diam. 71. Perianth tube thin walled, base expanded into a hollow stipe- leaves semi-terete, ca. 10 mm long 21. M. trichoptera 71. Perianth tube with a hard boss-like base; leaves elliptical to narrowly obovate, mostly 7-12 mm long 22. M. huniillima 70. Wing 15-25 mm diam., tube turbinate, base not expanded- leaves very- thick and fleshy ca. 20 mm long ... 26. M. murrayana 1. Maireana sclerolacnoidcs (F. Muell.) P. G. Wilson, comb. nov. Echinop'iilon sekrohenoides F. Muell., Trans. Phil. Soc. Inst. Viet. 2:75 —Eriochilon nomen.-CW/cu sxkrolaenoides (F. Muell.) F. Muell. ex Benth., FI. Austral. 5:192 ( 1 9.10).— Bassia sclerolaemides ( F Muell ) F. Muell., Census 1:30 (1 882).-flm-.vmcr/oc/;fm« Tate, Handb. FI. Extratrop S AustVa 51 and 218 ( 1 890).— .4«i-/rofias-vw .ukrolaenoides (F. Muell.) Ulbrich, Nat ftanzenf ed 16c:532 (19.34). Type: “Desert of Lake Torrens, of the Murray and ’ Darling “ (Lecto- Cudnaka, F. Mueller, MLL, n.v., fide E. H. Ising, Trans. Roy. Soc. S. Austral. 88:72 ( 1964)) Chenoka dallacinaria Benth., \.c. .—Bassia daUachvana (Benth.) F Muell Census 1 - 30 (1882). Type: Murray river, Dallwhy (MEL, tide Ising l.c., n.v.). ' ' Ascending to erect perennial to 30 cm high. Branches slender lanate Leaves alternate, narrow-oblong to semi-terete, 5-10 mm long, acute, appressed villous (to sericeous). Flowers solitary, hermaphrodite, densely silky lanate. 18 usually congested towards the apex of the branches to form dense leafy spikes. Fruiting perianth cartilaginous to bony, obscured by the dense silky Innate indumentum: attachment small; tube hemispherical. I mm high and 2 mm wide at apex, 5-ribbed with the rib opposite the radicle deeply grooved; wings 5. opposite the perianth lobes, linear-acuminate (spine-like) eventually spreading, ca. 3 mm long; appendages within wing 5, erect, opposite the perianth lobes, oblong, ea. 3 mm long, each shortly to deeply divided into two acuminate lobes, upper perianth convex, hard, 5-lobed. Utricle thick; pericarp mem- branous, villous; style slender, exserted, villous. (Fig, 4H F, Map 14.) Distribution: Western New South Wales, South Australia, Western Australia south of the latitude of Carnarvon, southern portion of the Northern Territory. New' Sou III Wales: Broken Hill, A. AW/.v 284 (AI)W'). Soi III Australia: Near Port Pirie. M. Koch 213 (PERTH). W'rsTERN Australia: II mi W of Yalgoo. A. S. George 7960 (PERTH): l.eonora. W. E. Hlackull 363 (PERTH); 3 mi S of Reiil, I Sept. 1962, 7'. E. II. .Iplin (PERT H). Maircana sclerolaenoUles docs not easily fit into any of the described genera. The wings are spiny (whereas in most species of Maircana they are Hat) and the erect appendages are strictly opposite the perianth lobes, unlike other species which possess similar appendages. Both Anderson (1923) and Ising (1964) placed M. sclerolaenoicics in Bassiu sect. Kriuchiton as the only species. 2. .Maircana criantha (F. Muell.) 1^. G. Wilson, comb. nov. Kochin erievutta F. Muell., Rep. Babbage's Exped, 20 (1859). - K. viHosii var criaiilha (F Muell.) Moore ct Bctchc, Handb. FT. N.S. Wales 110 (1893). Lectotvpc: Elizabeth Creek Babbage's Expedition (MEL), ' ’ ?A'. cunvciva Ising, Trans. Roy. Soc. S. Austral. 78:112(1955). Tvpe: Evelvn Downs F H Isiiif ,' (holo AD, iso NSW). ^ Subshrub to 50 cm high. Branchlcts closely pubescent bearing persistent leaf bases. /.e«rE'.y alternate. Ileshy, linear, obtusely trigonous in cross section, 10-30 mm long. 1-5 3 mm wide, acute, densely sericeous. Flowers pre- dominantly dioecious, solitary or in pairs, arranged in leafy spikes towards the branch apices; perianth cupular. .5-lobcd, denscK' villous outside and within at base; ovary ovoid, sparsely villous, style of two tiliform stigmas to 6 mm long; male novvers with prominently exserted stamens. anthers"l -5 mm long. Fruiting perianth densely long silky lanatc all over; attachment small; tube obconieal to cupular, ca. 3 mm high, chartaeeous, 5 I ribbed (an inter- mediate rib opposite the I'Eidiele) or occasionally 10 ribbed; wing coriaceous, horizontal, deeply and irregularly divided into 5 lobes which'arc opposite the perianth lobes, ca. 2-5 mm wide and 10 mm diam.; upper perianth lobed to tube, completely covering ovary; lobes thick and coriaceous in lower two- thirds, chartaeeous near apex. Utricle ■_ globose; pericarp crustaceous, sparsely woolly; seed oblique, embryo hippocrepiform, (Fig. 4G H. Map 24.) Distribution: South western Queensland, north western New South Wales, ciistern South Australia north of Pt. Augusta, southern portion of the Northern Territory. Queensland: S end of Bygrave Ra., 5 mi E of Warri Gate, 6 .tunc 1955, L. A. S. Johnson NhW South Waits: t owlers Ctap, Aug. 1955, A'. Allison (AD): 13 km S of Warri Gale. Sol III At:sTRALiA: 43 mi I-.Sl: of Lake I tart. A. C. licangicholc 20012 (AD): Coober Pedv. 5 ,luly I960. J. H. Clcloiul (AD); Balia Baltaiia Ck. 28 Sept, 1953, E. S. Froncis (AD); .Afkaringii Ck., 16 May 1891. R. Helms (Ml t,. NSW); L)ro ttIulT, Ytidnapinna, 13 JiiK 19,54^ F. AT Hilton (ADW); Evelyn Downs. E. H. Isiii.i; 3560 (NSW); Mt. Lvndlitirsl, Oet. 1895 ,V/. Koch 155 (MI:L); De Rose Hill Stn.. T. R. A'. I.o'hiim 853 (AD); 5 mi N of Dtilf Ck.! /. R. .\. Lothian 1358 (AD); 40 km NW of Leigh Creek. T. R. A'. Loihhm 3534 (AE))- Foot- hills of Emery Rd.. T. R. A'. Lothian 4795 (AD): 22 km I, of Pedirka, 7. R. ,V. Lothian 4819 (.AD); Lords Gorge. A. .Morris 1457 (.ADW, NSW); 20 mi N of Red l.akc, Stuarts Creek Sin., 2 Dee. 1960, D. Svmon (AITW); Andamooka, ./. zf. tVvber 1447 (.AD); Stokc.s Rd. to Cooper Ck, Wheetcr (MEL 42101); Woomera, /.. D. Williams 441 (AD). 19 Found principally on stony plains or on rocky hills. Maireana eriaiitha has a fairly uniform appearance but exhibits consider- able variation in fruit characters. The tube varies from turbinate to hemi- spherical and the wing from being shallowly to deeply lobed. The form which was described by E. H. Ising as a new species 'Kochia concava is only known from the type collection and this consists of only three short pieces. It differs from M. eriaiitha in the fruit shape and in having a short sericeous indumentum on the branchlets. The fruits of K. concava are in pairs and are firmly fixed to the stem, only to be removed with difficulty; the tube is obconical and shorter than in M. eriaiitha; the wing varies from slightly to deeply lobed and from -f horizontal to erect; it is thick and hard at its base; the upper perianth is convex. From the little material available it is not possible to say whether K. concava is a good species, a hybrid invohing M. eriaiitha, or a deformed state of the latter species. Typical M. eriaiitha has been found at the type locality of the former species. 3. Maireana cheelii (Anders.) P. G. Wilson, comb. nov. Kochia cheelii Anders., Proc. Linn. Soc. N.S. Wales 59:270 fig. 3-4 (1934). Type- Zara, Dec. 1913, E. Officer CNSW). Small erect caespitose perennial with a woody stock and swollen tap-root. Branches slender, striate, closely woolly when young. Leaves alternate, fleshy, slender, semiterete, ca. 6 mm long, glabrous, concave above, very shortly spurred at base. Flowers solitary or in pairs, hermaphrodite; perianth deeply lobed, shortly woolly. Fruiting perianth wheel-shaped, cartilaginous to woody, somewhat woolly above, in all 5-6 mm diam.; tube shortly convex (depressed in centre), ca. 2 mm diam., strongly lO-ribbed, with a solid boss- like base; wings 5, distinct, fan-shaped, cartilaginous, horizontally radiating, each up to 2-5 mm long; upper perianth flat, obscuring ovary, shortly lobed, the radicular slit extending to tube where it continues as a more prominent rib. Utricle discoid; pericarp thin, glabrous; style very short. (Fig. 5G-H, Map 26.) Distribution: Near the New South Wales-Victoria border between Bendiao and Flay. New' South Wales: I n-ii N of Murray R. due south of Moulamein, 7 Nov. 1947, D L W Henderson (NSW); Deniliquin, Sept. 193.5, J. C. McCaw (ADW); Tchelery Stn , 9 Dec 1957, W. A. Mnirhead (NSW); 24 mi S of Hay, J. C. DeNardi 358 (NSW). Victoria: 20 mi N of Bendigo, 29 Mar. 1947, D. Henderson (NSW). Maireana cheelii is a very distinct member of the genus and shows no close affinity with any other species. J. FI. Willis in "A Flandbook to Plants in Victoria" 2;103 (1972) records that the tap-root tastes of coconut when fresh. He suggests the vernacular name Chariot Wheels . alluding to the wheel-like appearance of the fruiting perianths. 4. Maireana luehmannii (F. Muell.) P. G. Wilson, comb. nov. M'^'el'-’Vict. Nat. 7:47 {mO).~Anstrohassm luehmannii (F. Muell.) 2. I6c:532 (1934). Type: Finke River, IF. F. Schwarz (MEL fide E. H. Ising, I.c., n.v.). ’ Woody divaricately branched perennial to 40 cm high. Branchlets woolly when young. Leaves alternate, succulent, obovoid, ca. 5 mm long shortly sericeous, narrowed into a petiole-like base. Flowers hermaphrodite’ in axillary pairs, pubescent. Fruit woody, somewhat flattened and appressed to stem; attachment broad and flat; tube short, convex, ca. lO-ribbed- wings 5 woody, 2-3 mm long, spreading and often irregularly curved each ’ divided into two (or more) spiny lobes; upper perianth flat, divided to base into 5 imbricate lobes; radicular canal prominent. Utricle glabrous; style short pilose: seed thick and discoidal, endosperm prominent. (Map'll.) ’ Distribution*: Northern South Australia, northern eremean Western Australia Northern Territory. ’ *Partly after E. H. Ising (1964). 20 South Australia: Dalhousie Springs, S. A. White (AO. n.v.). Western Australia; Lake Auld, A. S. George 9142 (PERTHi. Northern Territory: 20 mi SSW of Mongrel Downs, P. K. Latz 746 (PERTH). E. H. Ising (1964) placed this species in the nionotypic section Bassta sect. Spinosissimae. Its affinities are, however, obviously w'ith M. amoena which it resembles closely both in vegetative and tioral morphology. 5. Maireana amoena (Diels) P. G. Wilson, comb. nov. Kochia amoemi Diels, Bot. Jahrb. 35:183 (1904). Type; Near Bullabniling, Oieh 5200 I B, now destroyed). Brittle undershrub to 40 cm high. Branchlets woolly when young. Leaves succulent, globular to ovoid or terete. 5-15 mm long, glabrous to sericeous. Flowers hermaphrodite, in pairs, covered with a short dense indumentum. Fruii pilose, woolly, or glabrescent; base angular, truncate; tube short hemispherical, woody (rarely firm and spongy); wings 5, horizontal, oblong to flabelliform, (0-5) 1-5-3 mm long, chartaceous to coriaceous or spongy (rarely almost altsent); upper perianth lobed to base, coriaceous (to spongy), completely covering ovary, lobes exduplicate valvate. so as to form 5 radial ridges, or imbricate; radicular canal prominent; utricle hemispherical (flat above); pericarp cartilaginous above, otherwise membranous; seed thick, endosperm prominent. (Fig. 5C-D, Map 12.) Distribution: Western Australia, from the latitude of Carnarvon south to Norseman. Usually found in the sandy rises around gypseous or salt lakes. Western Au.stralia; 250 mile peg on Great Eastern Highway, K. M. Alien 111 (PERTH) Dundas Rocks, A. C. Beaiiglehole 13210 (CANS. NSW. PERTH); White Well. Glenorn Stn 15 .Aug 19.38, iV. T. Burhidge (PERTH); 75 mi E of Wiluna. A. R. lairall 1896 (PERTH)- II mi E of Carnegie HS. A. S. George 5521 (PERTH); Nanmne. Sept. 190.3. W. V. Fitzgerald S of Wiluna. C. A. Gardner 2400 (PF.Rl H); Lake Wooleen, 23 Sept. 1950 A W llnniphries (PERTH): Carnarvon. ./. N\ Hatchinson 2 (PERTH); Cowcowing. Sept' 1904 .V/ Koeh 1 166 (NSW. PERTH); 15 mi SW ol' Nannine. N. It. Speck 739 (PERTH): Yarra Yarra Lakes near Three Springs, R. A. Sallrey 631 (PERTH); 5 km W ot Pindar P. G. Wilson 9958 (PERTH'. This species exhibits great variability but it does not seem practicable to delineate infraspecific taxaT The type form, which is by far the most common, is found from Cowcowing east to Zanthus and north to Nannine and Laverton. In this form the leaves are globular to obovoid and glabrous (except when young). The fruit has five fan shaped wings which are j_ in contact, and a short broad tube. The perianth lobes are exduplicate valvate so as to form 5 short radial ridges in the centre of the fruit. Near Lake Wooleen (leg. A. W. Humphries) is found a form with sub- terete and acute leaves which are sericeous (at least when young). The fruit has a very short tube and a broad flat base; the wings are coriaceous, irregular, oblong to obovate and shortly fimbriate, and the radial ridges on the upper surface of the fruit are very prominent. Towards the west coast of Western Australia near Three Springs (R. A. Saffrey 631) is found a form with a spongy perianth and very short ‘wings' (which are practically absent). Plants intermediate in character between those found at Three Springs and the type form of K. amoena \rd\g been collected at Pindar (Wilson 9958) and at Carnarvon (J. N. Hutchinson 2). 6 Maireana scleroptera (J. M. Black) P. G. Wilson, comb. nov. Kochia scleroptera J. M. Black, Trans. Roy. Soc. S. Austral. 46:568 (1922). Lectolype; R Alberga, June 1913, Miss Staer ihO). ■ i u Prostrate to erect perennial up to 30 cm high with a woody perennial base. Branches slender, striate, woolly. Leaves scattered, narrow-oblong to narrow- obovate 5-10 (20) mm long, sericeous to glabrescent. Flowers hermaphrodite, arranged in pairs in the leaf axils; perianth depressed globose, densely woolly. 21 Fruiting perianth cartilaginous to woody, moderately to densely woolly all over; attachment small; tube convex. lO-ribbed, ca. 2 mm diam.; wings 5. horizontal, cartilaginous, broadly oblong, Hal, crcnulate on margin, 1-2 mm long; upper perianth lobed to tube, open in centre, thick and hard lorming a disc-like ring around the membranous horizontal tips. Utricle discoid, pericarp coriaceous above otherwise membranous, glabrous; style short, sparsely villous. (Map 1 5.) Distribution: Found in the central eremcan area of Australia. South Ai'stralia: .^6 mi SH of ttverard Park Sin.. O. E. Symon 2706 (ADW); Wintinna Ck., 18 mi N ot Ml. Willoughbv, A. C. lieuiigicliole 20198 (AD). Westtrn Australia: KIdcr Creek, 2 mi \V ot Warburton Mi.ssion, A. S. George 3799 (PERTH); 30 mi NW of Ml. Davies, «. //. Kiicliel 76 (AD). Northern Terriiory: IOJ mi SW ot Alice Springs. 27 Sept. 1958, G. Chippendale (NT); Kings Canyon, George Gill Range, A. C. Beaiiglehole 20269 (AD). Maireana scleroptera is similar to some forms of M. amoena, especially to the form found towards the eastern limit of the latter's distribution. It is obvious that the two species are closely related and may even at one time have formed part of a continuous dine. As noted in several other species (e.g. M. lanosa and M. lobijlora) the wall of the fruiting perianth is penetrated in the position opposite the radicle by a canal the outer opening of which is normally obscured by the woolly covering. 7. Maireana brevifolia (R. Br.) P. G. Wilson, comb. nov. Kochiu brevifolia R. Br,, Prodromus 409 (1810). Salsolu brachyphylla Spreng., Syst Veg. 1:924 (1825), based on above. Tvpe: Australia (south coast), R. Brown (syn MEL 41973). Suaeda taniuriscina Lindl, in Mitchell, Journ, Trop. Austral. 239 (1848). -Encliylaena tamar- isciiu! (Lindl.) Druce, Bol. Soc. E.xch, Cl. Bril Isles 4:621 Eochia tamariscina (Lindl.) Black. Trans. & Proc. Roy. Sac. S. Austral. 47:368 (1923). Type: T. L. Mitchell 196, 18 July 1846 (bolo: CGE. iso: AD. MEL). Kochia thrniilolia I.indl. in Mitchell. Journ. Trop. Austral. 56 (1848). Type: Mara Creek T. E MiirhcU. 12 l eb. 1846 (CGE, photo seen). Bushy shrub 0-2-1 m high. Brancfilels slender, striate, sparsely to moderately lanate and with axillary tufts of wool. Leaves scattered, fleshy, obovoid to slender fusiform, 2 5 mm long, narrowed at base into a short petiole, glabrous (or sparsely pilose when young). Flowers solitary, hermo- phrodite, ebracteolatc, glabrous apart from woolly ciliate lobes. Fruiting perianth glabrous; attachment small, not hollowed; tube shallowly hemi- spherical, thin walled, faintly lO-ribbed, ca. 2 mm diam., W'ings 5, horizontal, chartaceous, arising from base of perianth lobes, each flabelliform. 2-3 mm long, with delicate brown venation; perianth lobes free to tube, thick and fleshy (sharply demarcated from wings), not completely obscuring the dis- coidal utricle. (Fig. 5A-B. Map 5.) Distribution: Found in all mainland Australian states but generally south of 26° of latitude. Queensland: Armadilla, between the Warrego and Maranoa, W. Barton 86 (MEL). New South Wales: Condobolin, 26 Feb. 1951, H. K. C. A/o/>(NSW). Victoria: Corio Bay, April 1901, G. (I. Pescott (MEL). South Austrai ia: 32 kin N of Pt. Augusta, T. R. N. Lothian 2659 (AD). Western Auserai.ia: I kni W of Dtikin, P. G. Wilson 6456 (PERTH). Northern Terriiory: I lermannsburg. Finkc R.. G. F. Hill 64 (MEL, NSW). Found in light to heavy soils often under slightly saline conditions. An early colonizer of disturbed land. Maireana brevifolia is very similar to M. diffusa. They may be dis- tinguished by diflerences in leaf shape and by the absence of bracteoles in the former species (sec also note with M. diffusa). The original description of Suaeda tamariscina was very brief and was based solely on flowering material. Bentham (1870) considered it to be a synonym of Encliylaena tnicrophylla and subsequently the two names have 22 always been considered to be synonymous and have been applied in their combinations under Kochia to plants which should correctly be referred to either A/, decahcms or to M. niicrophylla. In the 19th century literature the epithet dnicrophylUr was adopted (following Bentham) while in the 20th century the earlier epithet ''lamciriscina' was taken up. J. M. Black included Kochia lamarsicina in the second edition of his Flora of South Australia (1948) and inadvertently correctly applied the name, for although his description refers to A7. niicrophylla (of which he had seen New South Wales material) the one South Australian specimen in his herbarium, determined by him as K. tamarischuL is in fact a flowering specimen of K. brcvifolia. This species and A7. diffusa appear to be most closely related to M . decalvans and M. suaedifoUa. 8. Maircana diffusa P. G. Wilson, sp. nov. Fruliciihis yirgaiiis 0-5-1 ni alius. Folia carnosa, obovoidea vcl angiisle lusiformia, .t-15 mm tonga, hreviler pctiolata, sparse appresse pubcscentia yet glalsra. F/orry solitarii. bibraclcolali, braeteolis angustc triangularibus, ca. 0-7 mm longis. Pcnaiiihiiim Iructihcans illabrum, tubus convexus, ca. 2 mm diam., parictc lenui'. alae 5, horizonlales, llabeUiformes, ca. 2 mm longae, lenuiter chartaceae; periamhii lobi libri, inllexi, crassi sod vix carnosi, ad alas continui. Tvpc- I km W of l^ukin near Cow'cowing Lakes, Western Australia, l.t Mar. 1968. /’. G. Wilson 6461 (holo; PERTH, iso: AH. CANB, K. MEL. NSW). Virgately branched subshrub 0-5 1 m high. Branches slender, striate, with a short, thin, woolly indumentum when young. Leaves scattered, tleshy. obovoid and ca. 3 mm long on the short lateral branchlcts. narrowly fusiform to 15 mm long on the erect branches, sparsely appressed pubescent to glabrous, narrowed at base into a short flaltened petiole. Flowers solitary, hermaphro- dite, depressed globose, glabrous, with fleshy imbricate inflexed lobes, sub- tended by a pair of narrowly triangular bracteoles ca. 0 -7 mm long. Lriiiiing perianth glabrous; attachment small; tube convex, thin-walled and weak, faintly 10-ribbcd. ca. 2 mm diam.; wings 5, hori/ontal. completely separate and arising from base of perianth lobe, each llabellitorm and slightly im- bricate, ca. 2 mm long, thinly chaiTaceous. with fine pale brown radiating veins when dry; upper perianth horizontal, divided to base, lobes broadly imbricate, woolly ciliate, thickened but scarcely tleshy, continuous with the wings, the radicular slit extending as an overlap a short way down tube; utricle dis- coid. glabrous, style short. (Fig. 7G FI, Map 24.) Distribution: Western Australia, between Dalwallinu and Merredin. Western Ai;.straeia: 2 mi W of Kununoppin. Dec. 196.5, C. V. Malcolm (PERTH); 57 km E of Southern Cross, P. G. Wilson 4069 (PERTH). Found in saline soil, frequently around salt lakes. This species is .similar to A/, brcvifolia but has a much more restricted distribution and is not such a forceful colonizer of disturbed lands. Maireana diffusa dilTers from M. brevi folia in having a more slender and lax habit, longer leaves on the older branches, and a pair of bracteoles beneath each flower. In the fruit of A/, diffusa the perianth lobes are imbricate, flat, and almost cover the utricle; in K. brevifolia they are open, thick and fleshy so as to be clearly demarcated from the wings, and do not completely covci the utricle. 9. Maireana oppositifolia (F. Mucll.) P. G. Wilson, comb. nov. Kochia opposi’ifalia F. Mueff., Trans. & Proc. Viet. Inst. Atfvancem. Sci. I :I54 (f855). Type: Spencer Gulf. I85t. F. Mueller (MEL 45944). Compact subshrub 0-3 I m high. Branchlcts finely woolly when young. Leaves opposite (or in inland cremean foims alternate and often m tight eius- ters), sessile, spurred, narrowly ovate and up to 4 mm long to deltoid and ca. 1 mm long (when clustered) obtusely trigonous in cross section, shortly sericeous to glabrcscent. Flowers predominantly dioecious, shortly lobed. Fruiting perianth glabrous except for the pubescent lobes; attachment small, slightly 25 hollowed; tube broadly obconical to convex, I -5-2 mm diam., thin walled, ribbed; wings 5, completely free and arising from base of perianth lobes, chartaceous. prominently nerved, unequal, flabelliform, spreading unevenly (the two smaller sub-erect), in all ca. 7 mm diam.; upper perianth very short, lobed to wings. Utricle depressed globose; pericarp thinly crustaceous, sparsely woolly; style firm, exserted. ca. 0-7 mm long, lanate. (Map 22.) Distribution: Western Victoria; coastal or near coastal South Australia; coastal Western Australia as tar north as Shark Bay and inland as far north as Lake Moore and Comet Vale. Victoria: Mildura, 25 Jan. 1937, IV. Zimmer {MEL). South Australia: Price, Yorke, Peninsula H. Copley 3038 (AD). Western Australia: 9 mi S of Pingrup, K. Newhev 3083 (PERTH); 38 km NE of Southern Cross on road to Koolyanobbing, P. G. Wii.son 6437 (PERTH). Maireana oppositifoUa is always associated with saline conditions. This species varies considerably in habit and in leaf shape and arrangement. The south coast plant is erect with steeply ascending branches and the leaves are opposite, 3-4 mm long. The eremean form found in Western Australia from Shark Bay south east to Norseman is a divaricately branched sub-shrub in which the leaves are alternate and often clustered into compact glomerules. Intermediate forms are found and, at least in part, the plant habit appears to be a phenotypic response to the environment. The fruit in plants from all areas is virtually identical. Two other species of Chenopodiaceae are similar in the vegetative state to the eremean form of M. oppositifoUa. The first is M. glomerifolia which may be distinguished by its more tightly compacted leaf-clusters that are also more globular. The second is an undescribed species of Roycea in which the leaves are similar to those of M. oppositifoUa but the branches have a coarser indumentum. The Roycea species has hermaphrodite flowers, and perianths which are lobed to the base; both it and M. glomerifolia have fruit which is unmistakably different from that of M. oppositifoUa. 10. Maireana enchylaenoides (F. Muell.) P. G. Wilson, comb. nov. Ba.s.s1a cnchytaeiwides F. , Muell., Sysl. Census FI. PI. 1:30 (1882), Inon B. vithm Wallirh ex G. Don (I837)i based on lollowing.^£/ie/n/(/e/i« villo.w F. Muell., Trans Phil Inst Mssar** <^^^f) ~Heterochtamy.s vittosa F. Muell., Sec. Gen. Rep. Gov. Bot. Victoria 0854) nomen.— C/ie/wWa enchylaenoides F. Muell., Fragm. 10:92 (1876), nom. illeg (suner- fluou.s name). -Aw/iw crassitoba R. H. Anders., Proc. Linn. Soc. N.S. Wales 51 :383 ( 1926- nom. ilieg. (superfluous niunc). — Chenoleo villosa (F. Muell.) Ewan, F) Viet 458n0'^n Diniah vdh.sniE Muell.) Ulbrich, Nat. Pfianzenfam. ed. 2. I6c:537 (1934) — bi/rio/a crassi- loha Beadle Mud FI. N.F New S. Wales pt. 2:199 (1972) nom. inval. Type: “In loamy places near Adelaide and m Bacchus Marsh" (syntypes MEL). ^ Small perennial to 20 cm high; branches slender, decumbent to erect sparsely puberulous to villous, arising from a woody rootstock. Leaves scattered, somewhat fleshy, narrowly oblong-elliptic, obtuse, 4-10 (20) mm long, sparsely pubescent to villous. Flowers solitary, axillary, hermaphrodite depressed gbbose. hirtellous, sometimes subtended by a pair of small narrowly tnanplar bracteoles. Fruiting perianth sparsely puberulous, drying black small; tube slightly convex, thin walled and coriaceous -F 10 ribbed, ca 3 mm diam.; wmigs 5, completely separate and arising from base of perianth lobes, only slightly exceeding tube, in all ca. 4 mm diam coria- ceous, each broadly lunate and unevenly imbricate (auriculate when overlao- ping) margins somewhat inflated and sometimes incurved; upper nerianth completely divided into small deltoid lobes which are continuous with the wings. Utricle discoidal; style short. (Map 25.) DUtrihution: Southern and eastern Australia in areas S of 28 latitude and with over 300 mm rainfall. Queensland: Nindigully, Dec. 1937, R. /tuclCANB); Gilruth Plains, G. H Alien (CANBI M7or!T4!{CA^B). Narrandera-Grong Grong, C. )F i.' 24 Victoria: Nathalia, Goulburn Valley,/. H. Willis 10 (MEL). South Au.stralia: MooIooIoo Stn., Oct. 1920, B. Beck (AD). Western Australia: Fitzgerald River valley, A. S. George 10012 (PERTH). The form taken by the fruit in this species is so similar to that found in M. hrevifolia and M. diffusa as to not justify it being placed in a separate genus. The nomenclature surrounding this species is somewhat involved, this being partly because all the synonyms cited above are based on the same type. The confusion commenced in 1876 when Mueller transferred the species from Enchylaena to Chenolea, for he should then have retained the epithet "v///rAvr/”; the combination "C. eiichylaenoides'' was therefore illegitimate. Subsequently (1882) on transferring the species to the genus Bassia (Chenopodiaceae) he was not able to use the epithet "viUosa” since it had already been used by G. Don* in a genus of the same name but belonging to the Sapolaceae. Mueller was therefore free to use the epithet ‘'euchylaeiwides" w'hich is to be treated as new. When Anderson in 1926 transferred the species to Kochici he should also have adopted the epithet ^'eiichylaenoides" (since the epithet "villosa" was not available). Anderson's new name "Koehia crassiloha" is therefore illegitimate being superfluous. In the genus Maireana I have preferred to reserve the epithet "villosa" for that species which has for long been known under this name in the genus Koehia: 1 have therefore taken up the next available epithet for the species here described but which unfortunately has been used in the genus Koehia for the plant now referred to M. tomentosa. II. Maireana lobiflora (F. Muell. ex Benth.) P. G. Wilson, comb. nov. Koehia hhiflora F. Muell. ex Benth., FI. Austral. 5:184(1870), Type: Darling River. Victoria Exploring Expedition. I Nov. I860, Beckler {ho: MEL). Decumbent to erect perennial with a woody stock, lo 0 -5 m high. Branches slender, striate, lanate. Leaves alternate, linear to very narrowly elliptical, 7-15 (25) mm long, sericeous, i similar in size and shape throughout the plant. Flowers solitary, hermaphrodite: perianth woolly, deeply 5-lobed; ovary woolly. Fruiting perianth shortly lanate, coriaceous to crustaceous, attach- ment small: tube slightly convex, faintly costate; wing horizontal, of 5 spathu- late to flabelliform lobes (when broad they may be contiguous along their margins), in all to 10 mm diam.; upper perianth slightly convex, closed over the utricle, radicular slit extending to a short way down tube; erect processes (on upper perianth) 4 ■ 2, alternating with the perianth lobes (the contiguous pair being placed with one on either side of the radicular slit), processes narrow- spathulate to clavate, to 4 mm long, frequently expanded or lacerated at the apex or deeply bilobed. Utricle discoidal, sparsely villous; style villous, shortly exserted. (Fig. 3A-B. Map 17.) Distribution: Western New South Wales, South Australia, Western Australia, southern portion of the Northern Territory. New South Wales: SE corner of Cobham Lake, /. C. DeNardi 824 (NSW, PERTH). South Australia: 6 km E of Argepena HS., T. R. N. Lolhian 3241 (AD). Western Australia: 14 mi E of Carnegie HS., A. S. George 5522 (PERTH). Northern Territory: 25 mi E of Heavitree Gap, Undoolya Stn.,./. R. Macoiiochie 1049 (NT). This species exhibits considerable variation over its geographical range. In New South Wales, central South Australia and east-central Western Aus- tralia the leaves are very narrowly elliptic to linear and the wings in the fruit have prominent intervening sinuses. This is the type form of the species. In the Northern Territory the leaves are narrow-elliptic and thin and the wings of the fruit are broad, touching, and without intervening sinuses, and they may sometimes be partly fused. This has often been identified as M. lanosa. but the latter species is afso found in the Northern Territory and has a quite different appearance and does not appear to hybridise with M. lobiflora. *Bassia villosa Wallich ex G. Don. Gen. Hist. 4:36 (1837). 25 In the southern portion of Australia, over the Nullarbor Plain and eastwards to the Flinders Ranges, is found a form which is decumbent in habit and has small leaves. The wing lobes in the truit of this form are broad and contiguous and each of the four normally 'single' erect processes is divided to the base. For comments on the fruit morphology, and intergradation with M. lanosa, see notes under the latter species. 12. Maireana lanosa (Lindl.) P. G. Wilson, comb. nov. Kochia lanosa Lindley in Miichell, Journ. Expect. Trop. Austral. 88 (1848) Type: [near the Narran R.] 8 Mar. 1846, 7'. /.. Miichell (iso; MEL). K. lanosa var. minor Benth., FI. Austral. 5:184 (1870). Type: Murchison River, Oldfield (MEL). Open perennial with a woody stock, to 0-5 m high. Branches slender, striate, lanate. Leaves alternate, somewhat lleshy, elliptic to narrow-elliptic or narrow-obovate, ca. 4 mm long in the upper (fruiting) branches, 10-20 mm long in the lower branches, sericeous. Flowers solitary, hemaphrodite, perianth densely woolly, 5-lobed. ovary woolly. FruiUng perianth sparsely lanate, coriaceous to crustaceous: attachment small; tube slightly conve.x, faintly costate; wing simple, horizontal. 7 12 mm diam., with a single radial (radicular) slit, margin entire or crenulate; upper perianth slightly conve.x, closed over the utricle, radicular slit extending to a short way down tube; erect processes (on upper perianth) 4-2, linear to subulate, 3-4 mm long, alternating with the perianth lobes (the contiguous pair being placed with one either side of the radicular slit). Utricle discoidal; pericarp chartaceous. sparsely villous; style villous shortly exserted. (Map 18.) Dislrihtilion: South-western Queensland, central and western New South Wales, eastern South Australia, north-western Western Australia, Northern Territory. Queensland; Yappunga near Tliargomindah. April 1885. Mrs. Spencer (MEL). New' South Wales: Bambamero ( Lake Pamamaroo) .t Nov. I860, Beckler (MEL). South Australia: Nilpena, R. Helms 53 (NSW). Western Au.stralia: 33 mi S of Learmonth, ,4. S. George 6536 (PERTH). Northern Territory: 90 km NE ot Alice Springs, M. L. Benda (.AO). Although very similar to M. lohiflora, over most of its range M. lanosa is distinct from that species (with which it yet appears to be sympatric). Maire- ana lano.sa dilTers from M. lohiflora in the following characters: (I) The leaves of the fruiting branches are usually noticeably smaller than those on the vegeta- tive portions of the plant. (2) The wing of the fruit is simple. (3) The erect processes arc simple and never clavate. In Western Australia, in the area between the Wooramel River and Boolardy Station, the diflerences usually found between the two species are not present and in the same region can be found plants which dilTer only in the degree of lobing of the wing, however, because of the distinctiveness of the two taxa elsewhere in Australia, 1 have retained them as distinct species. Near the west coast ot Western Australia between Learmonth and the Murchison River is found a form with smaller fruits; this is the ‘var. minor" of Bcntham. The characters which distinguish it from the type variety are not sufficiently clear for its status to be maintained. In both M. lanosa and M. lohiflora a passage is present in the wall of the mbe opposite the radicle, passing outwards to the radicular slit. This passage is identical in lorm to that lound in M. ciliala and related species and generally in others where the wall of the tube is thick and hard. The erect processes of the fruits of M. lanosa, as also in M. lohiflora, are presumably homologous to the spines found in most species of Sclerolaena (the Austialian members ot the genus Bassia ). In Sclerolaena the spines also alternate with the perianth lobes and, when six in number, also have the 4 -L (2) arrangement. 26 13. Maireana ciliata (F. Muell.) P. G. Wilson, comb. nov. Kochia ciliata F. Muell., Rep. Babbage’.s F.xped. 20 (IS.-ig). Type: Wonnomulla. Emu springs, between Stuarts Creek and Margarcth Creek, Babbage's F!xpedilion (MFI.). Decumbent to erect plant with woody perennial base and slender woody taproot, to 10 cm high. Branches slender, woolly. Leaxes alternate, (linear to) nan ow-oblong, acute, ca. 10 mm long, sericeous. Flowers solitary, her- maphioditc, in congested lealy spikes and also scattered along branches, densely silky villous; perianth patellilorm. Fritiiing perianth compressed lenticular, pentagonal, ca. 3 mm diam., horny, covered with .silky villous indumentum; attachment small; lower surtace almost Hat to slightly conve.x, faintly costate; upper sLirlace convex in centre above the utricle otherwise flat and wing- toiming \vith 5 prominent radiating intertepaline ribs which extend to the angles ol the wing, the radicular slit passing along one rib to the wing margin which is there notched; perianth lobes horizontal, triangular, not completely obscuring ovary. Utricle discoid, sparsely villous above; style firm, exserted. (Fig. 3G-H, Map 20.) Distribution: Far western New South Wales, central and north eastern South Australia. Nfav South Walks: Fowlers Gap Station, 29 Aug. 1952. E. G. Cii/hhertsun (NSW). South Australia: Farina Ck., Flinders Ra., 7’. R. N. Lothiuii 1037 (AD). 14. Maireana pentagona (Anders.) P. G. Wilson, comb. nov. Kochia penlagoiui R. II. Anderson, Proc. Linn .Soe .N.S. Wales 51 :385 t.27 (1926) --Chcuotca peiita.eoiia (Anders.) Ewart, FI. Viet. 458 (1931). Type: Trangie, A. Morris 1288 (holo' NSW, iso: ADW). Prostrate to decumbent plant with small perennial woody base and longHcshy taproot. Branches slender, woolly. Leaves alternate, linear, acute, 7-12 (20) mm long, appressed villous, often becoming glabrescenl. with age. Flowers solitary, hermaphrodite, densely woolly villous. Fruiting perianth woolly villous, discoid, ■ pentagonal in outline. 2-5-4 mm diam. (without the indumentum), cartilagin- ous to crustaccoLis: attachment small; lower surface convex, ribbed, eontinttous with the narrow horizontal rim (wing) which is sometimes divided into live very short truncate lobes; upper surface raised into a thick, hard pentagonal (or star-shaped) ridge or platform formed from the convex thickened bases ol the perianth lobes, within the ridge the lobes are depressed and membranous, radicular slit extending to margin of wing. Utricle di.scoid, pubescent; style slender, firm, exserted. (Fig. 3E F. Map 19.) Distribution: Central and south western New South Wales, western Victoria, south eastern South Australia. New South Walks: 55 mi N of Dcniliquin, /'. M. lytmile 1722 (NSW). Victoria: Kcrang, Ocl. 1887,./. Minchia (MEL). South Australia: Renmark, Jan. 1917,4. M. Black (NSW) Western Australia: 75 mi N of Carnarvon, J. S. Beard i5\ I (Kings Park). The ridge on the upper surface of the fruiting perianth is pentagonal to star-shaped in outline with the five angles extending towards the wing margin. If the wing is very narrow, it may extend as a short acute lobe opposite the angles. If the wing is more developed it may be slightly retu.se opposite the angles, thus dividing into five short, truncate lobes. An intermediate cond- ition gives an entire outline to the wing. The only record for M. pentagona from Western Australia comes from an area well isolated from the otherwise known distribution of this species. The specimen is very similar to the Eastern States' plant but has a greater devel- opment of the wing in the fruiting perianth. Maireana pentagona appears to be closely related to M. ciliata and M. coronata; it is possibly related (but more remotely) to M. platyptera. 27 J5. Maireana coronata (Black) P. G. Wilson, comb. nov. Kochia coronata .1. M. Black, Trans. Roy. Soc. S. Austral. 41:43 t.9 (1917). Lectotype; Cootanoorina Creek, 7 May 1891, R. He/nis (ho\o: AD, iso: MEL). Decumbent to erect plant with woody perennial base and slender woody ( ?) taproot, up to 15 cm high. Branches slender, woolly. Leaves alternate, linear, acute, 7-20 mm long, appressed villous or sericeous. Flowers solitary, hermaphrodite, in congested leafy spikes or scattered along the stems, densely long silky villous; perianth base broad and flat; upper perianth forming a membranous erect cup that is surrounded by a fleshy circular disc (this disc develops in fruit into the corona). Fruiting perianth like an inverted academic hat (‘ mortar-board"), horny, silky villous, with a flat base which is continuous with the narrow circular wing, in all ca. 4 mm diam.; upper surface with a cup-shaped outgrowth (corona) to 2-5 mm high which is soft on the margin but otherwise horny; upper perianth within and towards the base of the corona, horizontal, i completely obscuring utricle, the radicular slit extending through the corona to the notched margin of wing. Utricle discoid, occupying the base of the corona; style firm; exserted (from perianth), eventually deciduous (Fig. 2F-G. Map 2.) Distribution: Central and south western Queensland, north western New South Wales, north eastern South Australia, and the south eastern area of the Northern Territory. Queensland: Longreach. Mitchell District, S. T. Blake 6595 (NSW). 45 mi NNW of Wilcannia on road to White Cliffs, J. C. DeNardi 261 NSW). South Australia: Maciimba Stn., Nov. 1950, E. H. (AD). Northern Territory: 12 mi W of Huckitta Stn., M. Lazarides 5937 (NSW). Maireana coronata is similar to M. ciliata and. prior to its description of" h were identified as the latter species. Bentham (1870) cited three collections under Kochia ciliata, one of them (Babbage’s Expedition) IS the type of this species while the other two (Dallachy and Mit- chell) are of M. coronata. The two species may be readily distinguished for m truit the latter possesses a well-developed cup-shaped corona which develops as an outgrowth from the base of the perianth lobes. It is in this position (the base of the perianth lobes) that the wing is formed in most species of Mariana, and it is possible that the corona is homologous to the normal wing while the apparent wing corresponds to the membranous extension of the tube base as found m the perianth of M. polrptervgia. 16. Maireana dichoptera (F. Muell.) P. G. Wilson, comb. nov. MEL Small erect plant to 20 cm high with perennial woody base. Branches SiT ■ C/? ^^25 mm long, appressed villous_ flowers hermaphrodite, solitary or in pairs, very depressed with mflexed perianth lobes, shortly tomentose outside. Fruiting perianth sparsely pilosulose. Straw-coloured when dry, largely consisting tff the six radially positioned vertical xvangs and a horizontal wing; attachment small; tube flat ca. 2 -5 mm diam., 6-nerved; wings delicate, faintly veined; horizontal wing )■ the tube. ca. diam.. wil^h a smde adial (radicular) slit, vertical wings 4 i 2 on upper side of perianth and inter- eXer"Jide opposite the radicle with one either side of the ladicular slit) attached to the horizontal wing and to the arched upper perianth, ca. 3-5 mm high; upper perianth formfng a convex exteldm^\o"'w1n^'^/f T membranous lobes, the radicular slit ex ending to wing. Utiicle discoid; pericarp crustaceous above sparsely villous around style base. (Fig. 3C-D. Map 25.) ^parseiy 28 Distribution: Central Queensland. Queensland: Tambo, Barcoo, Schneider (MEL); near the Alice River, 1884 C W Birch (MEL), This species is very similar to M. pentagona and its allies in both vegetative and fruiting characters. This resemblance is obscured by the presence of the delicate wings in M. dichoptera and by the dense indumentum on the perianth of M. pentagona. The wings correspond, however, to the horizontal rim and vertical ridges found in the latter species and in M. ciliata. 17. Maireana eriosphaera P. G. Wilson, sp. nov. Ramuli graciles, laxi lanati. Folia linearia vcl angiisti oblonga, acuta, ad !0 mm longa, sericea. f-lorc.'i dense spicati in seriebus verticalibns disposili, Pcrianthinni fructificans pedicellalLim cum indumento dense sericeo-lanato: tubus hemisphaericus vel turbinatus, 1 ■5-2-5 mm altus, chartaceus, margine incressato et undulato vel perbreviter 5-Iobato vel peranguste alato, rima radiculari in tubum continua; perianthium supernum in lobos minutos acuminates divisum, inflcxurn vel demuni clevatum in pilis lanatis pericarpio producto. Type: Upper Rudall River area, ca. 500 km S of Broome, Western Australia, 22 35 'S, 122 lO'E, quartz gravel and clay, spike brittle with 3 vertical rows of (towers, hairs on flower white, stigma red, 16 Aug. 1971, P. G. Wilson 10564 (hole: PERTH, iso: CANB, K, NSW). Erect loosely branched perennial to 30 cm high, woody at base. Branches slender and loosely woolly. Leaves alternate, linear to narrow-oblong, to 10 mm long, acute, sericeous. Flowers solitary, hermaphrodite, densely covered with a silky-villous indumentum, arranged in vetrical rows in dense terminal spikes 2-10 cm long; floral bracts similar in size and shape to foliage leaves; perianth broadly hemispherical; style lanate with 2-3 long stigmatic lobes which, with the stamens, are long exserted. Fruiting perianth on a distinct slender pedicel 0-5-1 mm long, densely covered on tube, wings, and upper surface of perianth lobes by a long spreading silky-lanate indumentum forming a soft ball ca. 10 mm diam., inner surface of tube and of perianth lobes glabrous; tube chartaceous, (hemispherical to) turbinate I •5-2-5 mm high, 5-10 costate; margin of tube thickened and undulate or of 5 short rounded erect divisions (wing lobes) continuous with tube or sometimes these united to form a very narrow coriaceous horizontal wing, the radial (radicular) slit passing for a variable distance down tube; upper perianth of 5 acuminate lobes ca. 1 mm long, coriaceous at base but otherwise membranous, inflexed or eventually raised up by the dense indumentum which, arising from the pericarp, overflows from the tube. Utricle di.scoidal; pericarp thin, silky lanate above. (Fig. 9A-B, Map 2.) Distribution: Eremean region of Western Australia. Western Australia: 16 mi N of Norseman. T. E. H. Aptin 1844 (PERTH); Nannine Sept. 1903 W. y. /•/tz.ccraW (NSW); Rudall River, 22 35'S. 122 lO'E, P. O’. Wit.son 10564 (PERTH). Maireana eriosphaera is normally only found growing within small shrubs, where, presumably, it is protected from animals. It usually occurs in sub- saline soil on the margins of salt lakes or at the base of breakaways. M. eriosphaera is similar in appearance to M. carnosa with which it has been confused in herbaria. In the field the former may be distinguished by its more slender and erect growth and by the flower spikes in which the flowers are vertically arranged contrasting with the spiral arrangement in M. carnosa. Other distinguishing characters are the floral indumentum which in M. carnosa is finer and more curled (floccose), the perianth shape, and the presence or absence of a pedicel. M. eriosphaera exhibits considerable variation in the shape and size of the fruiting perianth. In some specimens the margin of the tube (i.e. the erect wing) is only undulate, whereas in others it is 5-lobed. Occasionally the wing is slightly spreading and almost entire. Part of this apparent variation may be due to the stage of development of the fruit for in other characters the species is remarkably constant throughout its range. 29 18. Maircana carnosa (Moq.) P. G. Wilson, comb. nov. EchinopiUon carnosns Moc]. in DC., Prod. 1.^ 2:136(1849 ). — Clicnolea cciniosa (Moq.) Bcnth., FI. Austral. 5:190 (1870). Ilassia carnosa (Moq.) F. Miicll., System. Census Austral. PI. 30 (1882). Kochia carnosa (Moq.) Anders.. Proc. Linn. Soc. N..S. Wales 48: 353 (1923). Type: "In Nova-Hollandia seeus Swan-River. { DrnninioncI! n.l26)." n.v. Bassia lanuginosa C. T. White. Queensid. Agrie. .lourn. n.s. 15:216 (1921). Type: Twenty six miles east of the junction of the Mayne and Diamantina Rivers, Western (Queensland /'. Peihcr (iso: MEL. NSW). Herbaceous pei'ennial with woody base to 30 cm high. Stems sometimes decumbent with ascending bratiches, loosely woolly. Leaves alternate (or sometimes opposite), fleshy, narrow to broad oblong, to 10 mm long (the floral leaves often broader than the stetn leaves), silky villous to woolly or glabrescent. Flowers solitary, herniaphrodite, spirally arranged in dense terminal leafy spikes, densely covered with long white floccose wool: staniinal filaments and style long and sletider (3 5 mni). Fruiting perianihs sessile, enveloped in a thick tnass of wool ca. 8 mtn diam. and fortning stout woolly spikes; attach- ment voy small; tube crustaceous, very shallowly hemispherical, 2-3 mm diam., faintly to prominently costate; wing ci)‘cular, horizontal, vei'y narrow, to 0-5 n)m wide and in all 3-4 )nm diam.. with a single radial (radicular) slit; upper perianth crustaceous, flat or slightly infle.ved in cenl)'e, shortly lobed and not completely covering ovai'y; j'adicular slit extending to wing; utricle discoidal, pericarp thin, woolly above. (Fig. 9C-D, Map 23.) Disirihiition: South east Quee)island, northern South Australia, and the tem- perate and subtropical eremean regions of Western Austi'alia. <5uei;nsi.and: 26 mi E of .lunclion of May)ie and Diamanlina Rivers, .Ian. 1921, F. Pether (MEL, NSW). South Allsdvada: Evelyn Downs, 19 Aug. 1954. E. H. Ising (AD). Western Au.straiia: 5 mi W of MeekathaiTTi, N. //. Speck 553 (CANB, PERTH). Typically an eremean species inhabiting salt flats and sandy subsaline loa)r,. The type of EehinopsHon carnosus was stated by Moquin to be Drum)nond no. 126. This I have not seen. Howevei', Bentham (1870) cited Drmnmond 246 and a duplicate of this number in herb. MEL ag)'ees with Moquin’s description and may in fact be an isotype. 19. .Maircana niarginata (Benth.) P. G. Wilson, comb. nov. Enchyiaena nuirginala Bcnth., FI. Austral. 5:182 (1870). Type: "Swan river, DrnmmoncL 1st coll." (holo; N). ?E. micruniha Bcnth., op. cit. 181. -Bm.vw niicramiia (Benth.) F. Muell., Syst. Census Austral. PI. 30 (1882). Type: Western Australia. Drnninioiut 4th coll. no. 253 (iso: MEL). Kochia inassonii Ewart, Proc. Rov. Soc. Viet. .ser. 2. 20:82 t.lOb (1907). Tvpe: Cowcowing M. Koch. Sept. 1904 (holo: MEL, iso: MEL, PERTH). Prostrate to decumbent herb with a perennial woody stock. Branches slender, somewhat woolly when young. Leaves alternate, fleshy, linear to narrow-oblong, 5-10 mm long, loosely villous. Flowers solitary hernia- phrodite, depressed globose, pubescent above, subtended by a pair of deltoid bracteoles ca. 1 -5 mm long. Fruiting perianth when IVesh depressed barrel- shaped and constricted in middle, ca. 3 mm high, 4-5 tiDii diam. at apex, fleshy, dr ITit above and below', papillose on sides and puberulous above, tube with lower half soniewhat bulging, smooth, upper half with ten rounded ribs (on drying the upper half of tube beconies very broadly turbinate and thin walled while the lower hall lorms an expatided slightly spongy base); wing a very narrow horizontal extension to upper perianth, cntii'e except for the radicular slit: upper perianth flat, not completely covei'ing utricle, lobed about half way, t'adial (radicular) slit extending down upper half of tube as a prominent ridge which niay itself be longitudinally slit. Utricle lenticular, glabrous; pericai'p thin; seed with copious endosperm. (Fig. FA-B, Map 11.) DLstrihution: Western Australia. South West Distidct from Morawa south-east to Ravensthorpe. 30 Western Australia: Morawa, July 1903, C. Andrews (PERTH); Cunderdin, Sept. 1908, J. B. C/e/awi/ (NSW): Ravenslhorpe, P. G. Wilson 7086 (PERTH). Usually found in heavy soil in woodland. The holotype of Encliylaeiia micraiului at Kevv (K) and an isotype in Melbourne (MEL) both lack any trace of flowers or fruit. The vegetative appearance of these specimens is, however, similar to that of M. margiuata. For a comparison of M. marginata with M. excavuta. to which it is closely related, see under the latter species. 20. Maireana c.xcavata (J. M. Black) P. G. Wilson, comb. nov. Kochiu excavala J. M. Black, Trans, el Proc. Roy. Soc. S Austral. 47:368 (1923). Type: Spalding, 4 Nov. 1920 (holo: AD, iso: NSW). A. villosa var. Innnilis Benlli., FI. Austral 5:187 (1870). —Ak lonientosci var. hiimiUs (Bcnth.) Black, FI. S. Austral. 197 (1924) not as to description. Type: "desert country of Victoria and adjoining portion of N.S. Wales" (syn; Da/lacliw MEL 2114; Darlings River, MEL 42137; Skipton, It'. Wluin, MEL 43929). Small perennial herb to 20 cm high with a woody base. Branches slender, loosely woolly when young. Leaves alternate, narrowly oblong-elliptic, 7-12 mm long, somewhat fleshy, tawny appressed villous. Flowers solitary, hermaphrodite, with or without a pair of narrow-oblong bracteoles to I -5 mm long. Fruiiing perianlh fleshy when fresh; depressed barrel-shaped and con- stricted in middle, ca. 3 mm high, Hat abo\e. glabrous; lube (on drying) with upper half very broadly turbinate to conve.x. lO-ribbcd, thin walled, lower half expanded into a spongy hollow' base which when attached to the plant is filled by the cushion shaped receptacle; vying charlaceous, horizontal, circular and entire, ca. 10 mm diam., faintly nerved and with a single radial (radicular) slit; upper perianth flat, almost obscuring the ovary, deeply lobed, ciliate on the margin otherwise glabrescent, the radicular slit extending from perianth to base of tube as a narrow slit. Utricle lenticular; pericarp thin, glabrous; seed with copious endosperm. (Fig. 6G-H, Map 18.) Distrihulion: South central New South Wales, western Victoria, south-eastern South Australia. New South Wales: Waiiganclla, July 1903, E. 0//)V-7 68 Map 26 M. cheelii ■ M. appressa o 69 TO Figure I. Bassio costata: (A) Dextrorse fruit and embryo, (B) Sinistrorse fruit and embryo. Maireana glomerifoUa: (C) Sinistrorse fruit and ernbryo, (D) Dextrorse fiuit and embryo. Enchylaena tomentosa: (E) Fruit, (F) L.S. of fruit. Roycea spinescens: (G) Fruit, (FI) Embryo. (A B from P. Wilson 8985, C-D from P. Wilson 7560, E-F from P. Wilson 1041 1, G-H from P. Wilson 10964). 71 3 mm Figure 2. (A-B) Maireana prosthecochaeta; (C-D) Dissocarpus biflorus, p = perianth lobes, s = spines: (E) Kochia stellarix, from below, dissected to show embryo; (F-G) Maireana coronata. „ „ (A-B from R. O’Farrell s.n., C-D from R. Kuchel 2863, E from P. Aellen 977, F-G from E. FI. Ising, 3 August 1955). 72 5 mm Figure 3. (A-B) Maireana lobiflora; (C-D) M. dichoptera; (E-F) M. pentagona; (G-H) M. ciliaia. (A-B from R. Helms s.n., C-D from Schneider s.n., E-F from Eckert s.n., G-H from M. Koch 122). 73 Figure 4. (A-B) Maireana campamilata; (C-D) M. spongiocarpa; (E-F) M. sclerohenoides; (G-H) M. erianiha. (A-B from R. Kuchel 1085, C-D from N. Lothian 4594, E-F from P. Wilson 3114, G-H from J. Z. Weber 1447). 74 A B 5 min 5mm D , 3 mm 3mm Figure 5. (A.-B) Maireana hrevifolia: (C-D) M. amoetia; (E-F) M. melanocarpa: (G-H) M. cheetii. (A-B) from T. E. H. Aplin s.n., C-D from P. Wilson 6479, E-F from P. Richards s.n., G-H from D. L. Henderson s.n.). 75 10mm D 10mm F 5mm 5mm^ G H Figure 6. (A-B) Maireana glomerifolia; (C-D) M. platycarpa; (E-F) M. trichoptcra; (G-H) M excavata. (A-B from P. Wilson 8826, C-D from C. V. Malcolm s.n., E-F from A. S. George 415), G-H from R. Wallace 186.) 76 Figure 7. (A-B) Maireana margiiiala; (C-D) M. polypterygia ; (E-F) M. melanocoma; (G-H) M. diffusa. (A-B from P. Wilson 7086, C-D from A. S. George 1250, E-F from A. S. George 999, G-H from P. Wilson 6420.) Figure 8. (A-C) Maireana geor^ei : (D-F) M. turhinata; (G-H) M. convexa: (T-J) M. murray- ana. (A-B from K. Newbey 1544, D-F from P. Wilson 7595, G-H from F. H. Sharr 2731, 1-J from D. L. Wilcox 164.) 78 4 mm Figure 9, (A-B) Maireana en'osphaera: (C-D) M. carnosa; (E-F) M. tomentosa subps. urceohta: (G-H) M. tomentosa subsp. tomentosa. (A-B from P. Wilson 10564, C-D from A. Ashby 2597, E-F from J. B. Cleland s.n., G-H from P. Wilson 10486). 79 5 mm 5 mm Figure 10. (A-B) Maireana pvraivuhta; (C-D) M. schistocarpa: (E-F) M. villosci. {A-B from P. Wilson 9894, C-D from P. K. Latz 3160, E-F from A. S. George 4936). 80 INDEX TO NAMES Anisacantha 4 Auslrohus.sia luehmannii (.V . MutW.) XSVouch 20 .sT/m)/a(?«o(Vfes (F. Muell.) Ulbrich 18 Babbagia 5, 7 Bassia .. 2 Bassia sect. CnVfr/i/toH 19 sect. Maireaiici .. . . .. . .. ... ... . 14 sect. Spiiiosissimci .... .... .... .. . . . . 13,14 hrachyplera (F. Muell.) Anders 55 co(7K).sa (Moq.) F, Muell 50 (Bentli.) F. Muell 18 enchytufnokie.t F. Muell 24 eriochilon Jmc 18 lamipinosa C. T, White .... ... .... .... 30 liiehiiiaiinii f-. Muell. ... . .. . . . .. . . 13,20 m/cra/(t/(a (Benth.) F, Muell, 30 sclcroltienoides iV. Muell.) F. Muell 18 .stelligera (F. Muell.) F. Muell 56 \ illosa Witllich e\ G. Don 24 Chenolea .. ... .... ... ... . . . 3 Cheiiolea bkickii Fwart .... .... .. . .... 48 carnosa Benth. 30 r/(j//of7n'«H« Benth. .... . .... ... .... .. .. . 18 ench.ylaenoicics F. Muell. .... 24 (Anders.) Ewart T! scelrulaciwicics (V . Muell.) Benth 18 v/Y/trsa (F. Muell.) Ewart 24 Coilucarpus .... .. . .. . ... .. . .. ... .. .. 5 Cyriiwhitea 5, 7 Didynianthus . . ... ... .... .... ... . . .. . . 5, 7 Dissocarpus .... . . . . .... .. . .... ... . . .. . . 4, 7 Diaiahi 5,14,24 v///().vo ( F. Muell.) Ulbrich 24 Echinopsilon 3 EchinopsHon caniosus Moq. ... .... .... . .. .... ... .. . .... 30 sclerolaenoides ¥ . Muell .... ... .... .... ... .... 18 Enchylaena .... .... ... ... . .. . .. . 8 Enchylaena sect. Hcterochlamys F. Muell. .... .. . .... .... .. . ... 14 decalviiiis Gand. .... .... .... .... .... .. ... ... 46 marffinata Benth 30 mkranthu Benth 30 mirmphrlla Moq 47 ( Findl.) Druce 22 tomentosa R. Br 8 var. leptophylla Benth .... .. . .... . .. .... .... 47 rilhsa F. Mtiell. 24 Kenlrop.sis .... .... ... .. .... ... .... ... .... .... 4 Kochia 2, 14 sect, 'iustrokochia .... .... .... .. . .... .... .... .... .... 14 sect. Duriala 14 sect. Maireana .... .... .... .. .... .... .... ... .... 14 amocna DieLs 21 aphyUa R. Br. 54 appretsa Benth 54 astrotrichu L. A. S. Johns. .... .. . .... .... .... .. . .. 35 aikkisiaim Fit/g. 44 brachvptcru {¥ . Muell.) Benth .... .... .... .... ... ... 55 breyi/olia R. Br .. . .... .... .... .... .... .... .... 22 broKnii I-. Muell 54 cannomi Black .... .... .... .... .... .... .... .... .. . 42 carnosa (Moq.) Anders 30 cheedii Anders. 20 ciliata F. Muell 27 concavu I.sing .... .... .... .... .... .... .... .... .. 19 coronata Black . .. .... .... .... .... .... .... .... .... 28 crassiloha Anders. . .. .... .... .... . .. .... .... .... .... 24 decaptera F. Muell. 55 decipiens Gauba . .. .... .... .... .... .... .... .... .... 39 dichopiera F. Muell. .... .... .. . .... .... .... .... .... 28 ( 4)44205 81 enchylaenoides (Black) Black ehantha F. Muell erioclada (Benth.) Gauba excayata Black var- irichoptera Black fimhriolala F. Muell. neorgei Diels f;loiiu'rif' CONTENTS A Taxonomic Revision of the genus Maireana (Chenopodiaceae) By P. G. WUson Publication date of Volume 1 Number 5