MUELLER IA VOL. 2, No. I. MARCH, 1969. 2 5 FEB 1969 6442/68, MUELLERIA An Australian Journal of Bota VOL. 2, No. I. MARCH, 1969 2 5 FE& 1969 NATIONAL HERBARIUM ROYAL BOTANIC GARDENS, MELBOURNE VICTORIA, AUSTRALIA R. T. M. PESCOTT, Director and Government Botanist 6442/68. CONTENTS Page The genus Villarsia (Menyanthaceae) in Australia — Helen I. Aston . . . . . . . . . . . . . . 3 A review of the genera Teloschistes and Xanthoria in the lichen family Teloschistaceae in Australia — Rex B. Filson . . 65 The publication of this number has been made possible by a direct grant from the Botanic Gardens Branch Research Trust Fund which was set up in 1965, as the result of a generous donation to the Royal Botanic Gardens by Miss Maud M. Gibson of Switzerland, formerly of Melbourne. The trust in addition to providing a grant of $1,000 towards the cost of printing, also assumed financial responsibility for the provision of Research Assistants for the two authors. THE GENES VILLARSIA (MENYANTHACEAE) IN AUSTRALIA by Helen I. Aston* SUMMARY The species of Villarsia Vent, found within Australia are described and the distinctions and distribution of each given. Stress is laid on the taxonomic importance of seeds within the genus. The correct application of the name V . exaltata (Soland. ex Sims) G. Don is explained and this species is shown to be quite distinct from V . reniformis R. Br. with which it has long been erroneously synonymised. Two new species and one new variety are described for the first time — these are V. submersa, V. urnbricola var. umbricola, and V. umbricola var. beaugleholei. The genus within Australia is seen to consist of 9 species endemic to south-western Australia, and 3 species and one variety endemic to eastern Australia. It is also possible that some species now placed under Nymphoides Hill may prove to be better placed under Villarsia Vent. INTRODUCTION Villarsia Ventenat, Choix Plant. 9 (1803) has been conserved against Villarsia J. F. Gmelin, Syst. Nat. 2: 306, 447 (1791). The type species is V. ovata (Li.) Vent., now correctly known as V. capensis (Houtt.) Merrill (Merrill 1938, p. 360). Although this type species is South African, and the only South African species of Villarsia. the genus is chiefly an Australian one. Apart from perhaps three species found in Cambodia, Laos, and the Malayan Peninsula, all others are confined to south-western and south-eastern Australia. N. T. Burbidge (1963) gives 9 endemic species for this country but the results of the present work increase this number to 12 species plus one good variety. In most older works and in State floras Villarsia has been placed in the family Gentianaceae, and, where the family has been subdivided, in the tribe Menyantheae. Hutchinson (1960, p. 451) removes the Menyantheae from Gentianaceae and gives it family status as Menyan- thaceae. The Menyanthaceae are characterised by alternate leaves and valvate or induplicate-valvate corolla-lobes, while opposite leaves and contorted or rarely imbricate corolla-lobes are shown by the Gentianaceae. This distinction has gained acceptance, and Villarsia is placed in the Menyanthaceae in the recent floras of Eichler (1965, p. 260) and Beadle, Evans, and Carolin (1962, p. 353). In early literature there has been confusion between the generic names Villarsia, Swertia, Liparophyllum, Limnanthemum and Nymphoides, and distinctions between these have not always been clear or agreed upon. Within Australia Nymphoides Hill (syn. Limnanthemum S. G. Gmel.) is the only genus containing species which might be confused 6442/68.-2 National Herbarium of Victoria. 4 Helen I. Aston: The Genus Villarsia ( Menyanthacecie ) in Australia with Villarsia. Typically Nymphoides can be distinguished by its aquatic habit, its non-paniculate inflorescence with the pedicels arising from a short, weak peduncle (or the peduncle absent so that the pedicels appear in a cluster) which is often subtended by one or a cluster of several sessile or near-sessile and floating leaves, and also by its non-valvate, indehiscent or irregularly breaking fruits. Villarsia spp., by contrast, although frequently aquatic are best considered as wetland plants, and most typically have erect, paniculate inflorescences and valved, capsular, dehiscent fruits. Nymphoides has generally been retained to date for our northern and north-eastern tropical and semi-tropical species, including 2 well-known ones which penetrate into south-eastern Australia, but the distinctions between the two genera are not always clear to the present writer, and the fruiting characters in particular show variation. From material seen it appears possible that at a later date a few species now placed under Nymphoides may be more correctly transferred to the genus Villarsia. It seems unwise to take any action in this direction at present without adequate coverage of specimens and without held experience of the family in northern and north-eastern areas of Australia. It is obvious from collections housed at the National Herbarium of Victoria that the genus Nymphoides itself is also in need of revision. Australian species which have at some time been considered under Villarsia but which are now placed under Nymphoides are not mentioned in this paper. METHOD OF WORKING Results shown in the present paper have been obtained from a personal study of all Villarsia specimens housed in the Australian state herbaria and several others, together with extensive field observations in Victoria and South Australia. Where necessary photographs or seeds of type material were obtained from overseas herbaria for direct comparison here, or else selected specimens were forwarded overseas for comparison with types and critical comment. For most species however, syntype material was available in the collections of the state herbaria and in these cases where typification was clear no further efforts were made to locate other existing types. Because all type material will not have been located or seen by the writer, lectotypes have generally not been chosen. The field work undertaken in Victoria and South Australia was most essential, and in fact the full study originated from the writer’s discovery of two distinct species of Villarsia growing side by side in a swamp near Cranbourne and thus belying the accepted understanding that only one species existed in the State. The field provided many measurements and observations not obtainable from Herbarium specimens, particularly regarding the habit, flower span, other floral characters, maximum variations of size, and distribution. Representative collections were made in all areas visited, and additional locality records were kept. All species (and variety) from eastern Australia have been examined in the field, but all those from western Australia are known to me only from herbarium material. This necessarily means that the range of Helen T. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 5 measurements given will not be as broad for western species, as only comparatively meagre material is available and whole populations have not been seen. Floral characters in this genus are impossible to ascertain from herbarium specimens as the flowers are very fugitive and deliquescent. Those given for western species have therefore been taken from collector’s notes or early descriptions. Similarly plant height and habitat has mostly been ascertained from specimen labels or from prior descriptions. DIAGNOSTIC CHARACTERS Habit. — The size, and erect, reclining, robust, slender, herbaceous, or stoloniferous habit of plants, and the erect or floating foliage, can be very useful where plants appear typical under good growing conditions for the particular species concerned. Foliage. — Leaf size, ratio of length to breadth, outline, shape of the base, margin, texture, matt or glossy surfacing, degree of dorsiventrality, and prominence of venation can be diagnostic in themselves, but are not always so and must then be used in conjunction with other characters. Inflorescence.- V. capitata and V . congestiflora are immediately dis- tinguished from other species by their sessile flowers in clusters or heads. Amongst the remaining species, the degree of openness or compactness of the panicle, of slenderness and length of the pedicels, and whether the pedicels of mature capsules are erect or reclining are of some use. Flowers. — -The strongly heterostylous character (Fig. 27) of V. exaltata contrasts with the homostyly of other species. It is possible that V . parnassifolia also shows heterostyly, but this requires held checking. Fig. 1 — Variation in the corolla span of eastern species of Villarsia. - 45 - 40 _ 35 _ 30 - 25 _ 20 _ 15 _ 10 _ 5 L 0 6 Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia Calyx . — This may consist of lobes which are virtually free from the base or attached only in the very basal region, or which spring from the summit of a pronounced calyx-tube. The tube may be relatively inconspicuous in flower but may enlarge as the capsule matures, and remains adnate to the lower portion of the capsule wall. This explains Fig. 2 — Variation in the corolla-lobe apex of V. exaltata, V. reniformis, and V. umbricola. All shapes may be found within the one population. Fig. 3 — Capsules of Villarsia spp., all approx, x 2, contained within the persistent calyces, a, V. exaltata, Aston 1577. b, V. reniformis, Aston 1583. c, V. umbricola, MEL16407. d, V. parnassifolia, J. W. Green 868. e, V. tatifolia MEL16452. f, V. lasiosperma, MEL877. g, V. calthifolia, J. Hale, Dec. 1939. h, V. albiflora, MEL16437. i, V. violifolia, MEL16485. j, V. submersa, R. D. Royce 3778. k, V. congestiflora, MEL16482. 1, V. capitata , R. D. Royce 5174. Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 7 why the length of mature capsules may be greater than that of the calyx- lobes yet the capsule may still not project beyond the lobe tips and may be said to be shorter than them. Width of the calyx-lobes is not always helpful but these are broad in V. calthifolia and slender in V . latifolia and V . lasiosperma. The membranous lobe margins of the latter species are more conspicuous than those of other Villarsia species. Corolla. — Corolla size (Fig. 1) is important in eastern species, but has not been examinable for those of the west. The span is measured from the tip of one corolla-lobe diametrically across the centre of the corolla while pressing the lobes outwards horizontally from their points of connection at the insertion of the anthers. Corolla colour is constantly Fig. 4 — Seeds of Villarsia spp., x 15, with enlargements of surface projections, a, V. reniformis, Aston 1434. b, V. umhricola var. umbricola, Aston 1531. c, V. umhricola var. beaugleholei, Aston 1445. d, V. congestiflora, MEL874. e, V. exaltata, Aston 1432. f, V. lasiosperma, M ELI 6493. g, V. violifolia, MEL16491. h, V. parnassifolia, MEL16384. i, V. submersa, R. D. Royce 3778. j, V. calthifolia, MEL16478. k, V. albiflora, MEL 16443. 1, V. capitata, R. D. Royce 2237. m, V. latifolia, MEL16456. riHL 390 360 330 300 270 240 210 180 150 120 90 60 30 0 I. Aston: The Genus Villarsia ( Menycinthaceae ) in Australia s V. RENIFORMIS V. EXALTATA V. UMBRICOLA var. UMBRICOLA V. UMBRICOLA var. BEAUGLEHOLEI I i SEED SIZE Fig. 5— Graph of seed size in eastern species of Villarsia. NUMBER OF SEEDS NUMBER OF SEEDS Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 9 V. SUBMERSA SEED SIZE SEED SIZE Fig. 6— Graph of seed size in western species of Villarsia. 0 10 Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia white or yellow for each species. The presence of a longitudinal keel or wing on the inner surface of each corolla-lobe is peculiar to V . lasiosperma. The shape of the corolla lobes (Fig. 2) in all eastern species is quite variable and of no diagnostic value. The lobe apex may alter from acute to obtuse to truncate to emarginate even within the one population. Ovules. — The number of ovules per placenta has not been well-investi- gated, but it appears that this character would have very little diagnostic significance. Capsule. — The shape, mode of dehiscence, thickness of the wall, degree of adherence to the calyx-tube, and length in comparison with the calyx are all useful features (Fig. 3). Measurements refer to the ripe, open capsule. Seed. — Seeds (Fig. 4) are of primary importance diagnostically. Size (Figs. 5, 6), shape, surface patterning, presence or absence of tubercles, types of tubercles, and presence or absence of a conspicuous caruncle are all valuable features. The term “jig-saw” (Fig. 7) has been coined as most appropriate for the surface patterning wherein each minute segment sends projections into those alongside as with the pieces of a jig-saw puzzle. The segments are usually distinctly outlined, but at times appear less-defined and resemble hills and spurs separated by reticulating gullies. Length was measured across the widest diameter of each seed, this usually commencing somewhat away from the point of attachment. Fig. 7— Diagram of the “jig-saw” surface pattern of seeds of some Villarsia spp. Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 11 CITATION OF SPECIMENS Except for several inadequate specimens all those examined in the herbarium are cited under their specific descriptions. For the sake of brevity, localities have often not been given in complete detail. For collections made by myself only the sheets retained at MEL are listed (except for types), but it is to be understood that all duplicates forwarded to other herbaria have also been examined. Herbarium abbreviations used are: — AD — State Herbarium of South Australia, Adelaide; ADW — Waite Agricultural Research Institute, Adelaide; BM — British Museum (Natural History), London; BRI- Botanic Museum and Herbarium, Brisbane; CANB— Division of Plant Industry, C.S.I.R.O., Canberra; CAN BLR — Land Research and Regional Survey, C.S.I.R.O., Canberra; CBG Canberra Botanic Gardens, Canberra; FI — Herbarium Universi- tatis Florentinae, Florence; HO — Botany Department, University of Tasmania, Hobart; K— Royal Botanic Gardens, Kew, England; L- Rijksherbarium, Leiden, Holland; MEL— National Herbarium of Vic- toria, Melbourne; MELU— Botany Department, University of Mel- bourne; NSW — National Herbarium of New South Wales, Sydney; PERTH— Western Australian Herbarium, Perth; PERTHU -Botany Department, University of Western Australia, Perth; SYD— Department of Botany, University of Sydney. The symbol (Sd) placed before a collection indicates the presence of seed. KEY TO THE AUSTRALIAN SPECIES OF VILLARSIA L Flowers sessile or near-sessile in few-flowered clusters or in dense heads, corolla yellow. Western species. 2. Calyx-tube and sometimes also the calyx-lobes covered sparsely or densely with long fine hairs, capsule thinly membranous, apparently breaking or dehiscing irregularly, seed smooth and shining, non-carunculate \ y capitata 2. Calyx glabrous, capsule thickly chartaceous, splitting into 4 recurved valves, seed strongly granular and dull, noticeably carunculate 2. V . congestiflora 6442/68.-3 12 Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia I. Flowers occurring singly on pedicels 4-70 mm. long in open panicles, corolla yellow or white. Western or eastern species. 3. Corolla white. Western species. 4. Extremely slender, tine-branched, totally submerged aquatic; radical leaves dz elliptic, to 6 cm. long, twice to three times as long as broad, dorsiventral; each corolla lobe without a longitudinal keel on the inner surface; seed small, 0 6-0 8 mm. long, smooth 3. V. submersa 4. Erect, tufted, reasonably stout, emergent plants with culms to 3 (-5) feet high. 5. Radical leaves lanceolate to ovate, non-cordate, to 9 5 cm. long, dz twice as long as broad, thick-textured to semi- succulent, not markedly dorsi-ventral, both surfaces matt (or the upper ? sometimes somewhat glossy); each corolla lobe with a longitudinal keel on the inner surface; seed large, 1 4—2 mm. long, the surface with moderately-dense. fine, tapered, spinescent tubercles to 0 2 mm. long 4. V. lasiosperma 5. Radical leaves zb sub-orbicular, cordate, to 7 cm. long, slightly broader than long or dimensions about equal, strongly dorsiventral, upper surface dz glossy-green, lower surface paler and dotted; each corolla lobe without a longitudinal keel on the inner surface; seed medium, 0 8—1 2 mm. long, densely covered in short, rigid, hollow tubercles ca. 0 5 mm. long (or rarely these reduced to granules) 5. V. al biflora 3. Corolla yellow. Western or eastern species. 6. Small, low, stoloniferous, mat-forming perennial with slender culms to 15 cm. high; radical leaves small, to 3 cm. long, thin- textured; capsule not circumscissile, opening from the apex in 4 acute valves; seed 1-4-1 *8 mm. long, smooth, glossy or matt, with a conspicuous, almost-basal, translucent caruncle. Western species 6. V. violifolia 6. Large, erect, robust, non-stoloniferous perennial, with culms to 7-8 feet high; radical leaves large, to 20 ( ?— 30) cm. long and broad, almost round, deeply cordate, the margin strongly and irregularly toothed, the teeth callose-tipped; capsule circumscissile near the apex, then opening by an apical pore or splitting into truncate-tipped valves; seed E5-18 mm. long, densely covered with short, hollow, tapered tubercles, without a conspicuous caruncle. Western species 7. V. calthifolia 6. Medium to large, erect or semi-erect, tufted perennials. Western or eastern species. Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 13 7. Western species. 8. Radical leaves thin-textured, =t glossy, the veins prominent beneath, edges crenate-dentate; capsule 2-5 mm. long, shorter than the calyx, the lower adnate to it, globose or sub-globose, thick-walled, opening by 4 distinct, recurved valves; seed large, 1 5—1 9 mm. long, smooth and usually glossy, with a prominent, ± circular, translucent caruncle 8. V. latifolia 8. Radical leaves thick-textured, apparently semi- succulent, matt-surfaced, the veins rather incon- spicuous, edges sub-entire to crenate-dentate; capsule 4-10 mm. long, usually 1-2 mm. longer than the calyx and adnate to it only at the base, narrow-lanceolate to narrow-ovate in outline and usually appearing slender, thin-walled, opening by valves only near the apex; seed small, 0 5-1 mm. long, faintly granular, without a conspicuous caruncle 9. V. parnassifolia 7. Eastern species. 9. Radical leaves erect, ovate-elliptic to broad-ovate, usually much longer than broad, base rounded to only shallowly cordate, thick-textured, matt and ± uniform on both surfaces; flowers heterostylous; capsule with the lov/er to \ adnate to the calyx-tube; seed large, ( 1 5-) 1 1-2-6 (-3) mm. long, light cream-fawn to dark brown-grey-black, mostly sparsely to densely tuber- culate (tubercles to 0 3 mm. long), conspicuously carunculate 10. V exaltata 9. Radical leaves erect or floating, ovate to reniform, mostly about as broad as long or either dimension somewhat greater than the other, base usually slightly to deeply cordate, matt-surfaced to glossy above, often strongly dorsiventral; flowers homostylous; capsule adnate to the calyx-tube only at the very base; seed smaller, (0 7-) 1 — 15 (-19) mm. long, pale never blackish, smooth to slightly granular (but tuberculate in V. umbricola var. beaugleholei q.v.), without a con- spicuous caruncle. 10. Plants stoloniferous when in water, radical leaves then floating flat, strongly dorsiventral, glossy and deep green above, pale and red-dotted" beneath; culms db erect and self-supporting; inflorescence chiefly terminal on the culm; flower span 19-43 (average 32-3) mm.; mature capsules on erect pedicels 1 1 . V . reniformis 14 Helen I. Aston: The Genus Villarsia (Menyanthaceae) in Australia 10. Plants non-stoloniferous; radical leaves on plants in water either erect or floating, but then not lying fully flat, surfaces not strongly dorsiventral, some- what yellowish-green; culms slender, weak, semi- erect or reclining against supports; inflorescence lax and spreading from much of the culm; flower span 11-31 (average 21 7) mm.; mature capsules on recurved pedicels. 11. Seeds almost smooth to granular 12. V. umbricola var. umbricola 1 1 . Seeds densely tuberculate, the tubercles to 0-75 mm. long 12. V. umbricola var beaugleholei TAXONOMIC TREATMENT Villarsia Vent. Choix Plant. 9 (1803) nom. cons. Type species not examined. Aquatic or dampland, glabrous, tufted, small to tall and robust, sometimes stoloniferous, perennial or annual herbs. Roots fibrous, the perennial species also mostly having a thickened, branched, com- pact rootstock. Radical leaves well-developed, erect or floating, on petioles winged and semi-sheathing towards the base, the blades simple, from mildly to strongly dorsiventral and often dotted beneath, variously elliptic, ovate, rotund, or reniform, from rounded to very strongly cordate at the base, with margins entire to crenate-dentate, texture thin to thick and semi-succulent. Culms 1 -several, erect or semi- reclining, from a few centimetres to 5 feet (or more) high, bearing mostly an open, paniculate inflorescence, or sometimes the flowers in clusters or heads; cauline leaves sometimes absent and represented by bracts only, but mostly the lower 1-few panicle branches are subtended by petiolate, radical-like leaves with those of the higher branches pro- gressively reducing to form narrower, sessile, semi-clasping bracts, the final bracts subtending each panicle branch or pedicel being 0 5 cm. or less long, elliptic and stem-clasping. Flowers homo- or hetero-stylous, mostly well-pedicellate, sometimes sessile in clusters or heads. Calyx persistent, 5-lobed, the lobes mostly distinct almost to the base in flower, and remaining so in fruit or the calyx-tube then enlarging to equal perhaps half the length of the fruiting calyx. Corolla yellow or white, very deliquescent, opening for 1 day only, conspicuously fimbriate at the throat, up to 4 cm. in diameter, gamopetalous, 5- (sometimes 4- or 6-) lobed, the lobes spreading, each with broad margins and sometimes with a longitudinal, vertical keel on the inner surface, induplicate- valvate in bud, the tube broad and approximately one third the corolla length (at least in eastern species). Ovary superior to semi-inferior. Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 15 unilocular with 2 (abnormally 3) parietal placentas, surrounded by 5 small, often orange, nectary glands at the base; style single, terminal; stigmas 2 (abnormally 3), flattened, papillate, exserted. Filaments exceedingly short or almost lacking, inserted on the corolla tube at the junction of the lobes; anthers 2-cefled, linear or linear-lanceolate, yellow (at least in eastern species), opening introrsely. Fruit a few- to many- seeded capsule splitting at the summit into 4 (or secondarily more) somewhat recurved valves, or sometimes breaking irregularly; capsule adnate below to the calyx-tube. Seeds smooth to tuberculate according to species. 1. Villarsia capitata Nees in Lehm. Plant. Preiss. 1 : 365 (1845). V. involucrata Hook. Icon. Plant, t.725 (1848). Rather small, erect, tufted, non-stoloniferous plant, with flowers clustered in dense heads. Apparently annual. Radical leaves erect, on petioles 3-11 (-20) cm. long; blades from < 1-4 cm. broad and long, almost rotund to reniform, usually broader than long, apex obtuse c~ mostly rounded, base truncate to cordate with the sinus broad and shallow to narrow and deep (ca. length of blade), edge entire to crenate or crenate-dentate, texture thin, under surface sometimes dotted. C ulms several, erect, 7-29 cm. high, branching dichotomously from low down; cauline leaves radical-like at the lower 1-4 nodes, on petioles to 5-6 cm. long, becoming more obtuse at the base of the blade and bract-like higher on the culm, the upper petioles short and sometimes broadly-winged, final bracts sessile. Inflorescence a dense, compact head of sessile flowers subtended by 2 broad, sessile, leafy, involucral bracts, or the bracts not always developed; heads 1—1 5 (-2) cm. wide, terminal on the main culm branch and short side branches, appearing sparsely or densely woolly from the hairs of the calyx; involucral bracts and peduncles glabrous to villous. Flowers homostylous; calyx-lobes 3 (-5) mm. long, narrow-to-broad-lanceolate; calyx-tube covered sparsely or densely with long fine hairs, these sometimes arising also from the outside of the calyx-lobes; corolla yellow, described by Nees as shorter than the calyx, without longitudinal keels on the lobes. Capsule small, ellipsoid, shorter than to almost equal to the calyx, the lower adnate to the calyx-tube, thinly membranous, apparently irregularly breaking or dehiscing. Seed small, (0 7-) 0 8—1 1 (-1 2) mm. long, straw-yellow through brown to finally black, broad elliptic, double-convex, smooth and shining, with very fine, reticulated ridging and pitting, without a conspicuous caruncle. Fig 8 — V. capitata, shape of radical leaves, x §. a, A. Morrison, 4.12.1900 b, C. A. Gardner, 27.9.1944. c, E. Salisbury, Aug 1949. d, e, MEL16474. 16 Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia Distribution : South-west Western Australia, in the vicinity of the Greenough and Irwin Rivers, and from Perth to Albany. Habitat: Swampy soils. Recorded on sand and clay. Flowering time: FI. and Fr. recorded Aug. -Dec. Typification : The following syntypes have been seen — (Sd) Preiss No. 1956 (MEL 16470-1 and NSW 90572), syntype of V . capitata, cited by Nees as “In paludosis ad flumen Canning, Perth, Novembri a. 1841. Herb. Preiss. No. 1956”; (Sd) Drummond No. 7 ( MEL 1 6468-9) , syntype of V. involucrata, cited by Hook as “Swan River, Western Australia. James Drummond, n. 7, 1845”. Discussion : The capitate heads of this species distinguish it from all others in the genus except V . congestiflora , with which it could readily be con- fused at first sight. It differs from that species however in the denser heads, woolly calyx and other inflorescence parts, non-valvate capsule, and smooth, shining, non-carunculate seeds. Fig. 9 — V. capitata, distribution map. Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 17 Sphcimens examined (excluding types) : Western Australia— (Sd) W.A.. (MEL16473) “wet sand"; (Sd) S.W.Austr., Mr. Clarke (MEL16472); Serpentine R.. W.A., Oldfield (M ELI 6475) “fl. yellow”; King George Sound. J. R. Muir (MEL16474); (Sd) Irwin's and Greenough's River. F. Mueller. Nov. 1877 (MEL21956); Cannington. R. Helms, 12.11.1898 (PERTH, and NSW77138); Midland Junction, Swan River. Alex Morrison, 16.11.1898 (MEL26380; NSW90569) “swamps"; (Sd) Swan district, E. Pritzel, Dec. 1900 (NSW570) “locis subhumidis"; (Sd) Avon district. Byfields Mill, Diels and Pritzel, Dec. 1900 (PERTH) ; (Sd) Midland Junction. Swan River. Alex Morrison. 4.12.1900 (PERTH) ; Kelmscott, Canning R.. Alex Morrison 10326. 14.12.1900 (PERTH) “Swamp": W.A.. A. G. Hamilton 518, 1902 (NSW90513); (Sd) Busselton, F. Stoward 224 , Nov. 1912 (NSW 90571); Perth. Mr. Carlour, 14.11.1917 (PERTH); Busselton, C. A. Gardner, 27.9.1944 (PERTH); (Sd) Coolup. R. D. Royce 2237, 13.10.1947 (PERTH) “Firs, yellow. Swampy clay soil”; (Sd) W.A., E. Salisbury, Aug. 1949 (PERTH); Lower Hill River. C. A. Gardner 10243, 10.10.1951 (PERTH); (Sd) Busselton. R. D. Royce 5174, 3.11.1955 (PERTH) “Firs, yellow, capitate, calyx sparsely hairy' 5 . 2 Villarsia congestiflora F. Muell. Fragm. Phyt. Aust. 6 : 141 (March 1868). Rather small, erect, tufted, non-stoloniferous plant, with flowers clustered, a few together or in dense heads. Apparently annual. Radical leaves erect, on petioles to 1 1 cm. long, these sometimes strongly red- dotted; blades 1-4 cm. long and broad, broad-ovate to zh rotund to ± reniform, sometimes broader than long, apex obtuse to rounded, base truncate to cordate, edge entire to crenate or toothed, texture thin, under surface sometimes strongly red-dotted. Culms few to several, erect, 10-40 cm. high, sometimes strongly red-dotted in basal regions, branch- ing from near the base; cauline leaves 2-4 radical-like, the lower on petioles to 8 cm. long, the bracts above these becoming broad, sessile, amplexicaule, and mostly somewhat toothed. Inflorescence glabrous, consisting of spaced clusters of few to several sessile or near-sessile Fig. 10 — V. congestiflora , shape of radical leaves, x f. a, b, MEL874. c d, MEL16483. flowers forming bundles or congested heads terminating the main branches or very-reduced side-branches, the clusters subtended by broad, entire or toothed involucral bracts. Flowers homostylous; calyx-lobes 4-5 mm. long, lanceolate, acute, glabrous; calyx-tube glabrous; corolla yellow, throat fimbriate, the lobes without any longitudinal wing or keel on the inner surfaces. Capsule small, 4-5 mm. long, rotund-ellipsoid, 1-2 mm. shorter than the calyx, the lower adnate" to the calyx-tube, thickly chartaceous, splitting into 4 recurved valves. Seed 1-1*4 mm. 18 Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia long, straw-yellow at first to black at maturity, broad-elliptic to ± circular in outline, turgid but compressed, dull, the surface densely and strongly granular, noticeably carunculate with pale cells at the base. Distribution : West coast of Western Australia, from Perth north to the Murchison River. Habitat: “Moist sand.” Flowering time: Fr. recorded in Nov. Typification : Syntypes seen— (Sd) Nr. Perth, Swan River, Oldfield (MEL872-3), and (Sd) Murchison River, Oldfield (MEL874). Discussion : The clustered flowers sometimes in dense heads distinguish this species from all others in the genus except V. capitata q.v. Specimen examined (excluding types) : Western Australia — (Sd) G reenough’s and Irwin’s River, F. Mueller, Nov. 1877 (MEL 16482-4) “moist sand”. Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 19 3. Villarsia subniersa H. I. Aston sp. nov. Herha gracillima, omnino submersa (praeter flores), caespitosa, radicata, perennis. Foliorum radicalium laminae ellipticae usque ad ovato-lanceolatas, 2-3-6 cm. x 1-2-6 (-3), quam latae bis terque longiores, dorsiventrales, petiolis ad 40 cm. longis praeditae. Culmus usque ad 60 cm. longus. Inflorescentia laxe paniculata, multiramosa, ramulis vel pedicellis 1-5 ad nodos simul emergenti- bus; pedicelli 20-70 mm. longi. Flores parvi; calycis lobi 3. 5-4. 5 mm. longi; corolla alba, quam calyx circiter bis longior, intus lanata vel pilosa. Capsula 3-3-5 mm. longa, quam lobi calycis circiter 1 mm. brevior, tenue membranacea, ellipsoidea, irregu- lariter dehiscens ut videtur. Semen parvum, 0-6-0-75 (-0-8) mm. longum, glabrum, ellipsoideum, aliquanto lateraliter compressum. stramineum ad fulvum. A speciebus omnibus aliis Villarsiae recedit ob habitum submersum gracillimumque et ob folia elliptica (quam lata multo longiora). Eius semina ad haec V. parnassifoliae (Labill.) R. Br. proxime accedunt. Extremely slender, fine-branched, tufted, apparently non-stoloni- ferous, aquatic perennial, totally submerged except for the flowers which float on the water surface. Radical leaves on very slender petioles to 40 cm. long, expanded at the base; blades elliptic to lanceolate-ovate, 2 3-6 cm. long x 1-2 6 (-3) cm. broad, twice to three times as long as broad, broadly obtuse and mostly slightly emarginate at the apex, rounded to semi-truncate at the base, entire-edged, dorsiventral, upper surface smooth, lower paler and finely-dotted. Culms several, to 2 feet long, extremely slender, without cauline leaves or large bracts. Inflores- cence a much-branched, open, very fine-stemmed "panicle, with 1-5 branches or pedicels arising together at the nodes, each subtended by a small, elliptic-ovate-lanceolate, obtuse, sessile bract 2-5 mm. long; pedicels of mature capsules 20-70 mm. long, capillary. Flowers small! homostylous; calyx-lobes 3-5-4-5 mm. long, elliptic to ovate-elliptic, obtuse or slightly emarginate; corolla white, apparently about twice as long as the calyx, with the lobes apparently having entire margins, lacking a longitudinal keel, and woolly or hairy with fine fimbriae over the whole of the inner surface; ovary unilocular with 2 parietal placentas. Fig. 12— V. submersa, shape of radical leaves, x f. a, b, usual shapes c d extremes. All from R. D. Royce 3778. 20 Helen I. Aston: The Genus Villarsia (Menyanthaceae) in Australia ovules ca. 12-15 per placenta, style short and thick, stigma bilobed; anthers linear, bilocular, shortly stipitate, 0 5-0 8 mm. long when dry, 0 9-1 mm. long when boiled. Capsule 3-3 5 mm. long, ca. 1 mm. shorter than the calyx-lobes, thinly membranous, ellipsoid, adnate at the base to the calyx, apparently irregularly dehiscing. Seed small, 0 6-0 75 (-0 8) mm. long, pale straw-yellow to brown-yellow, ellipsoid, turgid but laterally-compressed, without a conspicuous caruncle, surface smooth and somewhat shining with minute “jig-saw” patterns. Distribution : Known only from Boyanup, near Bunbury, south-west West Aus- tralia, and from King George Sound. Habitat: Rooted in mud in 18 inches of water. Flowering time: Buds, flowers, capsules and seeds present in early September. Typification : Holotype — (Sd) Boyanup, R. D. Royce 3778, 9.9.1951 ( PHRTH) “Firs, white, petals woolly; rooted in mud, leaves and panicle branches wholly submerged. Water 18 in. deep.” Isotype — MEL26022. Paratype -Boyanup, R^D. Royce 4343, 16.9.1953 (PERTH; K). Fig. 13 — V. submersa, distribution map. Helen 1. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 21 Discussion : With its entirely submerged habit (except for the floating flowers), very slender to capillary panicle-branches and pedicels, elliptic, quite non-cordate leaves, small white flowers, small and ? irregularly dehiscent capsules, and small smooth seeds, this species is quite distinct from all others of the genus. In seed characters it appears closest to V . parnas- si folia. Specimens examined (excluding types) : Western Australia — W. A., Drummond No. 17 (MEL27193): King Georce Sound. J. R. Muir (MEL27192). 4. Villarsia lasiosperma F. Muell. Fragm. Phvt. Aust. 6 : 137 (March 1868). Tall, erect, tufted, apparently non-stoloniferous perennial, dotted on the lower culm, petioles, and under-surfaces of leaves. Radical leaves erect, on petioles 2-20 cm. long; blades lanceolate to narrow- to broad- elliptic to ovate, longer than broad, 2 5-9 5 cm. long x 1 5-6 cm. broad, apex acute to obtuse, base acute to obtuse to zh truncate, edge entire and sometimes undulate, texture mostly thick and semi-succulent, both surfaces matt (or ? upper sometimes somewhat glossy), venation mostly rather obscure above, less so beneath. Culms 21 in.-3 ft. (-5 ft.) high, standing erect above the leaves, branching commencing from \ height of culm; cauline bracts sometimes with the lower leaf-like on petiole 2-6 cm. long, or else all bract-like with the lowest either sessile or shortly petiolate. Inflorescence an open, diffuse, dichotomous panicle; final bracts 2-3 mm. long, lanceolate, acute; pedicels of mature capsules 8-24 mm. long, slender. Flowers homostylous; calyx-lobes 4-7 mm. long, linear- to elliptic-lanceolate, narrow, glabrous or with fine hairs marginal or on the inside surface at the apex, the margins conspicuously membranous; corolla white, apparently about twice as^long as the calyx, fimbriate at the throat, each lobe with a longitudinal keel or wing on its inner surface. Capsule 4-6 mm. long, equal to or a little < calyx-lobes, globose-ovoid, thickly chartaceous, free from the calyx or attached only at the very base, splitting deeply into 4 acute, recurved valves. Seed large, 1 4—1 9 (-2) mm. long, pale straw-yellow in the immature seeds seen but probably darker on ripening (Mueller says “lividis”), ellipsoid, apparently turgid and laterally-compressed when mature, surface with minute “jig-saw” patterns and bearing conspicuous, moderately dense, long, fine, tapered, acute, spinescent tubercules to 0 2 mm. long, without a conspicuous caruncle. Distribution : South-west Western Australia, from Busselton to Albany. Habitat: “Heath swamps”, “Forest swamps”, “Edge of swampy depression”. 22 Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia Fig. 14 — V. lasiosperma, shape of radical leaves, x §. From MEL876. Flowering time: FI. and Fr. recorded Oct. -Dec. Typification : Syntypes seen — (Sd) Swamps at King George Sound, Oct. 1867; Albany; Swamps in the rear of Mt. Melville (various labels on sheet), coll. F. Mueller (MEL16493): Heath swamps north of Albany, F. Mueller , Oct. 1867 (MEL16494): Swamps at K.G. Sound, Oct. 1867; Forest swamps north of Albany (various labels on sheets), coll. F. Mueller (MEL 16495; MEL875). Discussion : V. lasiosperma seems quite closely related to V. exaltata (q.v. dis- cussion) of the eastern States, and the two species resemble each other in size, habit, and foliage, and some seed and flower characters. Flowever V. lasiosperma is readily distinguished by its white, homostylous flowers with slender calyx-lobes and with the corolla-lobes bearing longitudinal keels, by the capsules not or very scarcely adnate to the calyx, and by its non-carunculate seeds with fine, tapered tubercles. It should not be confused with any western species. Specimens examined (excluding types) : Western Australia — (Sd) King George Sound, G. Maxwell (MEL877); Blackwood, W.A., Mrs. McHard (MEL876); (Sd) Chokarup, F. Mueller , 20.12.1877 (MEL 16496): Near Wilson’s Inlet, also at Chokarup, F. Mueller, 22.12.1877 (MEL 16497) “Leaves thick, not . . Busselton, C. A. Gardner, 27.9.1944 (PERTH); (Sd) 2 miles south of Northcliffe, T. E. H. Aplin 1406, 12.12.1961 (PERTH) “Edge of swampy depression”. Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 23 Fig. 15 — V. lasiosperma, distribution map. 5. Villarsia albiflora F. Muell. Fragm. Phyt. Aust. 2:21 (Feb. 1860). Large, tufted, apparently non-stoloniferous perennial, with erect culms and floating leaves, the submerged portions usually conspicuously dotted. Radical leaves on petioles (2-) 4-23 (-28) cm. long; blades (1-) 2 5-6 (-7) cm. long x (1-) 2-6 (-7 5) cm. broad, broad-ovate to sub-orbicular to reniform, mostly slightly broader than long or dimen- sions about equal, apex obtuse to rounded, base cordate with broad rounded lobes sometimes overlapping, sinus narrow and deep (to -3 length of blade) to broad and shallow, edge entire to slightly crenate or sometimes markedly crenate or toothed, upper surface deeper green and ? glossy, lower surface paler and much-dotted. Culms erect, 8 inches to 4 feet high, stout in larger specimens, branching commencing from %-f height of culm; cauline leaves usually 1-3 with blades radical-like, on petioles to 9 cm. long. Inflorescence an open, many-flowered, dicho- tomous panicle; final bracts 2-4 mm. long, lanceolate, acute; pedicels of mature capsules (4-) 10-20 (-30) mm. long, slender. Flowers homo- stylous; calyx-lobes 4-7 mm. long, lanceolate to elliptic, acute; corolla white, apparently about twice as long as the calyx, fimbriate at the throat, the lobes without a longitudinal keel. Capsule 5-8 mm. long, from sometimes equal to the calyx lobes to usually 1 or 2 mm. longer than them, hard chartaceous, adnate to the calyx only at the base, splitting deeply into 4 valves, the valves erect or tips somewhat recurved. Seed 0 8—1 2 mm. long, straw-yellow to yellow-brown (cream-white 24 Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia when immature), ellipsoid, turgid, slightly compressed laterally, the surface with minute “jig-saw” patterns and also densely covered in short, rigid, hollow tubercles ca. 0-5 mm. long (or rarely these reduced to low granules giving a comparatively smooth seed), without a conspicuous caruncle. Fig. 16 — V. albiflora, shape of radical leaves, x §. a, MEL16430. b, NSW77139. c, H. W. Alcock. Sept. 1896. d, MEL16439. e, MEL16444. Distribution : South-west Western Australia, from about 80 miles north of Perth south through Perth and York to the Cape Leeuwin area, and also found in the Porongorups. Habitat: Fresh-water swamps, lake edges, streams, etc., in water to at least 1 foot deep and apparently also in boggy places. Recorded on laterite, in Jarrah forests, and among stones of a river bed. Associated with Melaleuca laterita (in one report). Flowering time: FI. and Fr. recorded Sept. -Jan. Typification : Mueller in his original description cites “In stagnis prope oppida Perth et Hampden. Clarke et Oldfield.” The National Herbarium of Victoria holds two sheets (MEL16430 and 16447) labelled “Hampden, W.A., Oldf.” and “Perth, W.Aust., Oldf.” respectively in Oldfield’s hand, both labels being annotated as V. albiflora by Mueller himself. These are regarded as syntypes. Both specimens show vegetative parts. Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 25 Fig. 17 — V. al biflora, distribution map. buds and flowers only, and as seeds are of such importance within the genus Kew Herbarium was approached to ascertain if further syntypes with seed or even young capsules were available there. A sheet bearing two specimens and labelled (amongst others) “fl. white. Monger’s Lake, Nr. Perth, W.Aust.” in Oldfield’s hand, with the additional "annotation “albiflora F. Muell.” again in Oldfield’s hand was located, but it bears no handwriting of Mueller. A photograph (Kew negative No. 8566) of the sheet has been compared with the Melbourne syntypes and is in agreement both concerning the specimens and the label handwriting. It must be regarded as possible syntype material, as Mueller did not always annotate sheets seen by him. However, it is most likely that Mueller did not see the Kew sheet at all, the material on it being sent directly to J. D. Hooker by Oldfield himself. As Mueller’s original description stated “Capsula adhuc ignota” (capsule so far unknown) it was expected that any material held by Kew would lack capsules and seeds, and Sir George Taylor in a letter dated July 6, 1966, confirmed this. Referring to "the Kew sheet mentioned above he stated “These specimens are evidently in the same state as your own with buds and only the first flowers opening. There are no capsules not even young ones developed, . . Fortunately Mueller, subsequent to his original diagnosis, made two further descriptive references to Villarsia albiflora, including references to the seed of the species and citing further material examined. In Fragm. Phyt. Aust. 26 Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 5 : 46 (1865) he cited “Haec planta ad fiumen Blackwood’s River occurrit”, and in l.c. 6 : 138 (1868) he stated “Hue planta Drummondi 4”. The National Herbarium of Victoria holds a sheet (MEL16434) collected by Oldfield at the Blackwood River and also a Drummond No. 4 specimen (MEL16435), both annotated by Mueller as V. albiflora and both with seeds agreeing with Mueller’s description. These sheets are here regarded as homoeotypes. With his original diagnosis, Mueller synonymised “K. reniformis, Nees in Lehm. pi. Preiss. 1.364, non R. Br.”. At this reference, Nees placed Herb. Preiss No. 1958 under V. reniformis R. Br. Sir George Taylor in a letter from Kew dated July 6, 1966, suggested that it could be argued that Mueller was basing his species partly on V . reniformis Nees in PI. Preiss. 1 : 364 (non R. Br.), and forwarded a Kew sheet of Preiss 1958 to Melbourne for examination. This sheet does not bear Preiss’ number but is labelled “. . . in stagnantibus prope Perth 1839.10.25” and “Swan River Preiss” in agreement with data given by Nees and has the identification “reniformis R. Br.” subsequently altered by Bentham to “V. albiflora F. Muell.”. It contains seed and both it and two other specimens of Preiss 1958 held at Melbourne (MEL16431 and 16433) also with seed, agree with the homoeotype material cited above. I am quite in agreement that the three sheets of Preiss 1958, and the syntype and homoeotype material cited are all conspecific, but cannot see why Preiss 1958 should be regarded as type material. If Nees had actually published V. reniformis Nees as his own name then certainly the Preiss collection would be a type of V. albiflora F. Muell. However, Nees had no intention of using a new name or describing a new species, but simply mistook Preiss 1958 as belonging to V. reniformis R. Br., and placed it under that name. The name V. reniformis Nees has there- fore never been published, has no standing, and cannot have any type. Mueller may not have intended his mention of it to be taken as a direct synonymy, but only to indicate that the entity Nees circumscribed belonged under V. albiflora F. Muell. rather than under V . reniformis R. Br. This view is now accepted by Sir George Taylor and is held also by Mr. J. H. Willis of Melbourne. Discussion : This species is very similar in foliage to V . reniformis and in seed to V. umbricola var. beaugleholei , but both of these are confined to the eastern States. V . albiflora is distinguishable however by its combination of densely-tuberculate, rough seeds, white corollas without any longi- tudinal wings on the inner surfaces of the lobes, and its cordate, dorsi- ventral foliage with blade length =t equal breadth. Two collections show variation from the typical, tuberculate seed. Oulina (Helena R.), Max Koch, 1907 (NSW90554) has two specimens, one with typical seed, the other with the tubercles reduced and in places absent, being represented then only by small granular swellings on the seed surface. Morley Park, N. G. Marchant f91, 1958 (PERTH) has no tubercles but only a granular surface. It is otherwise a good example of V. albiflora. Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 27 Specimens examined (excluding types) : Western Australia— (Sd) “In stagnantibus prope Perth”, Preiss 1958, Oct. 1839 (KEW); (Sd) Preiss No. 1958 (MEL16431 and 16433); Blackwood River, Walcott, Jan. 1868 (MEL16432) “Growing in swampy ground near Fennels station”; (Sd) Between Perth and York, F. Mueller, Nov. 1877 ( MEL 16437-8) “Swamps”, also [in Latin] Corolla white, lobe margins naked, throat fimbriate, lobes not longitudinally crested; (Sd) Swan River, F. Mueller , Nov. 1877 (MEL16439) “Jarrah forests”; (Sd) Swan River, Darling Ranges. F. Mueller, Nov. 1877 (MEL16436, 16444-5, 16448) “Vallies”, “Rivulets”; Serpentine River, F. Mueller, 1.12.1877 (MEL 16449) Corolla white, lobes ovate net crested, throat fimbriate [in Latin], “Cataracts, brooks . . .”; (Sd) Harvey’s River, F. Mueller, 5.12.1877 (MEL16443) Corolla lobes white . . . not crested. Leaves paler green beneath and less shining [in Latin]; (Sd) Blackwood River. W.A., Mrs. McHard, Dec. 1884 (MEL16441); (Sd) Swan District, H. W. Alcock, Sept. 1896 (PERTH); Bunbury, R. Helms, 27.9.1897 (PERTH); (Sd) Bunbury, Geographe Bay, R. Helms, 27.9.1897 (NSW77139); Cannington, R. Helms, Sept. 1898 (NSW90553) ; (Sd) Bayswater, A. Morrison 9429, 6.12.1899 (PERTH); Bayswater, low'er Swan River, A. Morrison 10330, 10.10.1900 (NSW90552); (Sd) Bayswater, Swan District, Diels & Pritzel 503, Nov. 1900 (PERTH); (Sd) Darling Range, J. B. Cleland, 190? (NSW90550); (Sd) Cannington, Swan District, Miss Lambert, Nov. 1901 (PERTH); (Sd) Mandurah, A. G. Hamilton 551, 1902 (NSW90555); (Sd) Near Perth, W. V. Fitzgerald, Oct. 1902 (NSW90551 ) “FI. white”; (Sd) Oulina (Helena River), Max Koch, 26.11.1907 (NSW90554) ; (Sd) York, J. B. Cleland, Dec. 1907 (NSW549); (Sd) Harvey, F. H. Steward, Nov. 1916 (PERTH); Collie River, Muja, C. A. Gardner, 8.4.1922 (PERTH); (Sd) Scott River, R. D. Royce 61, 17.1.1945 (PERTH) “Flowers white, leaves thick. Among stones in bed of river”; (Sd) Gnangarra, C. A. Gardner, Oct. 1945 (PERTH) “fls. white” “on shores of lake”; (Sd) York, northern turn-off, “The Lakes”, D. Churchill, Dec. 1957 (PERTHU) “in swamp over laterite assoc, with Melaleuca laterita”; (Sd) Walter Road, Morley Park, N. G. Marchant 191, Oct. 1958 (PERTH) “aquatic, leaves floating, water 9 deep, flow'ers white”; (Sd) Porongorup Range, slopes of Nancy’s Peak, G. G. Smith, 10.11.1960 (PERTHU); Chittering Valley, G. G. Smith. 27.9.1961 (PERTHU) “in open marsh in paddock, petals white— not keeled internally”. 6. Villarsia violifolia F. Muell. Fragm. Phyt. Aust. 6; 138 (March 1868). Small, low, stoloniferous, mat-forming perennial. Radical leaves on petioles 1 5-15 cm. long; blades small, (4-) 8-20 (-28) mm. long x (5-) 8-25 (-32) mm. broad, mostly semi-rounded to broader than long (up to 1 § as broad as long) but ranging from elliptic (rarely) to broad- ovate also, apex obtuse to rounded, base obtuse to truncate to slightly cordate, or rarely the sinus up to -J length of the blade, edge entire or almost so, texture thin, surfaces semi-matt, or the upper somewhat glossy. Culm slender, 4 5—1 1 (-15) cm. high, branches few, lower nodes with linear-lanceolate bracts only or sometimes cauline leaves on petioles 1-4 cm. long. Inflorescence 2-several flowered, the pedicels rising singly from the culm branches or paired, very slender, those of mature capsules 9-30 mm. long. Flowers homostylous; calyx-lobes 3-5 mm. long, narrow-lanceolate, acute; corolla yellow, a little longer than the calyx, fimbriate at the throat, the lobes without longitudinal keels or wings on the inner surfaces. Capsule 3—5 mm. lone, 1—3 mm. shorter than the 6442/68.-4 28 Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia calyx, the lower adnate to the calyx-tube, thickly chartaceous, open- ing into 4 distinct, recurved valves. Seed 14-17 (-18) mm. long, straw-yellow to fawn-brown or brown-black, ellipsoid, turgid, somewhat laterally compressed, smooth and glossy or matt, surface with minute reticulated patterning, carunculate with an almost-basal, circular, hollow projection of translucent, semi-papillose tissue. Fig. 18 — V. violi folia, shape of radical leaves, x f. a, b, MEL16491-2. c, d, NSW90559, e, f, NSW90560. Distribution : South-west Western Australia, from Perth southwards to the Pemberton and Lake Muir Districts. Habitat: In shallow freshwaters, and on wet soil. Flowering time: FI. and Fr. recorded Dec. -Feb. Typification : Syntypes seen — (Sd) Don River, Maxwell ( MEL 1 649 1-2) ; (Sd) S.W.Austr., Mr. Clarke (MEL16489); Tweed R., W.Austr., Oldfield (MEL16490) “pits, matted fl. yellow”. Discussion : A small, stoloniferous, mat-forming species, with short culms and open, few-flowered inflorescences. Similar in calyx-lobes, capsule, and seed to V. latifolia, but the caruncle rather less prominent. Specimens examined (excluding types) : Western Australia — (Sd) West Australia (MEL16488); (Sd) Upper War- ren River, F. Mueller, 12. 12. 1877 (MEL 16486— 7) "In water and at its edge ; (Sd) Lake Muir, Muir, 1878 (MEL16485); Kelmscott, Canning River, Alex Morrison 9428, 21.12.1899 (NSW90558; PERTH) “swamp": (Sd) Swan district, E. Pritzel, Dec. 1900 (NSW90561); (Sd) Burswood, W.A., W. V. Fitzgerald, Dec. 1900 (NSW90560) “wet soil Fl. yellow”; Big Brook, Max Koch, Nov. 1920 (PERTH) “grows in very moist places”; (Sd) Big Brook, Warren District, W.A., Max Koch 2545, 21.1.192? (NSW90559); (Sd) Jandakot, R. D. Royce 5831, 24.2.1959 (PERTH). Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 29 Fig. 19 — V. violifolia, distribution map. 7. Villarsia calthifolia F. Muell. Fragm. Phyt. Aust. 6 : 140 (March 1868). Large, erect, robust, apparently non-stoloniferous perennial. Radical leaves large, on petioles 1-2 feet long; blades 5-16 cm. long x 6-19 cm. broad on herbarium specimens but noted by Mueller as almost 1 foot long and broad, almost round or either dimension slightly exceeding the other, apex rounded, base deeply cordate, sinus narrow, to J (-%) length of blade, basal lobes sometimes overlapping, margin strongly and irregularly toothed or teeth sometimes less pronounced, the teeth tips callose, texture thin to medium, upper surface glossy or matt, lower surface lightly dotted, veins quite noticeable at least below. Culm erect, measured to 3 feet high on incomplete specimens while Bentham (1868, p. 374) cites Drummond as recording heights of 7-8 feet; cauline leaves present, at least the lower 4 radical-like, the blades to 7 5 cm. long x 10 5 cm. broad on petioles to 19 cm. long. Inflorescence a many- flowered panicle, the final branches and pedicels short, giving a semi- open to compact but not over-dense appearance; final bracts 5-7 mm. long, broad-lanceolate to elliptic, acute; pedicels of mature capsules 7-12 mm. long. Flowers homostylous; calyx-lobes 6-9 mm. long, broad- elliptic to broad-ovate, acute; corolla yellow, fimbriate at the throat, the lobes without any longitudinal keel or wing on the inner surfaces. Capsule (6-) 7-9 mm. long, equal to or slightly greater than to up to 1 5 mm. longer than the calyx, broad-ovoid to ± urceolate, adnate at 30 Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia the base to the calyx tube, the apex and style usually falling by circum- scission, the remaining capsule opening by a circular pore or by 4 (or secondarily more) valves at the summit, the valves truncate, remaining erect or turning slightly outwards, rarely somewhat recurved; capsule wall often thickened beneath the line of circumscission. Seed large, 1 5—1 8 mm. long, straw-yellow to brownish, ellipsoid, very turgid, somewhat laterally compressed, the surface with a minute, reticulated pattern and densely covered with short, hollow, tapered tubercles, without a conspicuous caruncle. Fig. 20— V. calthi folia , radical leaf, x §. a, MEL16481. b, A. C. Beauglehole 12790. Distribution : South-west Western Australia, confined to the Porongorup Ranges. Habitat: Moist, shaded or sheltered positions amongst granite boulders. Flowering time: FI. reported Oct. -Dec., Fr. recorded Dec. Typification : Syntypes seen — Porongorup, F. Mueller, Oct. 1867 (MEL 16480-1) “Cor. yellow”; (Sd) Porongorup, F. Mueller (NSW90574). Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 31 Discussion: A distinctive plant with its large and robust habit, large, circular, cordate, toothed foliage, more compact inflorescence, large, broad, circumscissile capsules with truncate valves, and large, turgid, densely tuberculate seeds without any conspicuous caruncle. Specimens examined (excluding types) : Western Australia— (Sd) W.A., J. Drummond No. 5 (MEL16476-8) ; Porongorup, Mrs. Knight , 1870 (MEL16479); (Sd) Porongorup Range, J. Hale, Dec. 1939 (PERTH) “Amongst granite rocks, in moist spots, fl. yellow”; Devil’s Slide, Porongorup Range, T. E. H. Aplin 2164, 20.10.1962 (PERTH) “in shaded crevices”; (Sd) Top of Nancy’s Peak, Porongorups, A. S. George 6477, 8.12.1964 (PERTH) “Perennial herb. Basal lvs. dying off. FIs. yellow. S. side of granite boulders”; Devil’s Slide, Porongorups, A. C. Beauglehole 12790 Sept. 1965 (Herb.A.C.B.) . in Endl. et al. Enum. Plant. Hueg. 8 8. Villarsia latifolia Benth, (1837). Tall, erect, tufted, apparently non-stoloniferous perennial. Radical leaves erect, on petioles 6-15 cm. long; blades 5-7 cm. Ions x 4-5 cm. broad, broad-ovate to =t= circular to a little broader than" lone, apex obtuse to rounded, base rounded to truncate, edge crenate-dentate, texture thin, upper surface ? somewhat glossy, lower surface ? also somewhat glossy with veins prominent. Culm erect, 1 ft.— 3 ft. 2 in. hieh, 32 Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia branching commencing from about height of culm; cauline leaves rising from the lower 2-6 nodes of culm, 1-4 arising together at each node, radical-like blades to 15 cm. long x 8 5 cm. broad on petioles 2-34 cm. long, the cauline bracts above these becoming bract-like and sessile. Inflorescence an open to semi-open dichotomous panicle, the pedicels arising singly or more usually the panicle branches and pedicels found in near-whorls of 2-5 together; final bracts narrow, lanceolate, acute, 2-3 mm. long; pedicels of mature capsules 5-11 mm. long. Flowers homostylous; calyx-lobes (3-) 5-7 mm. long, lanceolate, acute, noticeably slender; corolla yellow, from description possibly not much longer than the calyx, said to be smaller than that of V. parnas si folia, fimbriate at the throat, each lobe without a longitudinal keel or wing. Capsule 2-5 mm. long, 1-2 mm. shorter than the calyx, globose or sub- globose, the lower adnate to the calyx-tube, thickly chartaceous, opening into 4 distinct recurved valves. Seed large, (1 5-) 1 6—1 8 (-1 9) mm. long, straw-yellow to pale yellow-brown, ellipsoid, very turgid, smooth and very finely and minutely patterned only, glossy or sometimes matt, with a prominent, projecting, ± circular, translucent caruncle just lateral to the base. Fig. 22 — V. latifolia, shape of leaves, x §. a, cauline leaf, MEL16459. b, MEL16464. c, MEL16460. b and c are radical leaves. Distribution : South-west Western Australia, from near Harvey to near Pemberton, and at King George Sound. In water or on wet soil of swamps and river banks. Recorded in granite hills. Flowering time: FI. and Fr. recorded Dec. -Feb. Habitat: Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 33 Typification : Type not seen — King George Sound, Huegel. Specimens examined include several initialled and/or cited by Bentham (1868). These are indicated below by the symbol (B). Discussion : V. latifolia should be readily distinguished from the other large, loose-panicled, yellow-flowered species by its thin-textured, broad, crenate-dentate, non-cordate foliage with raised venation beneath, the several radical-type cauline leaves (at least on large plants), the usually grouped pedicels, the slender calyx-lobes, the capsule shorter than the calyx and adnate to it below, and the distinctive large, smooth, carun- culate seed which resembles only that of V . violifolia. V. violifolia is otherwise, however, a quite different plant. Specimens examined: Western Australia — (B) Preiss No. 1955 (MEL16461-2) ; (B) (Sd) West Austr., Drummond No. 3 (MEL16453-4) ; (B) (Sd) W.A., J. Drummond not numbered but probably also No. 3 (MEL16457); (B) W. Australia, Drummond No. 3 (NSW90568) ; (B) W.A., J. Drummond No. 163 (MEL16467); (B) W. Austr., Drummond No. 4 (MEL16464-5; NSW90567); (B) Swamps near the Blackwood, Oldfield 131 (MEL16463); (B) (Sd) Blackwood River, P. Walcott, Jan. 1868 (MEL16455-6) “River Banks of granite Hills 20 miles from coast” “Growing in Swamps near the Blackwood River. 15 miles from Fig. 23 — V. latifolia, distribution map. 34 Helen 1. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia coast and 30 from Cape Leeuwin”; Warren River, P. Walcott, Feb. 1872 (MEL16466) “Karri Hills and Banks of Streams”; (Sd) Preston River, F. Mueller, 5.12.1877 (MEL16450-1) in drying swamps, corolla lobes yellow, not crested, dilated terminally, leaves concolorous, shining on both sides [in Latin]; Banks of the Harvey River, F. Mueller, 5.12.1877 (MEL1 6458-9) corolla yellow', apex of the lobes truncate-obtuse, not crested, throat fimbriate. Foliage on both sides shining and deeply green, concolorous [in Latin]; (Sd) Preston & Collie Rivers, F. Mueller , 7.12.1877 (MEL 16452), Blackwood and Preston River, F. Mueller, 8.12.1877 (MEL16460) “4’. On banks (not in water) . . (Sd) Lowden, Max Koch 1993, Dec. 1909 (NSW90565) ; Lowden, Max Koch 1993, Jan. 1910 (PERTH); Big Brook, Max Koch, Jan. 1910 (PERTH); (Sd) Greenbushes, R. H. Pullein, Dec. 1917 (NSW90566) ; Yallingup, W. E. Blackall, Dec. 1930 (PERTH) “flowers yellow”; West of Harvey, C. A. Gardner, Sept. 1941 (PERTH). 9. Villarsia parnassifolia (Labill.) R. Br. Prodr . Flor. Nov. Holl. 457 (April 1810) p.p. Swert'a parnassifolia Labill. Nov. Holl. Plant. Specim. 1 ; 72, t.97 (1805). Erect, tufted, non-stoloniferous perennial. Radical leaves erect on petioles (1-) 2 5—14 (-18) cm. long; blades elliptic, ovate, sub-rotund, to sometimes broader than long, (0 7-) 1—4 5 (-5 5) cm. long x (0 4-) 0 7-3 5 (-4) cm. broad, apex acute to obtuse or rounded, base (acute to) rounded or truncate (to sometimes slightly cordate), margin sub-entire to crenate or crenate-dentate, sometimes markedly so, texture thick, apparently semi-succulent, not markedly dorsi-ventral although some- times smoother above, surfaces apparently both matt, the veins often inconspicuous or the major ones only visible. Culm (6-) 11-24 (-32) inches high, the lowest branch found at (V14-) (-|) height of culm; cauline bracts either all bract-like, the lowest sessile or with a broad, semi-stem-clasping petiole, or else the lowest leaf-like with petiole 1 -5-8 cm. long. Inflorescence an open, dichotomous panicle or occasion- ally by reduction of the internodes up to 4 panicle branches and pedicels arising almost together; final bracts lanceolate-acute, 2-3 mm. long; pedicels (of mature capsules) 7-24 mm. long, erect. Flowers described as heterostylous by Bentham (1868, p. 376) but no mention of heterostyly made by Labillardiere in his original description or by subse- quent collectors; calyx-lobes (3 5-) 5-7 (-9) mm. long, lanceolate to elliptic, acute; corolla yellow, apparently about twice as long as the calyx, fimbriate at the throat, the lobes acute to broadly-truncate at the apex, without a longitudinal keel. Capsule (4-) 6-8 (-10) mm. long x 2-3 mm. broad, from rarely slightly < to usually 1-2 mm. > calyx- lobes, membranous, lanceolate to narrow-ovate or pyriform in outline, usually appearing slender, adnate to the calyx-tube only at the base, the valves opening only at the apex and often appearing several. Seed 0 5-0 8 (-1) mm. long, straw-yellow to light-brown, turgid, compressed broad-ellipsoid to almost globular, without a conspicuous caruncle; surface with minute “jig-saw” or entire patterns, slightly granular, occasionally tuberculate with small, dense, blunt tubercles. Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 35 Fig. 24 — V. parnassifolia, shape of radical leaves, x §. a to d, MEL16369 and 16373. e, MEL16368. f, NSW77140. g, NSW90564. h, i, MEL16384. Fig. 25 — V. parnassifolia, distribution map. 36 Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia Distribution : South-west Western Australia, from the vicinity of Busselton and Cape Leeuwin eastwards to about 30 miles east of Esperance. Habitat: Shallow waters in or around freshwater swamps, etc., or on wet flats subject to periodic drying. Apparently mostly in sandy soil. Recorded once in association with Banksia littoralis. Flowering time: FI. reported Oct. -Jan., Fr. Oct. -April. Typification : Under V. parnassifolia Brown included both Swertia parnassifolia Labill. and Menyanthes exaltata Soland. ex Sims. The latter species is however quite distinct, and is here completely excluded from the circum- scription of V . parnassijolia (Labill.) R. Br. Type material “in terra Van-Leeuwin”, i.e., Esperance Bay, should be in Labillardiere’s Herbarium acquired by Webb and now at Florence. Professor Guido Moggi, Herbarium Universitatis Florentinae, Florence, forwarded photographs of a sheet found in the Webb Herbarium, and bearing two different collections. The right hand collection on the sheet is a Labillardiere collection, and the left hand one belongs to the Desfontaine Herbarium. The hand written label “Swertia parnassifolia t. 97 Coll. n. holl.” on the right hand collection is apparently Labil- lardiere’s own, and the specimens belonging with it should be regarded as type material of Swertia parnassifolia Labill. Jn addition, the British Museum (Natural History) holds a Labillardiere specimen with an original label “Swertia parnassifolia. n.h.pl. sp. tab. 97” and this agrees well in habit and appearance with the Florence photo and may be taken as a duplicate of the type collection in Florence. A photograph of the British Museum specimen was forwarded by Dr. W. T. Stearn to Mel- bourne where it is seen to agree well with a Swertia parnassifolia sheet collected by Labillardiere and previously sent as a donation by the British Museum to the National Herbarium of Victoria. This sheet (MEL16384) should also be taken as a duplicate of the type gathering. The National Herbarium of New South Wales, Sydney, has a similar duplicate (NSW90562) of the type gathering donated from the British Museum. The Florence, British Museum, Melbourne and Sydney collections are all in general agreement with the plate provided by Labillardiere except that the leaves are sometimes more strongly toothed than is shown in the illustration. In his description of Swertia parnassifolia Labillardiere described the seeds as many, globose, finely scrobiculate, pale, and straw-yellow. As seeds have proved to have great diagnostic value in the genus Villarsia , those of the four type collections have been matched against each other and appear identical. They also agree very well with Labillardiere’s description except for the colour being more pale-brown Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 37 than straw-yellow, possibly due to darkening with age. Those from the Melbourne collection are sub-globular and 0 5-0 6 mm. in diameter, and several were forwarded to the B.M. where Dr. Stearn compared them with the one seed available on the specimen there. He reported this (letter Jan. 2, 1967) as “subglobose, approximately 0 5 mm. in diameter, with the surface not tuberculate but marked into numerous minute areas exactly as in MEL16384 sent for comparison”. Seeds from the Florence collection were forwarded to Melbourne where comparison with M ELI 6384 showed them identical in surface patterning and apparently similar in shape and size. Precise measurement could not be made due to squashing of the seeds in transit. Seeds of NSW90562 agree well with those of Melbourne in all respects. Enquiries sent to the Director, Conservatoire et Jardin Botaniques, Geneva, revealed that there is no type material of Swertia parnassifolia there. From the four type sheets located I now select the major specimen held at Florence as lectotype. The British Museum, Melbourne, and Sydney specimens thus become isolectotypes. Discussion : This species has been erroneously recorded for South Australia by Black (1957, p. 693). I have seen no specimen of it from that State, and records of its occurrence there seem to be due to confusion with the newly-described species V. umbricola. V. parnassifolia has distinctively small, turgid, usually non-tuberculate seeds unlike those of other species, except V. submersa q.v. The thickish, crenate-dentate, often non-cordate foliage, and slender, membranous often pyriform capsules mostly well-exceeding the calyx are also helpful for identification. Specimens Examined (excluding types): Western Australia— (Sd) Preiss No. 1957 (MEL16375); (Sd) W.A. Drummond No. 6 (MEL16378; NSW90563); (Sd) S.W.Austr. (MEL16381)- (Sd) Orleans Bay, Maxwell (MEL16380) “Next bay east from Lucky Bay . . i.e. Mississippi Bay, not the present Duke of Orleans Bay; (Sd) S.W Austr Tone River (MEL16379); (Sd) Kalgun River, Oldfield 130 (MEL16368)- (Sd) Moirs Inlet, Phillips and Fitzgerald Rivers (MEL16377); (Sd) Near Cape le Grand, Maxwell (MEL16376) “Around fresh water swamps”; (Sd) Cape le Grand, Maxwell (NSW90564) ; (Sd) Bald Island, Maxwell (MEL16383); (Sd) Swamps around King George Sound and into the interior G Maxwell (MEL16382) ; (Sd) Wilson’s Inlet, F. Mueller, Dec. 1877 (MEL16369-73) “on places drying up” “Wet flats, periodically dry” “Lobes of corolla (yellow) entire, broadly truncate, not crested” “Almost terrestrial”; (Sd) Porongorup Mrs. Knight (MEL16446); (Sd) Busselton, Dr. T. C. Rosselloty (MEL16366) ’ (Sd) Lake Muir, W.A., Muir (MEL16367); (Sd) King Georges Sound, Webb Dec. 1882 (MEL16374); (Sd) King George’s Sound, J. H. Maiden Nov 1909 (NSW77140-1 ) ; (Sd) Nillup via Margaret River, R. D. Royce 13 14 1 1944 (PERTH) “Petals yellow, villous. 12-24 inches tall”; (Sd)' 3 miles N of Esperance, R. D. Royce 4115, 21.4.1953 (PERTH); (Sd) 4 miles E. of Albany /. W. Green 868, 1.12.1956 (PERTH) “Herb 80 cm. high; fls. yellow In water’ Swampy sand, assoc, with Banksia littoralis”; (Sd) 12 miles W of Denmark on Manjimup Road, A. R. Fairall 631, 12.10.1962 (PERTH) “Plant 2’ Yellow fl Succulent foliage” “Growing in water”; (Sd) 1 1 miles from Denmark towards 38 Helen 1. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia Manjimup, M. E. Phillips, 12.10.1962 (CBG) “‘In wetland patch in swamp. Sandy peat.” “yellow”; (Sd) Cape Augusta, A. R. Fairall 813, 18.10.1962 (PERTH) “Herb 18". Flowers yellow”; (Sd) Albany-Denmark road, 18 miles W. cf Albany, T. E. H. Aplin 2179 , 20.10.1962 (PERTH) “swampy soil”; 23 miles E. of Esperance, M. E. Phillips, 4.11.1962 (CBG) ‘T ft. high, in sandy soil.” 10. Villarsia exaltata (Soland. ex Sims) G. Don Gen. Syst. 4 * 169 (1838). Menyanthes exaltata Soland. ex Sims in Curtis's hot. Map. 26 : sub. 1. 1029 (1807), icone excluso. Erect, usually robust, tufted perennial, without stolons. Radical leaves rising erect, never floating, on petioles (3^-) 5-16 (-24) in. long; blades usually ovate-elliptic, broad-lanceolate, narrow-ovate or broad- ovate and longer than broad to rarely rhomboid or sub-rotund and length zb equal breadth, to very rarely the breadth slightly > length, outline often somewhat irregular, (4-) 6-12 (-13-5) cm. long x (1 -5-) 3-8 (—9 5) cm. broad, apex acute to obtuse or rounded, base mostly rounded to shallowly cordate, occasionally truncate or oblique, edge entire or sometimes slightly to zb strongly crenate-dentate, often un- dulate, surfaces matt and zb uniform in coloration or upper surface slightly darker, texture thick and at times almost semi-succulent, veins not markedly pronounced. Culms 1 -several, 18 in.-5 ft. high, standing erect well above the radical leaves; cauline leaves 1 -several, solitary at each node or several together, usually subtending a panicle branch but the lower ones not always doing so, lowest leaf arising at about the level of the blades of the radical leaves, in larger plants at least the lower 2 cauline leaves similar to the radical leaves, on petioles ( 1-) 2-7 (-1 1 ) in. long and semi-sheathed at the base. Inflorescence an open, many- flowered panicle; final bracts ovate, acute, (2-) 4-5 mm. long; pedicels (of mature capsules) 4-13 (-18) mm. long, remaining erect. Flowers heterostylous; calyx-lobes 4 5-8 mm. long, lanceolate-ovate, acute, (-3—) %-j (-§) length of the corolla; corolla yellow, 9 5—1 8 mm. long, 1 6—30 (average 22) mm. span, lobes 5 or sometimes 4, 7 5—12 mm. broad, variable in shape and apex (Fig. 2), without a longitudinal keel on the inner surface or very rarely having a reduced portion of one (Fig. 28); gynoecium normally with 2 placentas and 2 stigmas but occasionally 3 of each, ovules 10-19 per placenta. Long-styled flower with style slender, (3-) 4-6 mm. long; stigmas papillose, longer than broad, erect, zb ovate and with recurved edges, 1—2 5 mm. long, held well above the anthers; anthers 1 -6—2-2 mm. long, on short incon- spicuous filaments (ca. 1 mm.). Short-styled flower with style thicker, 0 5-2 mm. long; stigmas papillose, broader than long, semi-erect to broadly-spreading, irregularly rounded, somewhat undulate, 1-18 mm. broad, held below the level of the anthers; anthers 2-3 mm. long, on noticeable filaments 2-3 mm. long. Capsule 5-13 mm. long, from a little < to slightly > the calyx-lobes, thickly chartaceous, sub-globular to broad-ellipsoid, the lower -J — \ adnate to the calyx tube, dehiscing at the summit into 4 acute, slightly-recurved valves. Seed large, ( 1 5—) 1 7—2 6 (-3) mm. long, light cream-fawn to pale grey-brown to dark Helen T. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 39 Fig. 26 — V. exaltata, shape of radical leaves, x f. a to f, Aston 1457. a to c, less usual shapes, g, MEL16339. h, Aston 1579. 40 Helen T. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia Fig. 27 — Heterostyly in V. exaltata. a, short-styled flower and stigma, b, long- styled flower and stigma. From Cranbourne, Victoria. Flowers x 6, stigmas x 14. Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 41 brown-grey-black, compressed broad-ellipsoid, double convex, with a conspicuous, whitish to translucent, circular, hollowed caruncle; surface with minute “jig-saw" patterns, smooth to slightly granular, and mostly also strongly and conspicuously tuberculate, or the tubercles sometimes reduced in size and number or very rarely completely lacking; tubercles terete, scattered to ± densely arranged, up to 0 3 mm. long but usually half that length, each tubercle again minutely tuberculate at the extremity, its summit expanded. Fig. 28 — V. exaltata. Abnormal, rudimentary keels on midlines of corolla-lobes. Cranbourne, Victoria. Distribution : Near-coastal areas from about Gympie in south-east Queensland south through New South Wales and west to Cranbourne in Victoria, also in north-eastern Tasmania. There are only two known localities for this species in Tasmania, St. Mary’s and Scamander, but it very probably existed previously (and may still occur) near Georgetown. Hooker (1860, p. 272) when dis- cussing Villarsia in Tasmania wrote “Gunn considers that . . . [there] . . are two perfectly distinct species, and observes that one has floating leaves, and grows in lakes; the other has erect leaves, and grows in marshes only, near Georgetown, flowering a month earlier than the other”. I would consider that Gunn’s description of the Georgetown species applies very well to V . exaltata , including the reference to earlier flowering (whenever the two species concerned occur together under the same conditions). The species he describes first would be V . reniformis which is widespread in Tasmania. Sandy heath land; fresh-water aquatic in up to 2 feet (possibly a little more) of water in permanent swamps, or amphibian on low-lying areas subject to inundation in winter or to seepage, in summer surviving on dry or near-dry soil in such areas; open or semi-open situations receiving good sunlight, sometimes sheltered by denser, higher vegetation. The largest and most robust plants are found in water in open areas. Some associated species recorded are Banksia marginata, Leptos- permum juniperinum, L. myrsinoides, Melaleuca ericifoHa , M. squarrosa, Sprengelia incarnata, Epacris micro phylla, Bauera rubioides, Gahnia clarkei, Restio tetraphyllus, Lepidosperma sp., Juncus sp. sect, genuini, Scirpus nodosus, Haloragis micrantha. Cent el l a asiatica, Utricularia dichotoma , Drosera binata, Selaginella uliginosa, and many small sedges. Habitat: 42 Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia Fig. 29 — V. exaltata, distribution map. Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 43 Flowering time: FI. Aug. -May (in Victoria chiefly Oct. -Jan.), Fr. Oct. -May (in Victoria chiefly Nov. -March). Typification : The application of this name has created considerable confusion in the past. Mueller (1868, p. 139) and Bentham (1868, p. 376) both considered it synonymous with V . reniformis R. Br., which name they both retained. Since then, all floras of the eastern States have continued to accept this synonymy and have mostly used either V. reniformis or V. exaltata as the accepted name. However, these two entities are quite distinct, and are here redefined as separate species. The confusion over the application of V. exaltata has undoubtedly arisen because of the lack of clarity in the type description of its basonym Menyanthes exaltata and the publication with it of an erroneous plate. This plate agrees more with V. reniformis than it does with V . exaltata, and is here excluded from the latter species. The selection of lectotype material requires some explanation. It seemed logical to suspect that both Solander’s manuscript and the material in the Banks Herbarium referred to by Sims in his description of Menyanthes exaltata would apply to material seen and gathered by Banks and Solander on Cook’s Voyage of 1770. The furthest south they could have collected would be Botany Bay, New South Wales, for this was their first place of landing during the voyage northward up the east coast of the continent. Field work and examination of herbarium collec- tions showed that the species of Villarsia occurring (quite commonly) at Botany Bay and northward is quite distinct from the more southern species which agrees well with the illustration published by Sims. It was therefore highly probable that the plate and the Banksian material were referable to different species. Accordingly, samples of leaves and seeds of Aston 1430 and 1431 (representing the two species in question) were forwarded to Dr. W. T. Stearn of the British Museum together with an explanatory sheet concerning points of similarity and difference. Dr. Stearn, replying by letter dated 2 Jan. 1967, wrote: “Comparison of Banks and Solander material with that sent shows closest agreement in leaf with Aston 1431 the leaf-blades being distinctly longer than broad, their surfaces matt, the veining on the lower surface not raised. The seeds are about 2 mm. long by 18 mm. broad, with a conspicuous pale caruncle at base and a distinctly tuberculate surface, though not so profusely tuberculate as in Aston 1431 and also much darker in colour . . . They are unlike those of Aston 1430.” This comparison together with two seeds sent for my own examination, confirms that the Banksian material is conspecific with the entity here redefined as V. exaltata. In addition, examination of Solander’s original manuscript showed that the material was collected at Sting Ray’s Bay (now Botany Bay) between April 28-May 5, 1770, and this locality is verified by a note “ New South Wales: Botany Bay. J.B.”, i.e., Joseph Banks, on the back of the herbarium sheet. 6442 /68. — 5 44 Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia Having determined the true nature of the Banksian material the question arose as to the lectotype of Menyanthes exaltata Soland. ex Sims. Dr. Stearn (letter 2 Jan. 1967) advised “Normally one would have accepted the plant figured by Sims. Here, however, there is a definite reference to ‘Soland. MSS. et Herb. Banks’, with both of which Sims was well-acquainted, and there is nothing in the diagnosis which could not have been taken from the Banks material. This is followed by the statement ‘a native of New South Wales, where it was discovered by Sir Joseph Banks’. Thus it would seem logical as well as taxonomically convenient to typify Menyanthes exaltata by the material in Banks Herbarium (collected at Botany Bay) which Sims cites. My colleagues Mr. J. E. Dandy and Mr. R. Ross . . . independently made the same choice when confronted with the evidence . . . Botany Bay, New South Wales, then becomes the restricted type-locality”. I am in absolute agreement with this decision, and the Banksian material (BM) here referred to now becomes the lectotype of the species. A Banks and Solander duplicate collection (NSW90511) obtained by the National Herbarium of New South Wales from the British Museum becomes an isolectotype. It agrees well with a photo of the lectotype forwarded to Melbourne. Discussion : A very distinctive species readily separated from V . reniformis , the only species sympatric with it, by the erect, thick-textured, matt-surfaced, typically ovate foliage, the heterostylous flowers with smaller span, the capsule rather more globular and adnate below to the calyx-tube, and the larger, strongly-tuberculate seeds. In almost all situations the species can be distinguished by foliage alone. At Cranbourne, where both species occur side by side in a swamp, V. exaltata was found flowering a month earlier than V . reniformis, but I have insufficient evidence to determine if this is generally so. It is likely that variation in water level partly controls flowering time. (See also under distribution.) V. exaltata seems closely related to V. lasiosperma of south-western Australia. The two species are virtually indistinguishable vegetatively, and both have large tuberculate seeds although the tubercles are of different types. The corollas of V. lasiosperma, although white, possess a longitudinal keel or wing on the inner surface of each lobe, a feature suggested in some abnormal flowers of V. exaltata (Fig. 28) but not found in other species. No attempt has been made to ascertain the breeding system associated with the heterostylous nature of this species, but seeds have been forwarded to Dr. Robert Ornduff, University of California, Berkeley, U.S.A., who is studying this subject in Menyanthaceae. In the field both style types appear functionally bisexual, and set apparently viable seed in equal quantities. There is no noticeable distinction between flower, capsule, and seed sizes of short-styled and long-styled plants. I he only exception occurred in a small population near Tarwin Lower (Aston 1577) where capsule length varied from 5-9 mm. on one plant Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 45 to 9-13 mm. on another. As flowering was completed it was not possible to determine if this difference was associated with different style types, but habitats appeared identical. Ornduff (1966) is topical to this aspect. Specimens examined: Queensland — Durval (the present Toorbul), Leichhardt , 25.9.1843 (NSW90512); (Sd) Maroochie Swamps, F. M. Bailey, Oct. 1874 (BRI061778); Queensland, Eaves, 1874 (MEL16331); (Sd) Wellington Point, J. Wedd, Nov. 1891 (BRI061777) “This plant seldom has reniform leaves”; (Sd) Lake Cootharaba, /. Keys , 1911 (BRI061776); Coolangatta, C. T. White, Sept. 1912 (NSW90513) “Very common in swamps”; Swamps at back ol Coolangatta, C. T. White, Sept. 1912 (BRI061774); (Sd) Bribie Island, W. D. Francis, Nov. 1919 ( BRI06 1 772 ) ; Elimbah, C. T. White 3248, 12.9.1926 (BRI061775). New South Wales — (Sd) Hastings River, Dr. Beckler (MEL16330); (Sd) Manly swamps, W. Woolls (MEL16329) “leaves not floating”; Manly beach. W. Woolls (MEL16333); Port Jackson, Manly, E. Betche, 30.11.1881 (NSW90524); Waterloo marshes, Sydney, Coll.? T. Whiteleggc, Dec. 1884 ( MELl 6328) [Two small packets containing flowers only, one labelled “short style and long stamens”, the other “Long style with short stamens. The lobes of the stigma are longer than the short style one”]; (Sd) Broger's Creek, W. Bauerlen 24, May 1886 (MEL16327); (Sd) Richmond River, N.S.W., 1887 (MEL 16326) ; Kogarah, T. S. Cumfield, Oct. 1893 (NSW90528) ; (Sd) Wyong. J. L. Boorman, Jan. 1903 (NSW90521); (Sd) Byron Bay, J. H. Maiden & J. L. Boorman, Nov. 1903 (NSW90515); Coffs Harbour, J. L. Boorman, May 1909 (NSW90517) “A very common plant found everywhere on the coast in permanently wet places. Growing from less than 1-4 ft. high”; (Sd) Table Mountain near Milton, R. H. Camhage 4039, 27.12.1913 (NSW90534) ; Botany Swamps, L. Abrahams, April 1914 (NSW90529); Johns River, J. L. Boorman. Nov. 1915 (NSW90518); Johns River, J. L. Boorman , Dec, 1915 (SYD); La Perouse, H. M. Rupp, April 1916 (MELU); (Sd) Wyong, J. L. Boorman, Nov. 1916 (NSW90522) ; (Sd) Narrabeen, H. M. Rupp, Jan. 1917 (MELU); R.A.N. College, Jervis Bay, F. A. Rodway, Jan. 1918 (NSW91012); Bowen Island, Jervis Bay, F. A. Rodway, Nov. 1919 (NSW91015); (Sd) Bowen Island, Jervis Bay, F. A. Rodway, Dec. 1925 (NSW91014); (Sd) Botany Bay District, O. D. Evans, 8.12.1925 (SYD) “in a swamp” “erect” “flower yellow”; (Sd) Dee Why, O. D. Evans, 23.2.1926 (SYD) “margin of swamp” “flower yellow”; Fitzroy Falls, S. Coast, N.S.W., F. A. Rodway 233, 30.11.1930 (NSW91013); Byron Bay, C. T. White 10430, 24.8.1936 (BRI061773) “Fairly common on marshy ground. Rather a succulent plant with very brittle stems. Flowering stalks erect, branching, about 3 ft. high. Flowers yellow”; Cape Solander, L. A. S. Johnson, 20.10.1945, (NSW90530) “Common in wet sedge-swamp” “(Swamp) about V deep” “Per. herb, 60 cm., hydrophyte, leaves sub-aerial” “FI. yellow”; (Sd) Narrabeen, H. Salasoo 1027, 4.1.1953 (NSW90523) ; Centennial Park, G. Chippendale, 19.8.1953 (NSW90526) “in swamp”; (Sd) North of Boote Boote, L. A. S. Johnson, 13.10.1953 (NSW90519) “Petals yellow, fringed. Common on edge of Cladium swamp”; Byron Bay, per Shire Council, Sept. 1954 (NSW90516) “reaching pest proportions in low’ lying swampy heath land”; (Sd) Maroubra, N. C. Ford, Nov. 1954 (NSW90527) “among thick sedge vegetation in hind-dune quaking bog. FI. yellow'”; (Sd) Carrington Falls, E. F. Constable, 22.1.1955 (NSW30798) “Erect slender herb, 2-3', fls. yellow. Scarce. Sandstone. 1850'”; Pigeon House Range (between Braidwood and Jervis Bay), E. Gauba 008176, 21.11.1958 (CBG); (Sd) Little Bay, E. F. Constable, 22.11.1960 (NSW53341) “Erect herb 12-18", flowers yellow. Localised to one area. Swampy ground. Sandstone. Altitude 50 ft.”; (Sd) Picton Lakes, Thirlmere, E . J. McBarron 7119, 21.4.1962 (NSW90533) “Yellow flowered- fringe of lake rooted in water”; (Sd) Picton Lakes, D. Blaxed, 13.12.1962 (NSW90532) “Edge of lake — damp soil” “Sandy” “Height 5 ”; (Sd) Duranbah, Tweed River, H. S. McKee 11655, 6.10.1964 (NSW90514) “recently cleared damp sandv ground” "erect; leaves dark glossy green above, light glossy green below; leaf bases purple; flowers yellow”; Nelson Bay, G. M. Lithgow 96. 15.2.1965 46 Helen I. Aston: The Genus Vi liar si a ( Menyanthaceae ) in Australia (NSW90520) “Common in swampy areas" “taken from swamp margin"; Tabbigai, Kurnell Peninsula, B. G. Briggs, 13.8.1965 (NSW90531) ‘Flowers golden yellow. Bases of stems and petioles purplish. Beside pool in sedge swamp among dunes": (Sd) Kurnell. B. G. Briggs and L. A. S. Johnson, 13.1 1.1965 (NSW90508) “rooted in wet sand near deep swamp; at edge of scrub adjoining bare sand. Short-styled flowers". Also same collection (NSW90509) "Long-styled flowers (long and short styles apparently ± similar in numbers in the stand)". Also (NSW90510) “Long-styled flowers": Picton Lakes, Thirlmere, L. A. S. Johnson & B. G. Briggs, 21.11.1965 (NSW90507) “rooted in sand near water's edge and in shallow water. Flowers yellow" "alt. 1000 ft.": Myall Lakes, near Bulledelah, 3. Jacobs, 2.10.1967 (MEL26099) "Melaleuca quinquenervia swamp forest"; Wingecarribee Swamp, near Bowral. L. A. S. Johnson, 18.11.1967 (NSW97481) "Edge of peat bog only, in ca. 15 cm. water". Victoria — Snowy River, F. Mueller, March 1854 (MEL16337); (Sd) Australia Felix. F. Mueller. Jan. 1855 (MEL 16335); (Sd) Austral. Alps, F. Mueller, Jan. 1855 (MEL16338) locality considered incorrect — probably lower Snowy River; (Sd) New'merella, C. H. Grove, 12.2.1906 (M ELI 6334) “ovate-leaved form" “swampy ground"; Wilsons Promotory, high plain, Audas and St. John, 11.11.1908 (MEL16336); Reedy Creek. 3m. E. of Cann River, N. A. Wakefield 2869 (MEL25464); Mario to Cape Conran road, N. A. Wake- field 3785 (MEL25463); Cranbourne Swamp. E. J. Sonnenberg, 16.10.1936 (MELU); Lang Lang, coll.? 23.11.1962 (MEL 16339-41 ) ; Port Welshpool swamp, D. H. Ashton, Jan. 1963 (MELU); Cranbourne, H. I. Aston 1236, 27.10.1964 (MEL16343-4) ; (Sd) Cranbourne, H. I. Aston 1328 A, 19.1.1965 (MEL16342); (Sd) Cranbourne. H. /. Aston 1391, 17.11.1965 (MEL16356-9) ; (Sd) 7 miles S. of Lang Lang, H. I. Aston 1431, 10.12.1965 ( MEL16345-7) ; (Sd) Lang Lang. H. 1. Aston 1432, 10.12.1965 ( MEL16360-2) ; Reedy Creek, 4-5 miles S. of Cann River. H. 1. Aston 1455, 11.1.1966 (MEL16348); (Sd) 9 miles east of Mario. H. /. Aston 1457, 12.1.1966, (MEL1 6349-52) ; Stradbroke, H. /. Aston 1459, 13.1.1966 (MEL16363); (Sd) 4 miles S.S.E. of Foster, H. I. Aston 1461, 14.1.1966 ( MEL16353-5) ; (Sd) Tarwin Lower. H. I. Aston 1577, 2.3.1967 (MEL20425-6) ; (Sd) 6 miles S.E. of Tarwin Lower, H. 1. Aston 1578, 2.3.1967 (MEL20427- -9); (Sd) Cape Liptrap. H. 1. Aston 1579, 2.3.1967 (MEL20430-1) ; 7 miles S.S.W. of Fish Creek, H. I. Aston 1580, 2.3.1967 (MEL20432). Tasmania — St. Mary's, L. Rodway (HOB) “on land"; (Sd) One mile north of Scamander, T. E. Burns, 16.5.1966, collected alive and grown on at Melbourne from where material pressed (MEL20447-9) . 11. Villarsia reniformis R. Br. Prodr. Flor. Nov. Holl. 457 (April 1810). Menyanthes sarmentosa Sims in Curtis’s bot. Mag. 1328 (Oct. 1810). Stoloniferous perennial, conspicuously red-dotted on submerged portions, the culm erect or semi-erect, the radical leaves typically float- ing. Stolons usually strongly developed when plants aquatic in deeper, near-permanent waters and then 15 in. -6 ft. long, up to 15 cm. diam., rooting and shooting at intervals of several — 40 cm. along their length, several rising from the base and sometimes also one from the axil of the lowest cauline leaf when this is submerged; stolons usually not produced on plants occurring where waters are shallow and temporary. Radical leaves typically floating (sometimes erect) on plants in deeper waters, but normally erect on plants produced by shallow temporary water, on petioles 3-27 in. long; blades reniform, orbicular, or broad-ovate, mostly about as broad as long or slightly longer than broad, but varying from length equal § to % of breadth, length (3-) 4-8 (-10 or rarely 12) cm., apex obtuse to rounded or rounded-emarginate, base cordate, the lobes Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 47 mostly well-pronounced and up to £ (-<■£) of the leaf length, rarely as little as Vi 2 of the leaf length or the base ± truncate, sinus broad to extremely narrow or the lobes sometimes overlapping, edge entire to slightly (rarely strongly) crenate-dentate, texture mostly moderately thick; floating blades strongly dorsi-ventral, upper surface deep green and glossy, the lower pale cream-green and strongly dotted, often coloured with maroon-red, the veins clearly visible; erect blades as for the floating or often less dorsi-ventral with upper surface paler and non- glossy, the lower greener with dots inconspicuous to almost lacking. Culms solitary or few, 17 in.-3 ft. 8 in. high, standing erect above the leaves or in deeper water semi-erect and half-reclining amongst surface vegetation; cauline leaves 1 -several, solitary at each node, lowest leaf Fig. 30— V. reniformis , shape of radical leaves, x |. a to d, Aston 1428. e, Aston 1581. f, Aston 21.10.1960. g, Aston 1393. 48 Helen 1. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia (often absent from herbarium specimens) arising from near the culm base to above half the culm height, on larger plants the lowest 1-2 cauline leaves radical-like, on petioles from very short to 20 in. long. Inflorescence an open, many-flowered panicle, usually confined to near the summit of the culm; final bracts ovate, acute, ± 3-5 mm. long; pedicels (of mature capsules) (7-) 10-25 (-37) mm. long, remaining erect. Flowers homostylous; calyx-lobes 6 5-10 (—11) mm. long, lanceolate, acute, %-% length of the corolla; corolla yellow, 12-23 mm. long, 19-43 (average 32-3) mm. span, lobes 5 or sometimes 4 or 6, 7- 14 mm. broad, variable in shape and apex (Fig. 2), without a longi- tudinal keel; gynoecium with 2 placentas, ovules 41-88 per placenta, style 0 5-3 mm. long, stigmas 2, papillose, rather variable in shape and length-breadth ratio, 0 5—1 5 (-2 5) mm. long, held about level with the anthers; anthers 3-4 (-5) mm. long, on short filaments 1-2 mm. long. Capsule 7—1 1 5 mm. long, from a little < to > calyx-lobes, and sometimes exserted by as much as i the length of the calyx-lobes, thickly chartaceous, ovoid-ellipsoid, adnate to the calyx-tube only at the very base, dehiscing at the summit into 4 acute, somewhat-recurved valves. Seed ( 0 7—) 1 — 15 (-19) mm. long, pale cream-fawn-yellow to light reddish-fawn or reddish-brown, sometimes greyish-fawn but never dark grey or black, compressed broad-ellipsoid to almost compressed globular, double convex or thickened in the middle and with a slight intra-marginal depression, sometimes almost flat across the sides, point of attachment without a conspicuous caruncle; surface with minute “jig-saw” patterns, smooth to slightly granular, mostly dull but at times glossy, without tubercles. Distribution : South-east South Australia and Kangaroo Island, Tasmania, southern Victoria, and south-east New South Wales. Within South Australia the distribution of this species is in part uncertain. It is the common Villarsia of the south-east, around the Mt. Gambier-Kingston-Naracoorte region and might be located in future as far north as Bordertown if a series of wet seasons provided suitable swamps. A few collections from Kangaroo Island also agree with this species, although none of those seen by the writer shows the typical robust, stoloniferous nature characteristic of the plants of the south-east and elsewhere, while one of them (Hj. Eichler 18466) has seeds which by their slightly granular nature suggest affinity with V . umbricola var. umbricola. There are several mainland specimens from the vicinity of Adelaide-Macclesfield-Myponga which could possibly represent V. reniformis but which are either incomplete or limited in demonstrating essential characters. They might be referable instead to V. umbricola var. umbricola , but positive identification cannot be made. V . reniformis should be considered as only doubtfully present in the area concerned. Dr. Eichler of the State Herbarium of South Australia informs me that he has never seen V. reniformis in the Mt. Lofty Ranges or on the Fleurieu Peninsula. Further work there and on Kangaroo Island would be desirable. Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 49 Tilba, The Clyde, Bombala, and near Bowral, in south-east New South Wales, are the only four definite localities for that State There is a possibility that the species may have occurred in the Sydney district, but this rests only on two doubtful specimens. An early collection of Woolls from Parramatta has foliage only and its identity remains suspect, while a specimen labelled Kogarah 1893 is of doubtful locality due to its admixture with another collection. Specimens collected in 1805 by G. Caley and labelled New South Wales would almost certainly have come from Tasmania. Fig. 31— V. reniformis, distribution map. £ Definite records. A Doubtful records. Habitat: Swamps and depressions on varied soils, often sandy or peaty; aquatic in permanent freshwaters up to 2 feet deep or probably more’, or found in shallow seasonal waters, dying back when stranded by falling levels; open or semi-open situations receiving good sunlight. The largest and most robust plants are found in deep water in open areas. 50 Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia Some associated species recorded are Leptospermum juniperinum, L. lanigerum, Melaleuca erici folia, M. squamea , Acacia verticil lata, C l odium glomeratum, Gahnia clarkei, Eleocharis sphacelata, E. acuta, Restio tetraphyllus , Lepidosperma sp., Lepyrodia sp., Juncus maritimus , J uncus sp. sect, genuinii, Amphibromus recurvatus , Myriophyllum propinquum, M. muelleri, Triglochin procera , T. striata , Potamogeton tricarinatus, P. oc hr eat us, Scirpus fluitans, Claytonia austral asica, Utricularia flexuosa, Ranunculus rivularis, Haloragis brownii , Selliera radicans, Samolus re pens, Crassula helmsii , and small sedges. Flowering time: FI. Oct. -Jan. (-March), Fr. (Oct.-) Nov. -March. Typification : When investigating the confused typification of V. exaltata (q.v.) and V. reniformis selected specimens were forwarded to the British Museum for comparison with type material. Dr. Stearn, in a letter dated 2 Jan. 1967, wrote: “Brown’s specimen from Port Dalrymple collected in 1804 lacks flowers and the capsules have disintegrated but the leaf-blades are cordate, with a marked difference in colour between the upper and lower surfaces, the lower are conspicuously dotted and with evident veining as in Aston 1428; the largest is about 3 cm. long and broad. A better specimen written up by Brown as Villarsia reniformis collected at Port Dalrymple by Paterson agrees closely in size, etc., of leaf with Aston 1428. The seeds of this are smooth and glossy, about 12-14 mm. long”. This comparison adequately verifies the true nature of V. reniformis, and as Brown’s own collection is incomplete the Paterson collection is now chosen as lectotype. A photograph of this sheet was forwarded to Melbourne. As Menyanthes sarmentosa Sims was based on cultivated material the plate itself must be taken as the type. This agrees well with V. reniformis, and in addition Sims notes in his description . . the remarkable runners, several feet in length, putting out leaves and roots, at intervals of about a foot and a half . . a feature of the aquatic form of this species. Discussion : This species has long been confused with V. exaltata but in its typical form with glossy, reniform-orbicular, cordate, floating leaves and conspicuous stolons it is readily distinguished. See also under V . exaltata discussion and typification. For distinctions between V. reniformis and V. umbricola see under the latter species. There is a general uniformity of seed size amongst the plants of any one population, the maximum variation found being 0 5 mm. On the other hand there is often a noticeable variation in seed size between the plants of different populations, even when these populations occur within the same district. For example in south-western Victoria two separate swamps in the Edenhope-Langkoop area had seed sizes of Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 51 1 -5—1 -7 (av. 1 57) mm. for 42 seeds measured and 1 5—1 7 (av. 1 63) mm. for 56 seeds measured. At both places many more seeds noted by eye were similarly large in size. In contrast, another population in the Edenhope district, with 39 seeds measured had small seeds 0 9—1 2 (av. 1 08) mm. Other small-seeded populations at Heywood in south- western Victoria (81 seeds, 0 7—1 mm., av. 0 91) and at Dalmore south- east of Melbourne (42 seeds, 0 7—1 mm., av. 0 88) again illustrate how the variation in seed size between different populations is not related to geographical distribution. It could indicate a greater efficiency of vegetative spread over sexual reproduction within each small habitat zone, a feature which would be suspected from the stoloniferous habit of the species. Flowers of V . reniformis are quite bright yellow in colour. The only exception found was in a swamp in western Victoria where some plants with pale yellow corollas were scattered amongst the population. Most plants in the swamp were normally coloured. Specimens examined: South Australia — Yallum, ca. 15 km. W. ol Penola. Herb. R. Tate 106, Nov.? (AD966 1 00364) ; Harriet River. Kangaroo Is.. T. G. B. Osborn, 25.11.1923 (AD966 1 00362) ; Middle River, Kangaroo Is., R. L. Crocker, Dec. 1940 (ADW); Near Kelly Hill, Kangaroo Is., H). Eichler 15271, 7.11.1958 (AD96301032) ; 10 km. E. of Penola, P. Wilson 1293, 16.11.1959 (AD96022051 ) ; Bool Lagoon Lucindale road, D. Hunt 260, 21.10.1961 (AD96151 148) ; W. of Bool Lagoon, D. N. Kraehenbuehl 1037, 12.10.1963 ( AD96417201 ) ; 30 km. S.W. of Naracoorte. D Hunt 1738, 24.12.1963 (AD9640521 3) ; (Sd) Kelly Hill Caves Reserve. Kangaroo Is.. Hj. Eichler 18466, 23.12.1965 ( AD966 1 1 0340: MEL25436); (Sd) Between Joanna and Struan, M. A. Fagg 119, 28.1.1966 (AD9661 10322) ; (Sd) 4 miles S.E. of Tarpeena, H. I. Aston 1517, 15.12.1966 ( MEL20459-60) ; 6 miles S.E. of Tarpeena. H. I. Aston 1518, 15.12.1966 (MEL20458); (Sd) 6 miles W. of Kalangadoo, H. I. Aston 1519, 15.12.1966 ( MEL20455-7 ) ; (Sd) 7 miles E.S.E. of Beachport. H. 1. Aston 1524, 16.12.1966 (MEL20454); (Sd) 12 miles S.E. of Robe, H. I. Aston 1525, 16.12.1966 (MEL20453); (Sd) 3*5 miles E. of Hynam, H. !. Aston 1533, 20.12.1966 (MEL20452); (Sd) 8 miles E. of Struan, H. 1. Aston 1534, 20.12.1966 (MEL20451); (Sd) 7 miles N.E. of Penola, H. /. Aston 1535, 20.12.1966 (MEL20450 and 25437); (Sd) 10 km. S.E. of Mt. Burr township, /. B. Wilson 715, 11.3.1967 (MEL26379; AD96719058) “in shallow waters of swamp interspersed with Gahnia clarkei clumps”; Larrikin Lagoon, Kangaroo Island. J. B. Cleland , 28.10.1967 (AD96748176) . New South Wales — (S d) Tilba, N.S.W. Reader, Dec. 1879 (MEL16290); The Clyde, W. Bauerlen 78, Oct. 1884 (MEL 16292); Bombala, W. Bauerlen 279, Dec. 1886 (MEL26384); Wingecarribee Swamn, near Bowral, L. A. S. Johnson, 18.11.1967 (NSW97482) “In water ca. 20 cm. deep on peaty bottom. Leaf-blades emerging, not floating”. Victoria — Near Melbourne, coll.?, 20.9.1856 (MEL16268) “Marshy ground and waterholes” “Common”; Mount Mclvor and Mclvor Creek. F. Mueller, Nov. (MEL16267); Ballarat, Bacchus (MEL16270); (Sd) Near Mt. Gellibrand. F. Mueller, Mar. 1875 (MEL16266); Barwon, Queenscliff Road, 7. B. Wilson, 2.12.1882 and 1883 (MEL16271); Port Phillip. Reader, Dec. 1884 (MEL16272); Lilydale, A. H. S. Lucas, Nov. 1885 (NSW90539); (Sd) Near Lake King, C. H. Grove, 1887 (MEL16318; MEL27196); Wimmera, Davis, 1890 (MEL16259 and 16317); Upper Yarra, C. Walter, Dec. 1896 (NSW90540); (Sd) Frankston. C. Walter, Dec. 1897 (NSW90537); Hawkesdale, H. B. Williamson, Oct. 1898 (NSW90536); Dromana, 7. H. Findlay, 12.12.1898 (MEL 16325); Brim, W. S. Macpherson, 1898 (MEL16262) “springs lagoons”; (Sd) Dandenong, 52 Helen I. Aston: The Genus Villarsia (M enyantlwceae) in Australia W. R. Baker, 20.10.1899 (MEL16277); (Sd) Dandenong, W. R. Baker, 1.12.1899 ( MEL16276) ; Beaconsfield. W. R. Baker, 24.10.1901 (MEL16324); Springvale, C. S. Sutton, Nov. 1901 (MEL-16269); (Sd) Springvale, G. Weindorfer, Dec. 1901 (MEL 16263) ; (Sd) Cheltenham. K. Cowle (MEL16257-8) ; (Sd) Vermont. W . R. Baker , 8.12.1902 (MEL16273); (Sd) Pakenham, W. R. Baker, 26.11.1904 (MEL16279); (Sd) Mornington Junction 10.11.1905 and Officer 30.11.1905 (mixed collection), W. R Baker ( MEL16274-5) : Wilsons Promontory, A. D. Hardy, Dec. 1905 (MEL 16265); Frankston, W. R. Baker, 14.12.1905 (MEL 16323) ; (Sd) Newmerella, C. H. Grove, 12.2.1906 (MEL16278) '’floating form”; (Sd) Beaumaris, A. J. Tadgell, Mar. 1908 (MEL16264); (Sd) Dalmore. A. Meebold 21771, 12.2.1936 (NSW90538); Cranbourne Swamp, Dr. Patton, 13.11.1938 ( MELU) ; Gorae West, A. C. Beauglehole 7328, Nov. 1946 (HERB A.C.B.); Moora Moora Reservoir, Grampians, T. & J. Whaite 1513, 26.10.1953 (NSW90535) "aquatic herb, radical leaves, in swamp”; Beaufort, F. Swindley 41. 9.12.1959 (MEL 16255-6) "Growing both on wet land near the lake, and in the water. Petals yellow” "Leaves, of plant in water, floating”; Near Condah Swamp. A. C. Beauglehole 7050 and H. 1. Aston, 21.10.1960 (HERB. A.C.B.; MEL16251 -2); 5 miles S.W. of Casterton, H. 1. Aston 770, 23.10.1960 (MEL25444; AD961 10056) ; (Sd) N. of Wannon Bridge, Grampians, D. E. Symon 1757, 1.11.1961 (ADW25084); (Sd) Victoria Valley, Grampians, D. E. Symon 1808, 4.11.1961 (ADW25082) "Leaves floating”; (Sd) 4 miles S.W'. of Edenhope, F. Swindley 1187, 30.11.1962 (MEL255) "Small swamp”; Beaconsfield. H /. Aston 1228, 27.10.1964 (MEL16250); 4 miles N.E. of Morwell, E. F. Constable 5314, 28.10.1964 (NSW78582) "swampy ground” ‘‘flowers yellow” "Very localised”; (Sd) 4| miles S.S.E. of Yea, H. 1. Aston 1308, 7.12.1964 (MEL16253-4) ; 8 miles W. of Alexandra. H. /. Aston 1319, 8.12.1964 (MEL 16245-6) ; (Sd) Cranbourne, H. /. Aston 1328, 19.1.1965 (MEL16247-9) ; (Sd) 20 miles west of Bairnsdale, H. I. Aston 1344, 29.1.1965 (MEL16244): (Sd) 3 miles E. of Skipton, H. I. Aston 1393, 22.11.1965 (MEL16212-3) ; (Sd) 9 miles S.W. of Stawell, H. 1. Aston 1410, 23.11.1965 (MEL 16214); (Sd) Moora Moora Reservoir, Grampians, H. 1. Aston 1414, 24.11.1965 (MEL16215); (Sd) 16 miles W. of Casterton, H. 1. Aston 1418, 25.11.1965 (MEL162I6); (Sd) 4-5 miles N.N.W. of Dunkeld, H. I. Aston 1424, 26.11.1965 (MELl 6233-4) ; (Sd) 15 miles N.N.E. of Cavendish, H. 1. Aston 1426, 26.11.1965 (MEL16217-9) ; (Sd) 12.5 miles S. of Cherrypool, H. /. Aston 1428, 26.11.1965 ( MEL 16220-2 ) ; (Sd) Cherrypool, H. 1. Aston 1430, 26.11.1965 (MEL16235-6) ; (Sd) Lang Lang. H. I. Aston 1433, 10.12.1965 ( MEL16226-7) ; (Sd) Peterborough, H. I. Aston 1434, 13.12.1965 (MEL16223-5 ) ; (Sd) 4-5 miles W.N.W. of Gorae West, H. 1. Aston 1437, 14.12.1965 (MEL 16228); (Sd) Drik Drik, H. /. Aston 1440 and A. C. Beauglehole 6782, 14.12.1965 (MEL16229); (Sd) Heywood, H. 1. Aston 1444, 15.12.1 965 (MEL16230); (Sd) 6-6 miles W. of Dergholm. H. I. Aston 1446, 15.12.1965 (MEL16231); (Sd) 7-2 miles N.W. of Dergholm, H. /. Aston 1447, 15.12.1965 (MEL 16232); (Sd) 4 miles S.E. of Edenhope, H. /. Aston 1448, 16.12.1965 ( MELi 6237-9) ; (Sd) 6 miles W.S.W. of Horsham, H. 1. Aston 1452, 16.12.1965 ( MELl 6240-1 ) ; (Sd) 4 miles S.S.W. of Stradbroke, H. 1. Aston 1460, 13.1.1966 (MEL16242-3) ; (Sd) Penola to Dergholm road 1 km. on Victorian side of border, M. A. Fagg 118, 28.1.1966 (AD966100290; MEL25438); Port Campbell, /. Stone, 29.10.1966 (MEL20463-4) ; St. Leonards, H. I. Aston 1473, 2.11.1966 (MEL20468-71 ) ; 3 miles S.S.W. of Rosebud, H. 1. Aston 1481, 7.11.1966 (MEL20472) ; (Sd) 4 miles W. of Portland, H. I. Aston 1513, 13.12.1966 ( MEL20461 ) ; (Sd) Drysdale, A. Gibbon, 15-20.12.1966 ( MEL20473-4) ; (Sd) Darby River, Wilsons Promontory, J. H. Willis, 13.1.1967 (MEL20462) ; (Sd) Tooradin, H. /. Aston 1581, 3.3.1967 (MEL20433-4) ; (Sd) 6 miles N.N.W. of Anglesea, H. /. Aston 1582, 21.3.1967 (MEL20435); (Sd) Airey's Inlet, H. I. Aston 1583, 22.3.1967 (MEL20436); Stony Rises, 2j miles west of Pirron Yallock, H. /. Aston 1591, 11.4.1967 (MEL20437); Stony Rises, 3£ miles west of Pirron Yallock, H. /. Aston 1592, 11.4.1967 (MEL20438-40) . Tasmania — T asmania (MEL16314); Tasmania, R. C. Gunn, 1836 (NSW90544) ; Circular Head Neck, R. C. Gunn, in flower 10.12. 1 837, in fruit 23.1.1838 (NSW90541) ; George Town, R. C. Gunn, 22.10.1842 (HOB); (Sd) Helen I. Aston: The Genus Villarsxu ( Menyanthaceae ) in Australia 53 Tasmania, R. Gunn 1216/1842, 10.1.1843 (MEL16285; BR1061769); Van Die- mensland, R. Gunn (MEL 163 15); (Sd) Distillery Creek, R. C. Gunn , 1.1.1845 (HOB); (Sd) South Esk River, C. Stuart 167, 1848 or 9 (MEL16298); South Esk River, C. Stuart 273, 71849 (MEL16287); Barnard's Creek, 6 miles from Launceston, S. Hannaford 170, 26.11.1864 (MEL16280; HOB); (Sd) Tasmania, S. G. Hannaford (NSW90546); (Sd) Huon Road, S. G. Hannaford (HOB); (Sd) Tasmania, W . H. Archer (NSW90545); (Sd) Swanport, Dr. Story (MEL16310) ; (Sd) Between Bridgewater and New Norfolk, Feb. 1869 ( MEL16284) “in watercourses on marshes”; Lake St. Clair, T. Gulliver Mar 1873 (MEL16288) ; Lake St. Clair, T. & B. Gulliver, 1873 (MEL16291) “4000'"; (Sd) Huon Road, from A. Simpsons collection, Feb. 1878 (HOB); Cape Portland, Mrs. Bowen, Mar. 1885 (MEL16293); N. E. Tasmania, Mrs. Bowen, 1887 (MEL16295); Port Arthur. Rev. J. Bufton, 1892 (MEL16296); Tasmania! Rev. J. Bufton, 1892 (MEL16297); (Sd) Near Launceston, L. Rodway, Nov! 1894 (HOB); (Sd) Bellerive, A. H. S. Lucas, Dec. 1924 (NSW90543); (Sd) Little Hampton, F. H. Long 1133, 30.1.1932 (HOB) “Water herb” “height 2 ft." “Marsh lands” “Flow’er yellow” “Alt. 600 ft.”; St. Clair, W. M. Curtis, 8.2.1945 (MEL16321 ) “in swamp at HE. workings”; Scamander, H. D. Gordon, 16.11.1945 (MEL16322) “Marsh behind sand dunes”; Blackman River at Tun- bridge, W. M. Curtis, 14.12.1955 (HOB) “river flat”; (Sd) Meander River flats, 5 miles S. of Deloraine, J. Somerville, 2.1.1959 and 3.1.1959 (HOB) “swamp" “alt. c.800'”; (Sd) Between Lake Sorell and Great Lake, Hj. Eichler 16942, 6.2.1960 (AD96 109201 ) ; Cape Portland area, Mrs. M. Cameron, 6.11.1966 (MEL20467) ; (Sd) Rocky Cape, N. W. coast, J. H. Willis, 4.1.1967 (MEL20465-6) “abundant and luxuriant under shade of Melaleuca squarrosa in a heathland swamp . . 12. Villarsia umbricola H I. Aston sp. nov. Herba aquatica vel semiaquatica, erecta usque ad reclinatam, gracilis, caespitosa, perennis, plerumque 30-75 cm. alta, panicula patenti multifloraque praedita. Corolla flava, 11-31 (±21-7) mm. diam.; pedicelli capsulae maturae (5-) 10-40 (-62) mm. longi, leniter recurvantes; semina (0*8—) 1-1 -4 (-L6) mm. longa, pallide ochracea usque ad cinnamomeas, nunquam cinerea vel nigra A V. reniformi R.Br. recedit sic: absentia stolonum; laminis foliorum minus dorsiventralibus, tenuius textis, pallidius flavo-virentibus. undulatis etiamsi natantibus (turn denique super aquam non prone jacentibus) ; culmis tenuioribus, debilioribus, plerumque semireclinatis, pedicellis longis in verticillis opinatis (partum usque ad 6) saepe constipatis, recurvatis in capsulis maturis- diametro parviore floris expansi. In characteribus seminis etiani diflfert. Erect to reclining, slender, tufted perennial, without stolons. Radical leaves rising erect to spreading or reclining on land plants, erect to floating when in the water but then the blades remaining ±’ undulate and not lying flat on the water surface, on petioles 1-17 in. long; blades from elliptic (rarely) to ovate to broad-ovate (occasionally broader than long) to sometimes almost round, at times oblique, (1-) 2 5-8 (-12) cm. long x (0 7-) 1 5-6 (-1 1 ) cm. broad, apex obtuse to rounded, base from ± truncate or sometimes rounded to slightly or deeply cordate, the lobes mostly Y^-Ve of the leaf length but sometimes to sinus broad to narrow, edge entire and usually undulate, surfaces matt to semi-alossy, medium yellowish-green and ± equal in colouration or the upper surface a little darker, dotted beneath but not over-conspicuously, texture some- what thin. Culms several, (i-) 1-2| (-3i) ft. high, much lonaer than the radical leaves, slender, weak, semi-erect or reclining on both land 54 Helen I. Aston: The Genus Villarsia (M enyanthaceae) in Australia and water plants, usually supported by surrounding vegetation, the first branch arising from (V13-) (~i) of the culm height; cauline leaves 1 -several, subtending the panicle branches, the lower radical-like on Fig. 32 — V. umbricola, shape of radical leaves, x f. a to h, usual shapes found in both varieties, a to e, var. heaugleholei. a and d, Aston 1427. b and e, Aston 1445. c, Aston 1515. f to h, var. umbricola. f, Aston 1527. g, Aston 1529. h, AD96 102063. i, var. umbricola, Aston 1531. j to n, var. heaugleholei, Aston 1439 and A. C. Beauglehole 6781, reduced forms from plants in an open, roadside depression. Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 55 petioles from very short to 7 in. long, or sometimes all bractlike. In- florescence a slender, open, many-flowered panicle, branching dicho- tomously, each pedicel arising singly in the axil of a bract, or by reduction of the internodes the culm branches and pedicels arising in apparent whorls or near-whorls of up to 6 members, in small plants the panicle much reduced; final bracts ovate-elliptic, 3-5 mm. long; pedicels (of mature capsules) (5-) 10-40 (-62) mm. long, erect in flower, gently recurving as the fruits mature. Flowers homostylous; calyx-lobes (3-) 6-8 (-9) mm. long, narrow to broad elliptic-ovate, thinly mem- branous on the edges, %—% length of the corolla; corolla yellow, 7-15 mm. long, 11-31 (average 21 7) mm. span, lobes 5 or sometimes 4 or 6, variable in shape and apex (Fig. 2) without a longitudinal keel; gynoecium with 2 placentas, ovules 1 1-50 per placenta, style about 1 -5-2 mm. long, stigmas 2, papillose, ca. 0-5-1 mm. long, zb rotund to ovate and erect, held about level with the anthers; anthers 1 -5—2-5 mm. long, on short filaments 0 5—1 5 mm. long. Capsule (5-) 6-11 mm. long, from equal to the calyx-lobes to exserted beyond them by -3- (—£) their length (sometimes more), mostly noticeably exserted, chartaceous, globular- to ellipsoid-ovoid, adnate to the calyx-tube only at the base, dehiscing at the summit into 4 acute, somewhat-recurved valves. Seed ( 0 8—) 1-14 (-16) mm. long, pale cream to buff-fawn or light brown, never grey or black, compressed-ellipsoid to sub-cylindrical or sub- globular, without a conspicuous caruncle; surface slightly or strongly marked with minute “jig-saw” patterns, almost smooth to slightly granular, mostly glossy, also in the var. beaugleholei strongly tuberculate. var. umbricola. Semen paene glabrum usque ad parve granulatum. Seeds ellipsoid or irregularly so, compressed, double convex, mostly glossy; surface slightly or strongly marked with minute “jig-saw” patterns, almost smooth to slightly granular, without tubercles but some- times with some of the granules more pronounced and with several microscopic projections on the summit of each. var. beaugleholei H. I. Aston var. nov. Semen fortiter denseque tuberculatum. Seeds ellipsoid, somewhat compressed but turgid, at times almost sub-cylindrical or sub-globular; surface slightly or strongly marked with minute “jig-saw” patterns, and also strongly tuberculate, the tubercles hollow, to 0 75 mm. long, and each again minutely tuberculate at its summit, rarely the tubercles reduced to granules over the centres of the seed faces. Distribution : var. umbricola : — South Australia — Mt. Lofty Ranges from around Adelaide south to the extremity of the Fleurieu Peninsula, also Kangaroo Fsland and south-east of S.A. There is an outlying northern record from Wirrabara, north-east of Port Pirie. Victoria — Bridgewater Lakes west of Portland. 56 Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia var. beaugleholei : — South Australia — extreme south-east. Victoria — south-west, in the vicinity of the Grampians and the Glenelg and Wannon Rivers system. Habitat : Typically sheltered, shaded or semi-shaded situation on damp soil or in shallow fresh waters, sometimes in water to 18 inches deep. On damp soil in Eucalyptus obliqua and E. ovata forests (var. beaugleholei ), commonly edging or beside small creeks and in bogs, also in or beside Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 57 swamps. Mostly sheltered by taller shrubs and vegetation but sometimes in the open and then growing well if in water but suffering reduction if on damp soil only. Some associated species recorded are Leptospermum juniperinum, L. lanigerum, L. obovatum, Melaleuca decussata, M. squamea, M. squarrosa, Olearia glandulosa , Gahnia sp., Resdo tetraphyllus, Lepidos- perrna laterale, Car ex appressa, Schoenus tesquorum, Juncus articulatus, Goodenia ovata, Selliera radicans, Haloragis brownii, Hydrocotyle sp.. Ranunculus rivularis, Lobelia alata, Gratiola peruviana , Epilobium sp., Triglochin striata, Drosera binata, Blechnum sp., and Myriophyllum pedunculatum. Flowering time: var. umbricola : — FI. and Fr. recorded Nov. -April. var. beaugleholei : — FI. and Fr. recorded Nov.-Dee., but season apparently extends further as there are still many buds on plants seen in December. Typification : var. umbricola : — Holotype — (Sd) Ca. 4 miles direct line E.S.E. of Myponga, between Myponga and Hindmarsh Valley, H. I. Aston 1531 , 18.12.1966 (MEL20485). Isotypes — AD, NSW, K. Paratypes — (Sd) Mt. Lofty Railway Enclosure, just south of Mt. Lofty Rly. Stn., H. 1. Aston 1532, 19.12.1966 (MEL20484 and AD, NSW, PERTH, K, BM); (Sd) Callawonga Creek, about 1 3 road miles south of the Victor Harbor to Cape Jervis road, H. 1. Aston 1530, 18.12.1966 (MEL20486 and AD, NSW, K). The second paratype has the seeds rougher, with more pronounced granules, than have the other designated types. var. beaugleholei : — Holotype — (Sd) 13 miles west of Dergholm, along the road to Penola, H. 1. Aston 1445, 15.12.1965 (MEL16415). Isotypes— MEL 164 16- 17 and AD, NSW (2 sheets), K, BM, PERTH. Paratypes— (Sd) Ca. 4 5 miles N.N.W. of Dunkeld and 2 miles S. of Victoria Valley township, H. 1. Aston 1423, 26.11.1965 (MEL16422-23 and AD, NSW, K, BM, PERTH, CANB, L); (Sd) Malseeds Lake, ca. 2 miles west of Mt. Richmond Post Office, H. 1. Aston 1514, 13.12.1966 (MEL20480-81 and AD, NSW, K, BM). The second paratype shows smaller foliage than the other types selected, but lies well within the range of variation shown by the variety. Discussion : The erroneous recording by Black (1957, p. 693) of V. parnassifolia in South Australia seems to have been based on misidentiffcation of this previously unrecognised entity, V. umbricola. V. umbricola overlaps the range of V . reniformis and cannot always be distinguished from it by herbarium specimens alone as these have not always been collected adequately and seldom indicate flower measurements. Flowers themselves are rarely collected, and even if 58 Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia present with specimens are usually valueless because of their deliquescent nature. However, populations in the field are readily distinguished when flowering or fruiting, and can often also be separated when vegetative only. The less dorsi-ventral, thinner-textured, paler, usually more yellowish-green foliage, with blades not lying flat on the water surface even when floating, the slender, semi-reclining, weak culms with long pedicels often partially grouped in apparent near-whorls, the slender pedicels recurved in fruit, and the smaller flowers are features which separate V . umbricola from V . reniformis. In addition, the tuberculate seeds of var. beaugleholei are quite distinct from the smooth seeds of V . reniformis. The seeds of var. umbricola however, although tending to be less compressed and more granular than those of V. reniformis. are not considered reliably distinctive in all cases. The two varieties of V . umbricola are distinguished by seed alone. Within Victoria all collections of var. beaugleholei show the strongly tuberculate surface and generally are rather turgid and semi-cylindrical. At Rendelsham, between Millicent and Beachport in South Australia, both varieties were found growing intermingled in an extensive popula- tion (Aston 1520-23). Seeds of var. beaugleholei here showed a trend in characters towards those of var. umbricola, being more compressed and having the tubercles shorter and occurring only around the borders and face edges, the centres of the seed faces being granular. Some plants, however, retained the strongly tuberculate seed surface all over. Seeds of var. umbricola were either typically granular or had the granules around the borders and face edges somewhat more pronounced with each bearing microscopic projections on its summit suggestive of those occurring on the summits of the tubercles of var. beaugleholei. This variation in the degree of smoothness or the pronouncement of the granules of var. umbricola seeds occurs right throughout its range. Measurements made have shown tendencies for var. beaugleholei to have slightly smaller seeds and smaller flower span than var. umbricola. As the number of populations investigated is small, and the number of individuals available in each varied considerably, these tendencies should not be considered significant at this stage. They require fuller investiga- tion. Even if biologically significant, the different seed and flower span sizes are of no value for identification purposes as the extremes of measurement in each case are almost identical for both varieties, and the norms lie close together. Measurements are: — var. beaugleholei, flower span 11-29 (average 20 1 ) mm. for 82 individuals of 10 collec- tions, seeds (0 8-) 1 — 13 (-16) mm. for 639 indiv. of 20 collections; var. umbricola, flower span 14-31 (average 23 7) mm. for 69 indiv. of 6 coll., seeds (0-9- ) 1—14 (-16) mm. for 774 indiv. of 23 coll. Figures 1 and 5 should also be consulted. The distinctive turgid and rough seeds of the present V. umbricola var. beaugleholei were commented on by Mueller almost exactly a century ago when he wrote (Mueller 1868, p. 140) of a plant from the Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 59 Wannon River in the Grampians, Victoria. At that time he retained the collection concerned under V. reniformis, an understandable decision as he had no further specimens. The specific epithet is given in reference to the usual habitat preferred by this species as a whole, while the epithet of the var. beaugleholei is bestowed in honour of Mr. (Cliff) and Mrs. (Hilda) A. C. Beauglehole of Gorae West, via Portland, Victoria. Without Cliff’s splendid botanical knowledge and their combined assistance in offering local guidance and friendly hospitality on several occasions I could never have investigated the villarsias of their district so successfully. Specimens examined (excluding types) : var. umb r l col a : — South Australia — (Sd) Macclesfield, and Torrens River, F. Mueller, Nov. 1849 (MEL16404); (Sd) Torrens near Port Adelaide, Blandowski, 1850 (MEL16400) ; (Sd) Clarendon, Tepper 398, 1881 (MEL16398); Wirrabara, ca. 30 km. N.E. of Port Pirie, Herb. R. Tate, 19.11.1881 (AD966 1 00354 ) ; (Sd) Myponga, Herb. J. M. Black, 16.11.1922 (AD966100358) ; (Sd) Tooperang. C. M. Eardley, Dec. 1937 (ADW); (Sd) Callawonga Creek. F. M. Hilton , 25.11.1953 (ADW18798); (Sd) Mt. Compass, along the Nangkita road. D. J. E. Whibley 29, 15.11.1956 (AD957 12020) ; (Sd) Boat Harbour Creek, between Cape Jervis and Victor Harbour, Hj. Eichler 14417, 16.11.1957 ( AD95826012) : (Sd) 3 miles S. of the intersection of the Tunkalilla and the Victor Harbour- Cape Jervis roads, R. Schodde 601, 25.1.1958 (AD96029040; CAN BLR) ; (Sd) Deep Creek near mouth, Fleurieu Peninsula, E. N. S. Jackson 13, 11.1.1959 (AD96 102063) ; (Sd) Lake Bonney, 10 km. S.W. of Millicent, P. Wilson 1179 , 13.11.1959 (AD96020270; MEL16402); (Sd) Reservoir near Mt. Lofty Station. E. H. I sing, 22.3.1961 ( AD96151 103) ; (Sd) Reservoir near Mt. Lofty railway station, E. H. Ising, 8.12.1961 ( AD96207255 ) ; (Sd) At reservoir near Mt. Lofty railway station, E. H. Ising, 22.12.1962 (AD96421083) ; (Sd) Mt. Lofty railway reservoir enclosure, Hj. Eichler 17147, 5.1.1963 (AD96445390; MEL16401); (Sd) Between Myponga and Hindmarsh Valley, N . N. Dormer 950, 3.12.1963 (AD96414104: MEL16403); (Sd) Rocky River, Flinders Chase, Kangaroo Is., Hj. Eichler 18577, 5.1.1966 (AD9661 10341 ; MEL16399); (Sd) Strangw ; ays Falls, just N.E. of Inman Valley, D. J. E. Whibley 1731, 23.11.1966 (AD9665 1041 ; MEL20483) ; (Sd) Strangways Falls, just N.E. of Inman Valley, D. J. E. Whibley 1740, 23.ll.1966 ( AD9665 1 042; MEL20482); (Sd) 4-5 miles N.W of Rendef- sham, H. I. Aston 1521, 16.12.1966 (MEL20491); (Sd) 4-5 miles N.W. of Rendelsham, H. 1. Aston 1523, 16.12.1966 (MEL20490); (Sd) 2-5 miles ESE of Robe, H. 1. Aston 1527, 16.12.1966 (MEL20489); (Sd) Finnis, H. 1. Aston 1528, 17.12.1966 (MEL20488); (Sd) Willow Creek, 10 miles direct line west of Victor Harbour, H. 1. Aston 1529, 18.12.1966 (MEL20487). Victoria— (Sd) Bridgewater Lakes, about 11 miles west of Portland. F. Swindley 1590, 4.4.1963 (MEL1177); (Sd) Bridgewater Lakes, H. 1. Aston 1443 and A. C. Beauglehole 6783, 14.12.1965 (MEL1 6397-7A; Herb. A C B ) (Sd) Bridgewater Lakes, H. 1. Aston 1516, l4.l2.iVb6 ( MEL20492-3 ) . var. beaugleholei : — South Australia — (Sd) Between Kalangadoo and Millicent, J. B. Cleland 6.12.1922 (AD966 100417) ; (Sd) 4-5 miles N.W. of Rendelsham, H. 1. Aston 1520, 16.12.1966 (MEL20476); (Sd) 4-5 miles N.W, of Rendelsham, H. / Aston 1522, 16.12.1966 (MEL20475). Victoria— (Sd) Banks of the Wannon, C. Wilhelmi, Dec. 1856 (MEL16408L (Sd) Wannon, Coll. ? Wilhelmi or F. Mueller, (MEL16409); (Sd) Lower Glenele River, Eckert 62, 1891 (MEL16407); (Sd) Scrubby Creek. Victoria Valiev Grampians, H. 1 . Aston 1412, 24.11.1965 (MEL16419-21 ) ; (Sd) Ti-tree Creek 12-5 miles S. of Cherrypool, H. I. Aston 1427, 26.11.1965 (MEL16410-2) ■ (Sd)’ 60 Helen 1. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia Cherrypool, H. I. Aston 1429, 26.11.1965 (MEL1641 3-4) ; (Sd) 6 miles N.W. of Portland. H . I. Aston 1436, 13.12.1965 (MEL16424); (Sd) 4-5 miles W.N.W. of Gorae West School, H. I. Aston 1438 and A. C. Beauglehole 6780, 14.12.1965 (MEL 16426-7; Herb. A.C.B.); (Sd) 2-5 miles N.N.E. of Mt. Kincaid, H. I. Aston 1439 and A. C. Beauglehole 6781 , 14.12.1965 (MEL 16425; Herb. A.C.B.); (Sd) Drik Drik, H. 1. Aston 144J, 14.12.1965 (MEL16428); (Sd) Balmoral, H. I. Aston 1451, 16.12.1965 (MEL 1641 8); (Sd) Lake Bung Bung, 10 miles E. of Nelson, A. C. Beauglehole 22202, 12.6.1966 (Herb. A.C.B.; MEL25466); (Sd) Branch Creek, Victoria Valley, Grampians. A. C. Beauglehole 22, 201, 10.12.1966 (Herb. A.C.B.: MEL25465); (Sd) Glenelg River, 3 miles S.S.E. of Mt. Thackeray, Grampians. A. C. Beauglehole 22200, 10.12.1966 (Herb. A.C.B.); (Sd) Long Swamp, Nelson, H. I. Aston 1515, 13.12.1966 ( MEL20477-9) ; (Sd) Malseed's Lake, ca. 2 miles west of Mt. Richmond Post Office, A. C. Beauglehole 22203, 13.12.1966 ( MEL25505-6; Herb. A.C.B.). Variety uncertain as all specimens lack seeds ; — South Australia — North Adelaide, F. Mueller, 23.1.1848 (MEL16405): Lake Bonney, Miss Wehl, 1887 (MEL16406); Aldgate, Ashby, 19.11.1895 ( AD966041068) ; Eight Mile Creek. South East, C. M. Eardley, 4.2.1942 (ADW); Port MacDonnell, C. M. Eardley, 5.2.1942 (ADW); Callawonga Creek, Fleurieu Peninsula, D. E. Symon, 25.11.1953 (ADW18621); Edinburgh Swamp between Myponga and Hindmarsh Valley, Hj. Eiclder 12195, 23.2.1956 ( AD95903044) . Victoria — Fyans Creek, Grampians, 7. H. Warcup, 24.1.1961 (ADW23750); ± 7 miles N.W. of Zumsteins on main road to Horsham, D. E. Symon 1642, 30.10.1961 (ADW25085); Fyans Creek, Grampians, D. E. Symon 1852, 7.11.1961 (ADW25083); Near Glenelg River crossing on Casterton-Edenhope road, F. Swindley 1138, 29.11.1962 (MEL254). ACKNOWLEDGEMENTS I wish to thank all the herbarium directors and keepers who have made the specimens of their institutions available for my examination. I would also like to thank more specifically the following people who have aided me in this study — Dr. W. T. Stearn, Sir G. Taylor, Professor G. Moggi, and Dr. R. Weibel for assistance in locating and comparing types, and for advice on typification; Mr. T. E. Burns of Launceston who discovered the only location where V . exaltata is known to occur in Tasmania today, and who forwarded material of the species to Mel- bourne; Mr. and Mrs. A. C. Beauglehole of Gorae West for their hospi- tality and assistance in the field; Mr. R. D. Royce of the Western Aus- tralian Herbarium for informing me of his discovery of a new species of Villarsia in Western Australia and for giving me the opportunity to describe it; Dr. Hj. Eichler of the State Herbarium of South Australia for extending the facilities of his herbarium during a collecting trip and for helpful discussion; Mr. R. T. M. Pescott, Director of the National Herbarium of Victoria, for allowing me to undertake this work, and in particular Mr. J. H. Willis, Assistant Government Botanist of the National Herbarium of Victoria, for preparation of the latin diagnoses, for reading of the manuscript, and for his patient and willing advice extended on many occasions. Finally my gratitude is extended to Miss Barbara Taylor, Research Assistant employed at the National Herbarium of Victoria by the Botanic Gardens Branch Research Trust, who has cheerfully performed many somewhat tedious tasks in caring for all collections and taking seed measurements, and who has prepared all maps and graphs. Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia 61 REFERENCES Beadle, N. C. W., Evans, O. D., and Carolin, R. C. ( 1962)— Handbook of the Vascular Plants of the Sydney District and Blue Mountains (Armidale). Bentham, G. (1868)— Flora Australiensis 4 (London). Black, J. M. (1957)— Flora of South Australia, pt. 4, ed. 2 (Adelaide). Burbidge, N. T. (1963) — Dictionary of Australian Plant Genera (Sydney). Eichler, Hj. ( 1965)— Supplement to J. M. Black’s Flora of South Australia (second edition 1943-1957) (Adelaide). Hooker, J. D. (1860) — Flora Tasmaniae 1 (London). Hutchinson, J. (1960) — The Families of Flowering Plants 1, ed. 2, reprint (Oxford). Merrill, E. D. (1938) — A critical consideration of Houttuyn's new genera and new species of plants, 1773-1783. J. Arnold Arbor. 19 (4) : 291-375. Mueller, F. von (1868) — Fragmenta Phytographiae Australiae 6 (Melbourne). Ornduff, R. (1966) — The origin of dioecism from heterostyly in Nymphoides (Menyanthaceae). Evolution 20 (3): 309-314. 62 Helen I. Aston: The Genus Villarsia ( Menyanthaceae ) in Australia Plate 1 NATIONAL HERBARIUM Of VICTORIA $ll£hns