JOURNAL of the ADELAIDE BOTANIC GARDENS

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Volume 245 LELIN Government of South Australia Botanic Gardens and : q. +; Department of Environment State Herbarium 10 July 2012 is and Natural Resources

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A | OQOURNAL of the ADELAIDE

BOTANIC GARDENS

AN OPEN ACCESS JOURNAL FOR AUSTRALIAN SYSTEMATIC BOTANY

flora.sa.gov.au/jabg

Published by the STATE HERBARIUM OF SOUTH AUSTRALIA on behalf of the BOARD OF THE BOTANIC GARDENS AND STATE HERBARIUM

State Herbarium of South Australia

PO Box 2732

Kent Town SA 5071

Australia

Board of the

Botanic Gardens and

State Herbarium

a)

Government — of South Australia

Department of Environment, Water and Natural Resources

J. Adelaide Bot. Gard. 25 (2011) 1-4 =)

Gahnia halmaturina (Cyperaceae: Schoeneae), a new species from Kangaroo Island, South Australia

Russell L. Barrett®":° & Karen L. Wilson?

“Botanic Gardens and Parks Authority, Kings Park and Botanic Garden, West Perth, 6005, Western Australia E-mail: russell.barrett@bgpa.wa.gov.au ’ School of Plant Biology, Faculty of Natural and Agricultural Sciences, The University of Western Australia, Crawley, 6009, Western Australia “Western Australian Herbarium, Department of Environment and Conservation, Locked Bag 104, Bentley Delivery Centre, 6983, Western Australia

“National Herbarium of NSW, Royal Botanic Garden, Mrs Macquaries Road, Sydney 2000, New South Wales

Abstract

As a precursor to the fifth edition of the Flora of South Australia, Gahnia halmaturina R.L.Barrett & K.L.Wilson is described as a new species from Kangaroo Island. This species was previously known by the phrase name Gahnia sp. West Bay (B.M.Overton 2685) R.L.Taplin. A revised key to South Australian species of Gahnia 1s provided.

Keywords: Cyperaceae, Schoeneae, Gahnia, South Australia, Kangaroo Island, taxonomy.

Introduction

Systematic studies in Cyperaceae, tribe Schoeneae, have shown that there are a number of undescribed taxa in many of the Australian genera. A potentially new species on Kangaroo Island was first collected in 1998 by Bev Overton and was unknown when the previous edition of the Flora of South Australia (Jessop & Weber 1986) was published. Rosemary Taplin gave it the informal name Gahnia sp. West Bay (B.M. Overton 2685) R.L.Taplin in the South Australian Herbarium (Barker et al. 2005). Study of herbarium material of this taxon and comparison with collections of all other named Australian species of Gahnia J.R.Forst. & G.Forst. have confirmed that it represents a novel species. Molecular data have confirmed the relationships of this taxon (Barrett et al., unpubl.). The description is based on herbarium material.

Gahnia halmaturina R.L.Barrett & K.L.Wilson, Sp. NOV.

Gahniae hystrici /.M Black similis, sed habitu altiore, inflorescentia non contracta, foliis longioribus flexuo- sisque non pungentibus, antheris quattuor, differt. Gahniae microstachyae (R.Br) Benth. similis, sed foliis non applanatis, spiculis grandioribus pallidisque distinguitur. Ab ambobus speciebus foltis culmisque papillatis distinguitur.

Typus: South Australia: Kangaroo Island: Flinders Chase National Park, West Bay Creek S Tributary, 2.3 km E of West Bay, N side of West Bay track, 35°53'20"S, 136°37°50”E, 12 Apr. 1999, B.M. Overton 2744 (holo.: AD 99920049; iso.: NSW).

Gahnia sp. West Bay (B.M.Overton 2685) R.L.Taplin in W.R. Barker et al., J. Adelaide Bot. Gard. Suppl. 1: 189 (2005).

Tufted perennial with short, compact rhizomes. Culms and leaves spirodistichous; leaf to culm length ratio 0.6—0.9:1. Leaves flexuose, erect to spreading, margins tightly inrolled when dried, sometimes appearing terete with a groove on one side, finely striate, dull green (pale to cream at base), not glaucous, covered in very fine papillae, silica bodies prominent on upper surface, 23—70 cm long, 0.7—1.8 mm wide (when dried and margins inrolled), 0.2—0.5 mm thick (when flat); margins very finely scabrous due to papillae; sheath 3.5-10 cm long, pinkish brown to dark chocolate- brown, finely striate and covered in very fine papillae; ligule a pale band of tissue, glabrous, c. 0.5 mm high. Culms terete, often paler than leaves, otherwise as for leaves, 14-65 cm tall, 0.7—1.7 mm diam. /nflorescence + linear in outline, 7-16 cm long, 7—20 mm wide, with several lateral branches, | lateral branch per node; basal lateral branch 18—95 mm long (note majority of peduncle is hidden by sheath of involucral bract) with 9—19 spikelets; primary involucral bract 20-31 cm long, leaf-like. Spikelets 3.6-4.4 mm long, the upper flower bisexual, the lower flower functionally male; glumes c. 10, pale to dark brown, darkest at the keel, glabrous, apex obtuse, mucronate; lowest 6 glumes sterile; fertile upper glumes 2.5—3.1 mm long, c. 1.1 mm wide (when incurved). Stamens 4; anthers 1.3—1.7 mm long, 0.3—0.5 mm wide, apical appendage 0.2—0.3 mm long; filaments

J. Adelaide Bot. Gard. 25 (2011)

R.L. Barrett & K.L. Wilson

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Fig. 1. Gahnia halmaturina: A inflorescences; B culm bases. Scale bar: 1 cm. B.M. Overton 2744.

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J. Adelaide Bot. Gard. 25 (2011)

3.0—3.7 mm long at maturity. Style 3-fid (occasionally 1 of these further branched dichotomously, producing a 4-fid style), undivided portion c. 1.9 mm long, branches 1.6—2.2 mm long. Nut not held by the staminal filaments at maturity, cream to pale brown, smooth, narrowly ovate in outline, tapering towards the base, terete in cross-section to obscurely 3-angled, 3.0—3.3 mm long, 1.05—1.18 mm wide; epidermal cells narrowly oblong to subhexagonal or almost linear in outline. Figs 1, 2, 3.

Distribution and habitat. Endemic to Kangaroo Island, apparently restricted to the vicinity of West Bay in Flinders Chase National Park. This species is only known from the margins of freshwater creeks, growing on rocky areas along the banks and floodplains.

Phenology. Flowering recorded for April-May. Fruit collected in April, May and August.

Conservation status. Listed as ‘Rare’ (under its phrase name synonym) in Schedule 9 of National Parks and Wildlife 1972, 2008 Amendment (South Australia), as reflected in Barker et al. (2005). Based on collection notes, at least 1,200 individuals are known from several subpopulations over a distance of about 1.5 km. Given the limited distribution, the current listing 1s considered to be appropriate.

Etymology. Derived from Halmaturus, a generic name once applied to kangaroos (Smith 1870), ultimately derived from the Greek halme, a leap or bound, and commonly used as an epithet for species from Kangaroo Island.

New species of Gahnia (Cyperaceae)

Notes. Molecular data place this species as related to Gahnia hystrix J.M.Black, also endemic to Kangaroo Island, and G. trifida Labill. (Barrett 2012). G. halmaturina and G. hystrix are very easily distinguished by their growth habit and habitat, G. hystrix being a small cushion-forming plant growing on limestone, while G. halmaturina 1s a much taller, clumping plant growing on creek margins. The two are somewhat morphologically similar in having leaves exceeding the inflorescence and relatively few, large spikelets. Gahnia trifida is easily distinguished in the field from G. halmaturina by its much larger stature.

This species is superficially similar to Gahnia microstachya (R.Br.) Benth. from New South Wales, Victoria and Tasmania, differing from that species in the inflorescence being held below the level of the leaves, the leaves not clearly flattened and spread more evenly along the culms rather than all strongly clustered at the base, and the larger, paler spikelets. G. microstachya prefers dry rocky sites (Wilson 1993).

Both Gahnia hystrix and G. microstachya \ack the distinctive papillae found on the leaf blades and sheaths and culms of G. halmaturina.

Other specimens examined.

SOUTH AUSTRALIA. Kangaroo Island: Track to Cape Borda from West Bay, 1.7 km N of gate from West Bay track (200 m upstream), 3 May 2006, D./J. Duval 471, A. Quarmby & R.T. Taplin (AD; K n.v.); Flinders Chase National Park, 2.3 km E of West Bay camping ground, 2nd West Bay creeklet, 12 Feb. 1998, B.M. Overton 2685 (AD, PERTH); 12 Aug. 1998, B.M. Overton 2697 (AD, PERTH); West Bay, Flinders Chase National Park, at West Bay Creek fork, almost 2 km E

Key to Gahnia species in South Australia (adapted from Jessop & Weber 1986)

1. Inflorescence held within plant, shorter than the leaves, small, not paniculate; spikelets 4-19 2. Spikelets 9-19 in a shortly branched inflorescence; plant to 70 cm tall... 20... . ee ee G. halmaturina 2: Spikelets c, 4a asumple-spike-hke- imflorescenee? Gwark Plants ...4 ac. 2.59 5ach eleehs, exe ack dood bpp eeea elas ore a G. hystrix 1: Inflorescence distinctly longer than the leaves (occasionally less in G. ancistrophylla but then leaf tips are

distinctively coiled), paniculate; spikelets numerous

3. Mature nut adhering to the persistent staminal filaments; culms stout, usually 1-2 m high; glumes and bracts broadish, never long-acuminate; nut 2-4 mm long, shining

4. Glumes 12-17, the lowest 7-10 much shorter, all more or less obtuse, the lowermost without awns ......... G. clarkei 4: Glumes 5-10, of similar length, mostly acute, the lowermost long-mucronate ...................005. G. sieberiana

3: Mature nut never held by the staminal filaments; culms slender, usually 15—100 cm high; at least the outer bracts long-acuminate; nut c. 2 mm long or less, often dull or greyish

5. Spikelets 1-flowered, densely clustered

6. Lower leaf-like bracts of inflorescence smooth or minutely scabrid, conspicuously reddish-purple below; nut c. 5 mm long, more or less linear to very narrow-ovate in outline, triquetrous in cross-

section; pale vellow-browiiie veryidark BLOW .. as exc wes a Re SE CAN Oa EO Ee FR A G. filum 6: Lower leaf-like bracts of inflorescence coarsely scabrid, green or slightly yellow-tinted below; nut 2—2.5 mm long, narrow-obovate to narrow-elliptical in outline, trigonous in cross-section, grey to black .. G. trifida 5: Spikelets 2-flowered, more or less loosely arranged 7. Ligule a dark brown chartaceous band to 2 mm wide; panicle broadly oblong with branches somewhat SPrEACIne Or EV et CUTS me ree Sacetn dence eee cle FE WS EOE 2 Satin pei ad dom ODE SE OF AOE SMe pbk idiy ate EATS WE OEE G. radula 7: Ligule ciliate to woolly with hairs 2—4 mm long; panicle linear or linear-lanceolate with branches stiffly erect $.. Leaves very scabrous-at-least-on the marems m the-wpper halt oz... cscescetelsece gre aed Hoop tate elena eee 4 G. deusta 8: Leaves smooth 9, Leaves with curved or coiled filiform tips; leaf sheaths dull ...........0... 0... .0......005. G. ancistrophylla

9: Leaves with straight tips; basal leaf sheaths shining at least in part..... 20... 0.0... 0. cee eee G. lanigera

R.L. Barrett & K.L. Wilson

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J. Adelaide Bot. Gard. 25 (2011)

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STATE HERBARIUM OF SOUTH AUSTRALIA (AD) Cyperacese ( ADI9O92004G

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Fig. 3. Holotype of Gahnia halmaturina (AD), B.M. Overton 2744. Scale bar: 5 cm.

of west coast, 11 Apr. 1999, B.M. Overton 2743, D. Walters, C. Graham & S. Graham (AD); Flinders Chase National Park, West Bay Creek, upper portion of S tributary, 400 m E of gate to old Cape Borda track, 12 Apr. 1999, B.M. Overton 2745, D. Walters, C. Graham & S. Graham (AD, NSW); Flinders Chase National Park, West Bay Creek, N tributary, 1.7 km N of West Bay track on old Cape Borda track, 12 Apr. 1999, B.M. Overton 2746, D. Walters, C. Graham & S. Graham (AD); Southern tributary of West Bay creek, 2.3 km E of West Bay camp ground northern side of West Bay track, Flinders Chase National Park, 12 Apr. 1999, B.M. Overton BM 2749, D. Walters, C. Graham, S. Graham (AD, PERTH: 2 sheets).

Acknowledgements

We would like to thank Rosemary Taplin, Jiirgen Kellermann, Bill and Robyn Barker, and Helen Vonow for assistance at AD, and National Parks South Australia staff for assisting with associated fieldwork in South Australia. Neville Walsh (MEL) is thanked for taking RLB to examine G. microstachya in the field in Victoria.

References

Barker, W.R., Barker, R.M., Jessop, J.P. & Vonow, H.P. (eds.) (2005). Census of South Australian vascular plants: edition 5.00. Botanic Gardens of Adelaide & State Herbarium: Adelaide). [Journal of the Adelaide Botanic Gardens Supplement 1|.|

Barrett, R.L. (2012). Systematic studies 1n Cyperaceae tribe Schoeneae: Lepidosperma and allied genera. PhD thesis, The University of Western Australia.

Jessop, J.P. & Weber, J.Z. (1986). Cyperaceae. In: Jessop, J.P. & Toelken, H.R. (eds), Flora of South Australia 4: 2001- 2053. (South Australian Government Printing Division: Adelaide.)

Smith, W. (1870). Dr Smiths smaller English-Latin dictionary. (John Murray: London).

Wilson, K.L. 1993. Cyperaceae. In: Harden, G.J. (ed.), Flora of New South Wales 4: 293-396. (University of New South Wales Press: Kensington. ).

A | OQOURNAL of the ADELAIDE

BOTANIC GARDENS

AN OPEN ACCESS JOURNAL FOR AUSTRALIAN SYSTEMATIC BOTANY

flora.sa.gov.au/jabg

Published by the STATE HERBARIUM OF SOUTH AUSTRALIA on behalf of the BOARD OF THE BOTANIC GARDENS AND STATE HERBARIUM

State Herbarium of South Australia

PO Box 2732

Kent Town SA 5071

Australia

Board of the

Botanic Gardens and

State Herbarium

a)

Government — of South Australia

Department of Environment, Water and Natural Resources

J. Adelaide Bot. Gard. 25 (2012) 5—10

Boletus edulis (Boletaceae), a new record for Australia

Pamela S. Catcheside*’ & David E.A. Catcheside’

“State Herbarium of South Australia, DENR Science Resource Centre, P.O. Box 2732, Kent Town, South Australia 5071 E-mail: pam.catcheside@sa.gov.au

’ School of Biological Sciences, Flinders University, P.O. Box 2100, Adelaide, South Australia 5001 E-mail: david.catcheside@flinders.edu.au

Abstract

Fungi belonging to Boletus section Boletus, known in Italy as porcini, in France as ceps, are prized for their good flavour and command a high price. They are ectomycorrhizal, forming symbiotic associations with a wide range of host plants, and although endemic to the Northern Hemisphere have been introduced into South Africa and New Zealand. We report here that Boletus edulis Bull. has become established broadly across the higher parts of the Adelaide Hills in South Australia in mycorrhizal association with at least three species of exotic trees. ITS sequence and morphological data were utilised to confirm the identity of the fungus and a full description of the

Adelaide Hills collections 1s provided.

Keywords: naturalised and introduced fungi, edible fungi, Boletus, porcini, ceps, Quercus,

Pinus, Castanea, Australia.

Introduction

Boletus section Boletus comprising Boletus edulis and several allied species are of significant economic importance due to their excellent flavour both when fresh and dry and to their high nutritional value (Caglarlrmak et al. 2002; Ribeiro et al. 2008; Sitta & Floriani 2008). Annual worldwide consumption of porcini has been estimated to be between 20,000 and 100,000 tons (Hall et al. 1998) with a world market value exceeding $250 million. Prices range from $20-$80/kg in the northern hemisphere autumn but have reached as high as $231/ kg in a poor fruiting season in New York in 1997 (Hall et al. 2003).

Boletus sect. Boletus is characterised by the spongy hymenial surface of tubes and pores which 1s initially whitish due to the pores being covered with a white hyphal mass. The tubes and pores become yellow to olive-yellow and lose the hyphal web on maturity. The flesh does not discolour on cutting or bruising. The stipe is usually bulbous and has a raised reticulum, at least on the upper part. The pileus is hemispherical when young and brown to chestnut in colour (Watling 1970; Singer 1986; Breitenbach & Kranzlin 1991; Cortecuisse 1999; Beugelsdiyjk et al. 2008).

Species in Boletus sect. Boletus are ectomycorrhizal, erowing in association with a wide range of tree species in the families Fagaceae, Betulaceae, Malvaceae, Cistaceae, Salicaceae and Pinaceae (Hall et al. 1998; Agueda et al. 2006; Beugelsdijk et al. 2008; Dentinger et al. 2010). Boletus sect. Boletus is endemic in the northern hemisphere and does not occur naturally in the southern hemisphere. However, B. edulis has been

introduced into South Africa (Marais and Kotze 1977; Hawley 2008) and New Zealand (Wang et al. 1995; Hall et al. 1998; Stringer et al. 2001). B. edulis has not previously been reported from Australia (Watling & Li 1999; Roy Halling, pers. com., 2010) although an endemic true porcini with the provisional generic name “J/nferiboletus”, estimated to have diverged from Boletus 34 Mya, has recently been reported for Australia (Dentinger et al. 2010).

We first observed and collected material (PSC 2651) consistent with the macroscopic and microscopic characters of Boletus sect. Boletus in South Australia in May 2007 associated with Quercus robur L., although it is likely to have been present and known to European immigrants somewhat earlier. More material was found fruiting in May 2009, April 2010 and March 2011 (Fig. 1) at the same and a nearby site and another collection (PSC 3004) was made. Further reports in April and December 2010 of porcini-like boletes associated respectively with Castanea sativa Mill. and Pinus radiata D. Don, led to more collections (PSC 3273 and PSC 3458). Following early autumn rain in 2011, a report appeared in the Adelaide press on 2 Apr. 2011 of porcini being collected in the Adelaide Hills and sold at up to $120/kg in the Adelaide Market (Wilkinson 2011). A short survey on 12 Apr. 2011 led to finding a further six locations in the Adelaide Hills where the fungus was associated with QO. robur or P. radiata. A full evaluation of the collections was made due to the importance of porcini as a high value commercial crop and _ their potential to form mycorrhizal associations with a wide range of tree species.

P.S. Catcheside & D.E.A. Catcheside

J. Adelaide Bot. Gard. 25 (2011)

Fig 1. Boletus edulis. Adelaide Hills, March 2011, under Quercus robur. See back cover of journal for colour reproduction of this photograph.

Methods

Morphology

Habitat and associated plant communities were noted in the field. Collection locations were recorded by GPS (Garmin GPS12) and in situ photographs taken using a Nikon 4500 camera. Macroscopic characters were described directly from fresh material. Colours are described in general terms and more precisely according to the Royal Botanic Gardens Edinburgh Colour Chart (1969) (given as colour descriptor and number e.g. rust 13) and Kornerup & Wanscher (1978) (page number, column letter, row number e.g. 2B4). Fresh material was dried in a food dehydrator at 35°C for 24 h (Hydraflo LOOOFD).

Sections of fresh and dried material were mounted in 5% aqueous KOH, then stained with ammoniacal Congo Red. Measurements were made at <400 or x 1000 with an ocular micrometer. Illustrations of microscopic characters were made using an Olympus drawing tube system. Measurements are the normal range observed with outliers, if any, in brackets. Spore dimensions are given as length range < width range (n = 30). The length: width ratio (Q) of individual spores is presented

as the range and mean of Q values. Measurements do not include the apiculus. Basidia and cystidia dimensions are recorded as length range < width range (n = 20). All illustrations are based on collections PSC 2651, PSC 3004, PSC 3273 and PSC 3458. All collections have been accessioned into the State Herbarium of South Australia (AD).

DNA Extraction, amplification and processing

DNA was extracted from 5—10 mg of dried specimens by freezing with liquid nitrogen and grinding in a pestle and mortar with 500 ul of pH 8.0 isolation buffer (50 mM Tris-HCl, 170 mM EDTA, 1% N-lauroylsarcosine). The frozen paste was allowed to thaw, transferred to a 1.5 ml Eppendorf tube and incubated at 65°C for 5 min. Following addition of 300 ul 7.5 M ammonium acetate, the tubes were mixed by inversion, incubated on ice for 10 min. and then centrifuged at 13,000 g for 5 min. The supernatant (700 ul) was transferred to a fresh tube, mixed with 500 ul of isopropanol and held on ice for 10 min. Following centrifugation at 13,000 g for 3 min., the supernatant was discarded and the tubes drained by inversion on paper towel. The pellet was dissolved in 250 ul Tris EDTA buffer (10 mM Tris lmM EDTA

J. Adelaide Bot. Gard. 25 (2011) =

Fig 2. Boletus edulis, Adelaide Hills, PSC 3273, reticulum on stipe, under Castanea sativa.

pH 8.0), if necessary by incubation at 50°C for 5 min. and brief vortex mixing. PCR amplifications (20 ul) employed Phusion polymerase (New England Biolabs) in HF buffer, ITS1 and ITS4 primers (White et al. 1990) and | ul of a 1/20 to 1/100 dilution of the DNA extract. Amplification employed 5 min. at 98°C followed by 40 cycles (98°C 30 s, 57°C 15 s, 72°C 15 s) then 5 min. at 72°C prior to storage at 4°C. PCR products were purified using a PCK-1 kit (AdBiotec) and sequenced (AGRF) on both strands using the PCR primers. DNA sequence was assembled from unidirectional reads and sequences compared using Sequencher 4.9 (Gene Codes Corp., Ann Arbor, Michigan, USA) and blastn searches for similar sequences in GenBank conducted using NCBI software.

Results

Morphology

Specimens examined: PSC 2651, 19.v.2007; PSC 3004, 31.v.2009; PSC 3273, 15.1v.2010; PSC 3458, 27.xi1.2010.

Description. Pileus (70—) 100—250 (—275) mm diameter, (22—) 40-50 mm high, more or less hemispherical when young, later becoming plano-convex, irregularly plane to slightly concave; brown, light brown, 6D4—7, 7D8, 7E6—8, 7ES5—6 (Kornerup & Wanscher), fulvous 12, rust 13, rusty-tawny 14, brick 15 (Edinburgh Colour Chart); initially covered with white-grey bloom; dry becoming lubricous in centre; smooth to almost phlebioid in patches; margin rather irregular, projecting beyond tubes, wavy, finely revolute or involute, smooth, even, margin edge whitish with whitish bloom. Flesh of cap thick, solid, becoming rather spongy; white, cream, 4A2; not discolouring though turning slightly brown or reddish immediately under cap cuticle but not above stipe. Zubes emarginate to adnexed, with sulcus around stipe; separating easily from flesh; deep, 8-25 mm in centre; parallel; white, cream, becoming pale yellow 3A3-4, dull yellow, greenish-yellow 2B4, 2C4, 3B3-—5,

Boletus edulis in Australia (Boletaceae)

: erie. SS. la .

a 7 — To :

Fig 3. Boletus edulis, Adelaide Hills, PSC 2651, showing extended reticulum on stipe, under Quercus robur.

darkening to greyish-yellow 3C3 on cutting, not blueing. Pores initially white, cream, with whitish plug of tangled hyphae, later losing plug and becoming concolourous with tubes; not changing colour on bruising; small, 1—3/mm; rounded to angular, some irregular, slightly elongated; dissepiments thin to medium. Stipe (80—) 135-195 mm long, 30-68 (—98) mm diameter under cap, (32—) 70-98 mm in centre, 17-56 mm diameter 1 cm above base; ventricose or with bulbous base, occasionally cylindric; whitish, dull white-grey, pale brown, paler than cap, 5C4, 6C3—-4, 7D4, creamy-brown under cap, creamy clay-pink, closest to 30, 5B4, paler, whitish-brown at base; surface covered with whitish reticulum (Fig. 2) to half way down stipe, rarely over most of stipe (Fig 3), reticulations raised, white, white- beige, slightly greyer than 2B; reticulations small at top, approx. 1 <x 0.5 mm, larger below to 4.5 x 2 mm and more when most of the stipe is reticulated, lacunae shallow; surface rather waxy. Flesh of stipe thick, solid, white, not changing colour on bruising. Spore print olive brown, yellowish-brown, hazel 27, SE5—7. Spores (Figs. 4A, 5A) fusiform-ellipsoid, cylindrical, more or less smooth, rather thick-walled, pale yellow-brown in KOH; with oil globules; (12—) 13.6—18.4 x (4.0-) 4.5—-5.6 um, mean 16.8 x 5.3 um; Q: 2.4-3.8, mean Q: 3.17. Basidia (Figs. 4B, 4C, 5B) clavate, (24—) 34— 53 x (7—) 8-10 um, mean 35.8 x 9.6 um; with (2—) 4 sterigmata; without clamp connections. Cheilocystidia (Figs. 4D, 5C) fusiform, lageniform, ventricose-digitate, 40-72 <x 5.5-10 um, hyaline in KOH. Pleurocystidia similar to cheilocystidia. Pileipellis a trichoderm of irregular and regular interwoven hyphae, hyphae 3—10 um, many hyphae ascending, erect, exserted; slightly gelatinised; terminal elements 4-10 um; septa without clamp connections. Hymenial elements in reticulum of stipe (Figs. 4E, 5D): caulocystidia abundant; fusiform, cylindric, subclavate, 42—75 (—85) x (6.5—) 8-13 um, mean 62.2 x 8.9 um (n=10); basidia clavate, 2—4 spored but mostly 3-spored, 36—44 x 11—14 um. Hymenophoral trama of parallel hyphae.

P.S. Catcheside & D.E.A. Catcheside

J. Adelaide Bot. Gard. 25 (2011)

: . ' | : ' «* 7 — 2 { _ : |

- - i e ww . >» * 4 e. r

- = é “. i

Fig 4. Boletus edulis micrographs of PSC 2657. A spores; B, C basidia; D cystidium; E caulocystidia. Scale bar: 10 um.

Habitat. PSC 2651 in soil and deep leaf litter under Quercus robur L. with Amanita muscaria (L.) Lam., Lepista nuda (Bull.) Cooke and Russula sp., Araucaria bidwillii Hook. was nearby; PSC 3004 in litter under Quercus robur L.; PSC 3273 under 25 year old Castanea sativa Mill.; PSC 3458 in soil and pine needle litter under Pinus radiata D.Don. with Suillus granulatus (L.) Roussel, Rhizopogon rubescens (Tul.) Tul., Lactarius deliciosus (L.) Gray. Populus alba L. and Eucalyptus microcarpa (Maiden) Maiden were also nearby.

Molecular data

PSC 2651, 3273 and 3458 each had the same ITS sequence (Genbank accession JQ277466). Blast searches of the NCBI database at 1 November 2011 identified 39 entries having a sequence identical to JQ277466: DQ990838 (Peintner et al. 2007), EU417846, EU417847, EU417849, EU417851, EU417852, EU417855, EU 417856, EU417857, EU417858, EU417859, EU417861, EU417862, EU417863, EU417864, EU417868, EU417869, EU417874 (Beugelsdiyk et al. 2008), GU373493 (von Crautlein et al. 2011), GU198977, GU198981, GU198982, GU198991 (Korhonen et al. 2009), JF728991, JF728992, JF728994, JF728995, JEF728999, JEF729002, AY680981 (De la Varga et al. 2011), AY680983, AY680984, AY680985, AY680991, AY680992, AY680993, AY680994 (Leonardi et al. 2005), DQ002921 and DQ131622 (Agueda et al. 2006).

Discussion

The porcini mushrooms, Boletus sect. Boletus, comprise a number of described species (Singer 1986; Wang et al. 1995; Beugelsdijk et al. 2008; Dentinger et al. 2010) whose relationships have been clarified considerably by molecular data (Leonardi et al. 2005; Beugelsdik et al. 2008; Dentinger et al. 2010). B. edulis is now recognised as a taxon having variable morphology, a wide range of mycorrhizal partners and an extensive geographical range in the Northern Hemisphere. The molecular data suggest B. edulis includes the previously recognised species: B. betulicola, B. chippewaensis, B. persoonii, B. quercicola and B. venturi.

Morphologically, the Adelaide collections conform well to descriptions of Boletus edulis sensu stricto

|

(Watling 1970; Singer 1986; Breitenbach & Kranzlin 1991; Cortecuisse 1999; Beugelsdiyk et al. 2008) with respect to their microscopic and macroscopic characters. They differ from descriptions of the three other lineages recognised by Beugelsdijk et al. (2008) in Boletus sect. Boletus: B. aereus, B. reticulatus and B. pinophilus. None of the Adelaide collections have the dry, velvety darker brown cap and darker brown stem with a rust- brown reticulum of B. aereus; all had whitish or, at most, a pale brown stipe and the white to off-white reticulum usually reached no further than the stipe’s centre, unlike the reticula of both B. aereus and B. reticulatus which cover the whole stem. Collection PSC 3458 was found under Pinus radiata but no specimens had the thickly gelatinous cap pellicle or wide terminal elements of B. pinophilus; those in all our collections had terminal elements to 10 um while those of B. pinophilus may reach 27 um.

ITS sequences identical to those of the South Australian collections PSC 2651, PSC 3273 and PSC 3458 include AY680991, AY680992, AY680993, AY 680994, AY 680984, EU 417846, EU17847, EU17849, EU417851, EU417852, EU417855, EU417856, EU 417857, EU417858, EU417859, EU417861, EU417862, EU417863, EU417864 and EU417874 which are in the clade recognised by Beugelsdyyk et al. (2008) as B. edulis and also DQ002921 which is within the clade recognised by Dentinger et al. (2010) as B. edulis sensu stricto. This sequence is dissimilar to the ITS sequence of any of the additional taxa within the Boletus sect. Boletus clade recognised by Dentinger et al. (2010). There is thus no doubt that B. edulis has arrived in Australia.

Matching ITS sequences to the Adelaide Hills B. edulis biotypes have been found in collections made in Austria, Belgium, Finland, Italy, Netherlands, Spain and Sweden, variously associated with Betula, Castanea, Cistus, Fagus, Picea, Pinus and Quercus. In consequence, the currently available molecular data provide insufficient information to identify the source of the introduction into South Australia.

All currently known populations of B. edulis in Australia are within 8 km of one another and within 7 km of Mount Lofty at altitudes ranging from 330 m to 470 m above sea level. The extent of this range and

J. Adelaide Bot. Gard. 25 (2011)

association with three species of mature trees suggests that the introduction is not recent.

B. edulis is an obligate ectomycorrhizal fungus. Fruiting bodies have not been produced in culture, although efforts to cultivate the fungus in association with known plant hosts have had some success (Hall et al. 1998). These authors suggest that plantation forests of Pinus would be ideal habitats for B. edulis. In later work, Hall et al. (2005) observed Amanita excelsa ectomycorrhizas (ECMs) together with rhizomorphs of B. edulis penetrating the mantle around roots of their host plant and suggest that such co-symbionts may be mutually beneficial for all partners.

The discovery of Boletus edulis 11 South Australia raises the possibility of harvesting and managing this potentially valuable resource. A problem that might arise from deliberate cultivation of an introduced ectomycorrhizal fungus with a wide host range, such as B. edulis, 1s the possibility of its switching host to native trees. Although Malajczuk et al. (1982) found that ECMs of B. edulis did not form on any of the eleven species of eucalypt they tested for ECM formation, the possibility of B. edulis forming such symbioses with other Australian native species should not be discounted. Dunk et al. (2011) report on the invasion of Amanita muscaria into native Nothofagus forests in Tasmania and express the concern that such invasion may have detrimental effects on the ecosystem. Dunk et al. (2011) and Orlovich & Cairney (2004) comment on the lack of knowledge about the effect of the spread of exotic ECMs and emphasise the need for research into the long term consequences of the introduction or encouragement of potentially invasive mycorrhizal species.

Although recognition of the presence of the choice edible Boletus edulis in South Australia provides opportunities for exploitation of a new resource, there are corresponding risks which should be assessed before implementation of any actions to expand its distribution.

Acknowledgements

We thank Teresa Lebel and an anonymous reviewer for their helpful comments on an earlier version of this manuscript, members of the Adelaide Fungal Studies Group for their help finding the first specimens in 2007, Roy Halling for his comments on images of those specimens, and those who alerted us to probable additional sites in 2010.

References

Agueda, B., Parladé, J., de Miguel, A.M. & Martinez-Pena, F. (2006). Characterization and identification of field ectomycorrhizae of Boletus edulis and Cistus ladanifer. Mycologia 98(1): 23-30.

Beugelsdijk, D.C.M., van der Linde, S., Zuccarello, G.C., den Bakker, H.C., Draisma, S.G.A. & Noordeloos, M.E. (2008). A phylogenetic study of Boletus section Boletus in Europe. Persoonia 20: 1-7.

Breitenbach, J. & Kranzlin, F. (1991). Fungi of Switzerland Vol. 3 Boletes and agarics. (Verlag Mykologia: Lucerne).

Boletus edulis in Australia (Boletaceae)

Caglarlrmak, N., Unal, K. & Otles, S. (2002). Nutritional value of edible wild mushrooms collected from the Black Sea region of Turkey. Micologia Aplicada International 14: 1-5.

Cortecuisse, R. (1999). Collins Guide to the Mushrooms of Britain and Europe. (HarperCollins: London).

De la Varga, H., Agueda, B., Martinez-Pefia, F., Parlade, J. & Pera, J. (2011). Quantification of extraradical soil mycelium and ectomycorrhizas of Boletus edulis in a Scots pine forest with variable sporocarp productivity. Mycorrhiza. DOI 10.1007/s00572-011-0382-2.

Dentinger, B.T.M., Ammirati, J.F., Both, E.E., Desjardin, D.E., Halling, R.E., Henkel, T.W., Moreau, P-A., Nagasawa, E.., Soytong, K., Taylor, A.F., Watling, R., Moncalvo, J-M. & McLaughlin, D.J. (2010). Molecular phylogenetics of porcini mushrooms (Boletus section Boletus). Molecular Phylogenetics and Evolution 57: 1276-1292.

Dunk, C.W., Lebel, T. & Keane, P.J. (2011). Characterisation of ectomycorrhizal formation by the exotic fungus Amanita muscaria with Nothofagus cunninghamii 1n Victoria, Australia. Mycorrhiza. DOI 10.1007/s00572-011-0388-9.

Hall, I.R., Lyon, A.J.E., Wang, Y. & Sinclair, L. (1998). Ectomycorrhizal fungi with edible fruiting bodies-2. Boletus edulis. Econ. Bot. 52(1), 44—56.

Hall, I., Stephenson, S.L., Buchanan, P.K., Wang, Y. & Cole, A.L.J. (2003). Edible and poisonous mushrooms of the World. (Crop & Food Research Ltd: Christchurch).

Hall, I.R., Zambonelli, A. & Wang, Y. (2005). The cultivation of mycorrhizal mushrooms — success and failure. Fifth International Conference on mushroom biology and mushroom products. http://wsmbmp.org/ proceedings/S5th%20international%20conference/pdf/ chapter%204.pdf [Accessed 17 Nov. 2011].

Hawley, G.L. (2008). Ectomycorrhizal characterisation, species diversity and community dynamics in Pinus patula Schelcht. et Cham. plantations. http://eprints.ru.ac. Za/1165/1/Hawley-PhD.pdf [Accessed 11 April 2010].

Korhonen, M., Limatainen, K. & Niskanen, T. (2009). A new boletoid fungus, Boletus pinetorum, in the Boletus section Boletus from Fennoscandia (Basidiomycota, Boletales). Karstenia 49: 41-60.

Kornerup, A. & Wanscher, J.H. (1978). The Methuen book of colour. (Eyre Methuen: London).

Leonardi, M., Paolocci, F., Rubini, A., Simonini, G. & Pacioni, G. (2005). Assessment of inter- and intra-specific variability in the main species of Boletus edulis complex by ITS analysis. FEMS Microbiology Letters 243: 411-416.

Malajczuk, N., Molina, R. & ‘Trappe, J.M. (1982). Ectoycorrhizal formation in Eucalyptus I. Pure culture synthesis, host specificity and mycorrhizal compatibility with Pinus radiata. New Phytologist 91:467-482.

Marais, L.J., & Kotzé, J.M. (1977). Notes on ectotrophic mycorrhizae of Pinus patula in South Africa. South African Forestry Journal 100: 61-71

Orlovich, D.A. & Cairney, J.W.G. (2004). Ectomycorrhizal fungi in New Zealand: current perspective and future directions. New Zealand Journal of Botany 42: 721-738.

Peintner, U., Lotti, M., Klotz, P, Bonuso, E. & Zambonellt, A. (2007). Soil fungal communities in a Castanea sativa (chestnut) forest producing large quantities of Boletus edulis sensu lato (porcini): where is the mycelium of porcini? Environmental Microbiology 9(4): 880-889.

Ribeiro, B., Andrade, P.B., Silva, B.M., Baptista, P., Seabra, R.M. & Valentao, P. (2008). Comparative study on free amino acid composition of wild edible mushroom species. Journal of Agricultural and Food Chemistry 56: 10973—- 10979,

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a

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D

Fig 5. Boletus edulis, drawings of PSC 2657. A spores; B basidia; C cystidia; D caulocystidia. Scale bar: 10 um.

Royal Botanic Garden Edinburgh (1969). Flora of British fungi. Colour identification chart. (HMSO: Edinburgh). Stringer, A., Wang, Y., Bulman, S., Hall, I.R. & Orlovich, D. (2001). Boletus edulis sensu lato down under. In: Hall, ILR., Wang, Y., Zambonelli, A., Danell, E. (Eds.), Edible Mycorrhizal Mushrooms and Their Cultivation. (New Zealand Institute for Crop and Food Research Limited: Christchurch). [Proceedings of the Second International

Conference on Edible Mycorrhizal Mushrooms}.

Singer, R. (1986). The agaricales in modern taxonomy. Fourth edition. (Koelz. K6nigstein: Germany).

Sitta, N. & Floriani, M. (2008). Nationalization and globalization trends in the wild mushroom commerce of Italy with emphasis on porcini (Boletus edulis and allied species). Economic Botany 62: 307-322.

von Crautlein, M., Korpelainen, H., Pietilainen, M. & Rikkinen, J. (2011). DNA barcoding: a tool for improved taxon identification and detection of species diversity. Biodiversity & Conservation 20: 373-389.

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Wang, Y., Sinclair, L., Hall, IR. & Cole, A.L.J. (1995). Boletus edulis sensu lato: a new record for New Zealand. New Zealand Journal of Crop and Horticultural Science 23(2): 227-231.

Watling, R., (1970). Boletaceae: Gomphidiaceae: Paxillaceae. In: Henderson, D.M., Orton, P.D., Watling, R. (eds), British fungus flora, Agarics and Boleti, Vol. 1. (Royal Botanic Garden: Edinburgh).

Watling, R. & Li T-H., (1999). Australian Boletes. A preliminary survey. (Royal Botanic Garden: Edinburgh).

White, T.J., Bruns, T., Lee, S. & Taylor, J. (1990). Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In: Innis, M., Gelfand, D., Sninsky, J. & White, T. (eds.), PCR Protocols: a Guide to Methods and Applications, chapter 38. (Academic Press: Orlando, Florida).

Wilkinson, S. (2011). Wild about porcini - at $120 a kilo. The Advertiser April 02, 2011. http://www.adelaidenow.com. au/ipad/wild-about-porcini-at- | 20-a-kilo/story-fn6br97]- 1226032325058 [Accessed 2 Nov. 2011].

A | OQOURNAL of the ADELAIDE

BOTANIC GARDENS

AN OPEN ACCESS JOURNAL FOR AUSTRALIAN SYSTEMATIC BOTANY

flora.sa.gov.au/jabg

Published by the STATE HERBARIUM OF SOUTH AUSTRALIA on behalf of the BOARD OF THE BOTANIC GARDENS AND STATE HERBARIUM

State Herbarium of South Australia

PO Box 2732

Kent Town SA 5071

Australia

Board of the

Botanic Gardens and

State Herbarium

a)

Government — of South Australia

Department of Environment, Water and Natural Resources

J. Adelaide Bot. Gard. 25 (2012) 11-15

The genus Alisma L. (Alismataceae) in South Australia

John G. Conran

Australian Centre for Evolutionary Biology and Biodiversity, Benham Building DX 650 312, School of Earth & Environmental Sciences, The University of Adelaide, SA 5005 E-mail: john.conran@adelaide.edu.au

State Herbarium of South Australia, DENR Science Resource Centre, P.O. Box 2732, Kent Town, South Australia 5071

Abstract

The genus Alisma L. (Alismataceae) is revised for South Australia, with several populations previously incorrectly identified as the introduced weed A. lanceolatum With. (narrow leaved water plantain) found instead to be the native A. plantago-aquatica L. (common water plantain). The distinguishing features for these taxa and related Alismataceae in South Australia are given and the implications for their conservation and/or weed status are discussed.

Keywords: South Australia, Alisma plantago-aquatica, Alisma lanceolatum, Alismataceae,

taxonomy.

Introduction

The monocot family Alismataceae consists of herbaceous aquatic macrophytes and is represented by 11-14 genera and ~100 species worldwide (Haynes et al. 1998, Jacobson & Hedrén 2007, Mabberley 2008), of which five genera and 11 species occur in Australia (Jacobs & McColl 2011). The genus Alisma L. (water plantains) has 9-11 species (Bjérkquist 1968, Wang et al. 2010), mostly widely distributed throughout the Northern Hemisphere of the Old World (Wang et al. 2010), but with three native species recognized for North America (Rubtzoff 1964, Haynes & Hellquist 2000). In particular, A. plantago-aquatica L. and A. lanceolatum With. are widespread throughout Eurasia with the former extending to Australia and the latter to North Africa, and both species are also introduced weeds in a number of places including North America (Haynes & Hellquist 2000) and New Zealand (Allan Herbarium 2000). Alisma plantago-aquatica has a natural range extending from Europe across to subtropical and temperate areas of SE Asia including China, Burma, Japan, Korea, Thailand and Vietnam and south-eastern Australia and its wide distribution (and presence in Australia) may be the result of long-distance transport of the fruits by migratory wetland birds (Green et al. 2002), but this requires further study.

In Australia, A. plantago-aquatica is considered to be native to at least New South Wales and Victoria (Aston 1973), as well as possibly introduced to south- east Queensland and Tasmania (Jacobs & McColl 2011), but its natural distribution requires further investigation.

1]

Alisma has been the subject of some taxonomic controversy in South Australia and it is unclear if it is native or an early introduction. However, based on the native distribution of A. plantago-aquatica in Victoria and its presence in the upper Murray and Wimmera River drainage systems, it should be expected to occur naturally in South Australia. Alisma plantago-aquatica was recorded in South Australia by Black (1922, as A. plantago non L.); however, there are early 20" century collections at AD made by him of both Alisma plantago-aquatica and A. lanceolatum. In contrast, Eichler (1965), Aston (1973), Jessop (1986) and Jacobs & McColl (2011) considered that the South Australian Alisma accessions all represented the introduced weed A. lanceolatum, also apparently failing to notice the presence of collections of both species.

As a result of field work by the author in wetlands across the Southern Lofty and South East regions, it quickly became apparent that there were two Alisma species in South Australia: the introduced A. lanceolatum, but also the native A. plantago-aquatica. Examination of all the A/isma collections at AD further supported this, as well as providing historical data on some now extinct localities for A. plantago-aquatica. Because the recent Flora of Australia account only lists one species in the State (Jacobs & McColl 2011), a revised account for South Australia is necessary both to update distributions, as well as to detail known localities of the native species for conservation. This is especially relevant as several native populations are the target of eradication programs due to misidentification as the introduced weed.

J.G. Conran

Taxonomy

Key to the species of Alismataceae in South Australia 1. Carpels weakly coherent into a star-like ring

Damasonium minus (R.Br.) Buchenau 1: Carpels distinct, forming heads or rings on areceptacle 2

2. Flowers unisexual; carpels in a convex head ......... * Sagittaria platyphylla (Engelm.) J.G.Sm. 2: Flowers all bisexual; carpels arranged in a flattened ring

3. Leaf blade narrowly elliptic to lanceolate, apex acute to acuminate, base cuneate, tapering; petals usually acuminate 2. *Alisma lanceolatum With.

3: Leaf blade ovate to broadly lanceolate, apex bluntly acute to rounded, base + rounded to cordate; petals usually broadly rounded.................004.

4. Flowers pink; style longer than the ovary

|. Alisma plantago-aquatica L. 4: Flowers white; style shorter than or equalling the ovary Alisma triviale' Pursh

Alisma L. Sp. Pl. 1: 342 (1753).

Perennial, evergreen or geophytic, glabrous, rhizomatous herbs. Leaves emergent to submerged and floating, sessile or petiolate; lamina linear to ovate, margins entire, base attenuate to rounded or cordate, apex obtuse to tapering-acute. Inflorescence an erect, generally emergent, pyramidal compound panicle of 2-10 verticillate whorls; bracts acuminate. Flowers bisexual; pedicels spreading; pedicel bracts lanceolate, short, acuminate; sepals erect to spreading, sometimes recurving in fruit; petals pink or white, entire to apically subdentate; stamens 6, in pairs opposite the petals, filaments filiform, glabrous; carpels 12—20 around margin of flattened receptacle; ovule 1; style lateral to terminal. Fruits laterally compressed, abaxially 2-3- ribbed achenes. x = 7.

|. Alisma plantago-aquatica L.

Sp. Pl. 1: 342 (1753).

A. lanceolatum auct. non With.: Jessop in Jessop & Toelken, FI. S. Austral. 4: 1709 (1986), partly.

‘A. plantago’ orth. var.: L., Syst. Nat. ed. 10, 2: 993 (1759), cf. Richt., Codex Bot. Linn. 351 (1835-1839); R.Br. in Flinders, Voy. Terra Austral. 2: 592 (1814); Woolls, Contrib. Fl. Austral. 150 (1867); Benth, Fl. Austral. 7: 185 (1878); F.Muell., Syst. Cens. Austral. Pl. 121 (1882) & Syst. Cens. Austral. Pl. ed. 2, 205 (1889); J.M.Black, Natural. Fl. S. Austral 143 (1909); Fl. S. Austral. 1: 49 (1922) & Fl. S. Austral. ed. 2, 1: 52 (1943).

Perennial aquatic herbs to 1.5 m, rhizome thick, corm-like, to 5 cm diam. Leaves emergent, petioles 5—40 cm long, 4-16 mm diam.; blade lanceolate to broadly

' Although not recorded as naturalised in Australia, A. triviale is cultivated at the Mount Lofty Botanic Gardens (accession no. G843495; listed in Sandham & Kellermann 2010 as A. plantago-aquatica var. americanum Schult. & Schult.f.), where it self-seeds freely in an artificially maintained bog and along an associated creek and dam (along with Sagittaria platyphylla). It is keyed here because of its very close resemblance to A. plantago-aquatica when not in flower and its possible weed potential.

2

J. Adelaide Bot. Gard. 25 (2012)

elliptic or ovate, 2.5—30 x 1-12 cm, apex bluntly acute to rounded, base obtuse-tapering to cordate; venation acrodromous, mostly with 2 basal pairs and a single, strong suprabasal pair of secondary veins. Inflorescence to 150 x 50 cm. Flowers numerous in ~10 mm diam. clusters of 2—5, pedicels 3-45 mm; sepals ovate- lanceolate, green, 1.5—3.5 <x 1.5—2 mm; petals broadly obovate, purplish-white to pink, 3.5-6.5 x 3.5—6 mm, margins entire to denticulate, apex obtuse to broadly rounded; anthers ellipsoid, 0.7—1.4 mm; style + straight, 0.5—1.5 mm, ovary 0.4—0.6 mm long. Fruiting heads 4—7 mm diam.; achenes obovoid, |.7—3.1 mm long, abaxial keels broadly rounded, with usually 1 median groove, beak + erect. 2n = 14. Common water plantain. (Fig. 1A-C).

Distribution. S.A.: NL, SL, SE; Qld; N.S.W.; Vic; Tas; Europe and Asia. Naturalised in North and South America and New Zealand.

Habitat. Grows in damp ground or shallow water. Plants in permanently damp areas are evergreen, those in drier areas dying back to the thick, corm-like rhizome.

Conservation status. Rare in South Australia and apparently locally extinct at several localities. Threatened by habitat loss and eradication due to misidentification as A. lanceolatum.

Flowering and fruiting period. Dec.—Mar.

Affinities. Species taxonomy in Alisma has _ been problematic (Hendricks 1957, Voss 1958, Rubtzoff 1964), with the native North American former representatives of the very widespread and variable Old World taxon A. plantago-aquatica now regarded as separate species on stylar and other morphological characteristics (Bjérkqvist 1967, Haynes & Hellquist 2000), although the Eurasian species is also naturalised there (Haynes & Hellquist 2000). This separation is further supported cytologically, as A. plantago-aquatica is a diploid (2n = 14), whereas the North American segregate species A. triviale is a tetrapoid (2n = 28), as 18 A. lanceolatum (2n = 26, 28) (Baldwin & Speese 1955, Jacobson & Hedrén 2007).

Molecular analyses of evolution in the genus are still unclear, with evidence of relatively low genetic divergence between many taxa. The A. plantago- aquatica species complex was distinct and thought to be possibly more derived phylogenetically than A. lanceolatum (Jacobson & Hedrén 2007), but the origins of most of the polyploid taxa are uncertain. However, the phylogenetic study of Alisma by Jacobson & Hedrén (2007) did not include any native Australian material of A. plantago-aquatica, so its precise relationship to the European and Asian members of the complex are still unknown.

Note. The name of this species was published by Linnaeus as “Alisma Plantago AX”, which translates to “Alisma plantago-aquatica”’ (Art. 23.3, McNeill et al.

J. Adelaide Bot. Gard. 25 (2012)

2006). However, from the 10" edition of his Systema Naturae (1759), Linnaeus only used “plantago” as epithet of the species, a fact already noted by Richter (1835-1839). This orthographic variant of the name has been used by numerous authors throughout the years; the major publications relevant to Australia are listed in the synoymy.

Specimens examined.

SOUTH AUSTRALIA. Northern Lofty: | km from Wirrabarra Forest Headquarters, 25 Apr. 1994, R./ Bates 37061 (AD 99522058). Southern Lofty: Reedbeds at Fulham (c. 8 km W of Adelaide), 18 Oct. 1908, JM Black s.n. (AD 97623463); Reservoir near Mount Lofty Railway Station, 13 Apr. 1960, E.A [sing s.n. (AD 96220135, AD 96220136); Cresswell Park, Gilberton, 14 Jan. 1976, A.G.Spooner 4525 (AD 97621406); Thomas Gully, Mt Bold Reservoir Track 43, 14 Nov. 2007 D.J.Duval 946 & T:S.Te (AD 214871, AD 214872); Woorabinda Res., Stirling Linear Park, 8 Oct 2011, J.G.Conran 3142 (AD, ADU). South East: Millicent, town drain running toward Lake Bonney, 27 Nov. 1996, D.Guerin s.n. (AD 212157, AD 99650378).

2. *Alisma lanceolatum With.

Arr. Brit. Pl. ed. 3, 2: 362 (1796).

A. plantago-aquatica auct. non L.: R.Br. in Flinders, Voy. Terra Austral. 2: 592 (1814); Woolls, Contrib. Fl. Austral. 150 (1867); Benth, Fl. Austral. 7: 185 (1878); F.Muell., Syst. Cens. Austral. Pl. 121 (1882) & Syst. Cens. Austral. Pl. ed. 2, 205 (1889); J.M.Black, Natural. Fl. S. Austral 143 (1909); J.M.Black, Fl. S. Austral. 1: 49 (1922) & FI. S. Austral. ed. 2, 1: 52 (1943)., pro parte, as ‘plantago’ (orth. var.).

Perennial aquatic herbs to 1.5 m, rhizome thick, corm-like, to 5 cm diam. Leaves emergent, petioles 5—50 cm long, 4-16 mm diam.; blade lanceolate, 3—25 x 1-6.5 cm, apex acute to acuminate, base cuneate, tapering; venation pinnate, mostly with | basal pair and two well-spaced suprabasal pairs of secondary veins. Inflorescence to 120 <x 50 cm. Flowers numerous in clusters of 2-5, ~9 mm diam., pedicels 3-45 mm; sepals ovate-lanceolate, green, 1.5—3.5 mm; petals broadly oblanceolate to obovate, purplish-white to pink, 3.5— 6.5 mm, margins entire to irregularly subdentate, apex acuminate to rounded acute; anthers ellipsoid, 0.6—1.1 mm; style + curved, 0.5—1.5 mm, ovary 0.4—0.6 mm long. Fruiting heads 4-8 mm diam.; achenes obovoid, 2—3 mm long, abaxial keels broadly rounded, with 1-2 median grooves, beak + erect. 2n = 28. Narrow leaved water plantain. (Fig. 1D-F).

Distribution. S.A.: SL; W.A.; Qld; N.S.W.; Vic; Tas.; native to Europe, North Africa and Asia.

Habitat. Grows in damp ground or shallow water. Plants in permanently damp areas are evergreen, those in drier areas dying back to the thick, corm-like rhizome.

Conservation status. A naturalised weed which appears to be spreading widely throughout the Onkaparinga River catchment

Flowering and fruiting period. Dec.—Mart.

13

Alisma in South Australia (Alismataceae)

Affinities. A phylogenetically somewhat isolated tetra- ploid species of uncertain origin (Jacobson & Hedrén 2007).

Specimens examined.

SOUTH AUSTRALIA. Southern Lofty: Leslie Creek, Mylor, 7 Mar. 1986, R.J. Bates 6754, (AD 98631080); Mylor, banks of Onkaparinga River, 4 Feb. 1979, L.D. Williams 10196 (AD 98426341); Rliver] Onkaparinga, n[ear] Longnook, 31 Dec. 1906, S.A.White s.n. (AD 97919317); Bridgewater, 20 Jan. 1919, J.M.Black s.n. (AD 97623464); Onkaparinga River, 31 Dec. 1906, H.H.D.Griffith s.n. (AD 97623462); Mylor, “Rockford” (property of Mr C.G. Little), at the Onkaparinga River, 14 Dec. 1967, Hj.Eichler 19715 (AD 97616010); Clarendon Weir, 3 Dec. 1928, /.B.Cleland s.n. (AD 97236070); Onkaparinga River at Mylor, 2 Dec. 1938, J.B. Cleland s.n. (AD 97236071); Onkaparinga River, near Mylor, c. 20 km SE of Adelaide, 9 Dec. 1944, J.B.Cleland s.n. (AD 967510057, AD 95852022, AD 95852023); Onkaparinga River near the bridge of the road from Mylor to Echunga, 19 Mar. 1957, Hj. Eichler 13699 (AD 95814005); Mylor, Aldgate Creek, near the bridge on the road to Aldgate, 19 Mar. 1957, Hj.Eichler 13690 (AD 95813059, AD 95813049, AD 95813050, AD 95813051, AD 95813052, AD 95813043); Onkaparinga River at Verdun, 15 Feb. 1967, B.Grivell s.n. (AD 96932472, AD 96932463, AD 96932459, AD 96932456); Aldgate Valley reserve in creekline, 12 Mar. 2010, C.J. Brodie 1528 (AD 235326).; SE corner of Happy Valley Reservoir c. 15 km S of Adelaide, 11 Jan. 1959 R.Schodde 1069 (AD 96027072, AD 96027073, AD 96027074, AD 96027075, AD 96027076, AD 96027077); Creek crossing just S of Verdun on Onkaparinga Valley Rd c. 100 m N of turnoff to Mount Barker Rd, 2 Jan. 2012, J.G.Conran 3240 (AD, ADU); Aldgate Valley Reserve, 7 Jan. 2012, J.G.Conran 324] (AD, ADU); Leslie Ck, Mylor Parklands, cnr Stock Rd and Strathalbyn Rd, 7 Jan. 2012, J.G.Conran 3243 (AD, ADU); Cooper Reserve, Silver Lake Rd, Mylor, 7 Jan. 2012, J.G.Conran 3244 (AD, ADU); Outfall creek/drain from Happy Valley Reservoir near cnr Chandlers Hill and Serpentine Rd, 18 Jan. 2012, ..G.Conran 3245 (AD, ADU).

Discussion

All the Alismataceae listed here mainly occur in seasonally moist to permanently wet bogs and wetlands, Where they grow in shallow water along creeks, lake or swamp margins, or in ditches. Although the Alisma species are all perennial, they can also behave as annuals and/or geophytes in climates with seasonal drought (Jacobs & McColl 2011).

Germination in A/isma often requires a combination of stratification and alternate drying/wetting cycles (Byérkqvist 1967); A. plantago-aquatica, however, has different and more specific germination requirements than Alisma lanceolatum (Moravcova et al. 2001). Nevertheless, these studies were conducted for European populations where there are strong seasonal effects and the germination biology of the species in southern Australia has not been studied in detail.

Alisma lanceolatum, along with the noxious weed Sagittaria platyphylla and the native Damasonium minus are all considered to be weeds of rice fields in New South Wales (McIntyre & Newnham 1988), with S. platyphylla also regarded as a weed in Western Australia (Sage et al. 2000). Alisma and Damasonium species

J.G. Conran

J. Adelaide Bot. Gard. 25 (2012)

Fig. 1. Alisma species in South Australia. A-C A. plantago-aquatica: A habit; B leaf showing sub-cordate leaf with two pairs of basal and one suprabasal secondary leaf vein pair and percurrent tertiary venation; C flower showing rounded petals and more or less straight stigmas. D-F A. lanceolatum: D habit; E leat showing lanceolate leaf with one pair of basal and two pinnate suprabasal secondary leaf vein pairs and percurrent tertiary venation; F flower showing + acuminate petals and curved stigmas. Scale bars: A, D 20 cm; B, E5 cm; C, F5 mm.

have been found to show herbicide resistance and are the subject of studies for biocontrol (Ash et al. 2008), but this also has implications for the conservation of the native species in southern Australia.

The collections of Alisma plantago-aquatica in the State Herbarium of South Australia (AD) indicate that from the few collections made, it was probably never very common. Populations are no longer present in the Torrens River catchment and the vast majority of suitable habitat locations for the species in the Onkaparinga River catchment only appear to support A.

14

lanceolatum. The single report for A. plantago-aquatica from the Northern Lofty in the Wirrabarra Forest needs investigation to see if the species is still extant there.

The South-East Region collections from Miullicent were reported from the town drain running toward Lake Bonney, where it was noted on the specimen (AD 212157) that the population had grown rapidly and become locally dominant by 1996, despite an attempt to control with metsulfuron-methyl in 1994.

There are potential issues with the superficial similarity of sterile or juvenile A. plantago-aquatica

J. Adelaide Bot. Gard. 25 (2012)

to the native Plantago major L. (Plantaginaceae) and the introduced Zantedeschia aethiopica (L.) Spreng. (Araceae), both of which grow in the same environments. Nevertheless, they are easily separated on leaf venation, as the tertiary veins in A. plantago-aquatica are strongly percurrent (ladder-like) between the secondaries (Fig. 1B), whereas they are reticulate in these others.

It is unfortunate that the failure in the past to realise that there were two Alisma species — one native and one weedy has meant that an already uncommon native plant in South Australia has possibly been targeted for removal due to its misidentification as a weed. Given that there are now very few localities for this species in the State, mainly due to habitat alteration/loss, it is probably just as well that it is resistant to the ALS inhibitor class herbicides that are commonly used to control aquatic weeds (Ash et al. 2008, Figueroa et al. 2008). Measures should be taken to encourage its conservation at those places where it still occurs, rather than physical or chemical removal.

More research, particularly cytological and genetic is needed to determine whether the specimens of A. plantago-aquatica are native to South Australia or an early introduction, as well as whether the species in Australia is the same as that in the Northern Hemisphere of the Old World. However, for the moment the species should be considered native until proven otherwise and conserved accordingly.

Acknowledgements

The School of Earth & Environmental Sciences, The University of Adelaide is thanked for provision of resources for this research. The Chief Botanist, State Herbarium of South Australia (AD), 1s thanked for access to specimens.

References

Allan Herbarium (2000). Nga Tipu o Aotearoa — New Zealand Plant Names Database. (Landcare Research, New Zealand: Lincoln). http://nzflora.landcareresearch.co.nz/ [Accessed: 11 Dec. 2011].

Ash, G., Chung, Y., McKenzie, C. & Cother, E. (2008). A phylogenetic and pathogenic comparison of potential biocontrol agents for weeds in the family Alismataceae from Australia and Korea. Australasian Plant Pathology 37: 402—405.

Aston, H.I. (1973). Aquatic plants of Australia: a guide to the identification of the aquatic ferns and flowering plants of Australia, both native and naturalized. (Melbourne University Press: Carlton).

Baldwin, J.T., Jr. & Speese, B.M. (1955). Chromosomes of taxa of the Alismataceae in the range of Gray’s Manual. American Journal of Botany 42: 406-411.

Byjorkqvist, I. (1967). Studies in Alisma L. I. Distribution, variation and germination. Opera Botanica 17: 1-128. Bjérkquist, I. (1968). Studies in Alisma L. II. Chromosome studies, crossing experiments and taxonomy. Opera

Botanica 19: 1-138.

Black, J.M. (1922). Flora of South Australia, part 1.

(Goverment Printer: Adelaide).

15

Alisma in South Australia (Alismataceae)

Eichler, H. (1965). Supplement to J.M. Blacks Flora of South Australia (second edition, 1943-1957). (Government Printer: Adelaide).

Figueroa, R., Gebauer, M., Fischer, A. & Kogan, M. (2008). Resistance to bensulfuron-methyl in water plantain (Alisma_ plantago-aquatica) populations from Chilean paddy fields. Weed Technology 22: 602—608.

Green, A.J., Figuerola, J. & Sanchez, M.I. (2002). Implications of waterbird ecology for the dispersal of aquatic organisms. Acta Oecologica 23: 177-189.

Haynes, R.R., Les, D.H. & Holm-Nielsen, L.B. (1998). Alismataceae. In: Kubitzki, K. (ed.), The families and genera of vascular plants 4: 11-18. (Springer: Berlin).

Haynes, R.R. & Hellquist, B. (2000). 190. Alismataceae. In: Flora of North America Editorial Committee (eds), Flora of North America North of Mexico 22: 7—25. (Oxford University Press: New York and Oxford).

Hendricks, A.J. (1957). A revision of the genus Alisma (Dill.) L. American Midland Naturalist 58: 470-493.

Jacobs, S.W.L. & McColl, K.A. (2011). Alismataceae. In: Wilson, A.J.G. (ed.), Flora of Australia Volume 39: 3-126. (Australian Biological Resources Study/CSIRO Publishing: Collingwood, Victoria).

Jacobson, A. & Hedrén, M. (2007). Phylogenetic relationships in Alisma (Alismataceae) based on RAPDs, and sequence data from ITS and ¢trnL. Plant Systematics and Evolution 265: 27-44.

Jessop, J.P. (1986). Alismataceae. In: Jessop, J.P. & Toelken, H.R. (eds), Flora of South Australia 4: 1709-1710. (Government Printer: Adelaide).

Mabberley, D.J. (2008). Mabberleys plant-book. A portable dictionary of plants, their classifications, and uses, ed. 3. (Cambridge University Press: Cambridge).

Mclntyre, S. & Newnham, M.R. (1988). Distribution and spread of the Alismataceae in the rice-growing region of New South Wales. Cunninghamia 2: 25-38.

McNeill, J., Barrie, F.R., Burdet, H.M., Demoulin, V., Hawksworth, D.L., Marhold, K., Nicolson, D.H., Prado, J., Silva, P.C., Skog, J.E., Wiersema, J.H., Turland, N.J. (2006). International code of botanical nomenclature (Vienna Code). A.R.G. Gantner: Rugell. [Regnum Vegeta- bile 146]

Moravecova, L., Zakravsky, P. & Hroudova, Z. (2001). Germination and seedling establishment in Alisma gramineum, A. plantago-aquatica and A. lanceolatum

under different environmental conditions. Folia Geobotanica 36: 131—146. Richter, H.E.F. (1835-1839). Caroli Linnaei Systema,

Genera, Species Plantarum uno volumine, editio critica, adstricta, conferta sive Codex Botanicus Linnaeus [...] in usum botanicorum practicum editit brevique adnotatione explicavit. Otto Wigand: Leipzig.

Rubtzoff, P. (1964). Notes on the genus Alisma. Leaflets of Western Botany 10: 90-95.

Sage, L., Lloyd, S.G. & Pigott, J.P. (2000). Sagittaria platyphylla (Alismataceae), a new aquatic weed threat in Western Australia. Nuytsia 13: 403-405.

Sandham, J. & Kellermann, J. (eds) (2010). Botanic Gardens of Adelaide catalogue of plants 2010: Adelaide, Mount Lofty and Wittunga Botanic Gardens. (Board of the Botanic Gardens and State Herbarium: Adelaide). | Journal of the Adelaide Botanic Gardens, Supplement 2|

Voss, E.G. (1958). Confusion in Alisma. Taxon 7: 130-133.

Wang, Q.-F., Haynes, R.R. & Hellquist, B. (2010). Alismataceae. In: Wu, Z.-Y. & Raven, P.H. (eds), Flora of China 23: 84-89. (Science Press & Missouri Botanical Garden Press: Beijing & St. Louis).

A | OQOURNAL of the ADELAIDE

BOTANIC GARDENS

AN OPEN ACCESS JOURNAL FOR AUSTRALIAN SYSTEMATIC BOTANY

flora.sa.gov.au/jabg

Published by the STATE HERBARIUM OF SOUTH AUSTRALIA on behalf of the BOARD OF THE BOTANIC GARDENS AND STATE HERBARIUM

State Herbarium of South Australia

PO Box 2732

Kent Town SA 5071

Australia

Board of the

Botanic Gardens and

State Herbarium

a)

Government — of South Australia

Department of Environment, Water and Natural Resources

J. Adelaide Bot. Gard. 25 (2012) 17-25

Malva weinmanniana (Besser ex Rchb.) Conran, a new name for the pink-flowered form of M. preissiana Schitdl. (Malvaceae) =

J.G. Conran* “, R.M. Barker’ “ & E. Rippey’

“School of Earth and Environmental Sciences, Benham Building DX 650 312, The University of Adelaide, South Australia 5005 Email: john.conran@adelaide.edu.au

’State Herbarium of South Australia, DENR Science Resource Centre, P.O. Box 2732, Kent Town, South Australia 5071 Email: robyn.barker@sa.gov.au

“School of Plant Biology, The University of Western Australia, 35 Stirling Highway, Crawley, Western Australia 6009

“ Australian Centre for Evolutionary Biology and Biodiversity, The University of Adelaide, South Australia 5005

Abstract

The two morphotypes of the Australian endemic Malva preissiana Migq. (Austral Hollyhock) are here regarded as separate, distinct species with the white-flowered M. preissiana a coprophilic halophyte on seabird-dominated near-shore islands, or rarely mainland shores or inland salt lakes used by seabirds. Malva weinmanniana (Besser ex Rchb.) Conran, based on Lavatera weinmanniana Besser ex Rchb., is the new name applied to the more common, pink-flowered, largely mainland taxon. The names L. weinmanniana, L. plebeia var. eremaea J.M.Black., L. plebeia var. tomentosa Hook.f., and M. behriana Schltdl. are lectotypified and their statuses discussed. A key to the taxa and a table of comparisons 1s supplied.

Keywords: Malva, Lavatera, Malvaceae, hybrid, nomenclature, taxonomy, southern Australia, South Australia, Western Australia, Victoria.

Introduction

The two ecotypes of the Australian endemic Malva preissiana Mig. (Austral Hollyhock) discussed in Barker & Conran (2007) are here regarded as separate, distinct species, based on morphological and ecological differences (Moore 1994, Lewis 1999). The white- flowered M. preissiana 1s primarily found on near-shore islands of the southern coast of Australia (Rippey ef al. 2002, Zed et al. 2006), or more rarely from mainland shores or inland salt lakes associated with seabirds. Malva preissiana has for some time been recognised as being threatened by competition from, and hybridisation with, the introduced M. arborea (L.) Webb & Berthel. across its range and is in need of conservation (Rippey et al. 2002, Rippey 2004). The pink-flowered taxon, M. weinmanniana (Besser ex Rchb.) Conran, does not appear to be threatened but there is the necessity to be able to distinguish between all of the taxa concerned for conservation and monitoring purposes.

The differences between these taxa are summarised in Table 1 and a comparison of the flowers 1s shown in Fig. 1.

La:

Key to taxa in the Malva preissiana complex and their hybrid with M. arborea 1. Leaves well-spaced along stems, upper surface much less tomentose than lower ......... Malva weinmanniana 1: Leaves crowded together on stems and densely tomentose on both surfaces 2. Flowers deep cerise pink with a dark purple to black SOUS a on telat pate gn ate en Sebel Wena cain ews Malva arborea 2: Flowers white to pale pink with a pale centre 3. Epicalyx lobes shorter than the calyx lobes ......... Aa Ey LCE AER Hod 0 ce late amend ee bb HE Malva pretssiana 3: Epicalyx lobes and calyx lobes of similar length .... Matlva arborea X< M. preissiana

The Malva preissiana complex

Images of most type specimens cited are availabe on the following web-sites: ¢ herbarium.univie.ac.at/database/search.php (HAL) ¢ apps.kew.org/herbcat/navigator.do (K) ¢ www.lu.se/biological-museums/botanical-museum/databases (LD) ¢ www.plants.jstor.org (LD, K, TCD). Specimens accessible through these databases are indicated with an asterisk.

J.G. Conran, R.M. Barker & E. Rippey J. Adelaide Bot. Gard. 25 (2012)

Fig. 1. Flowers of the two native Malva species, M. arborea and the natural hybrid. A M. preissiana (Conran 857, ADU); B M. weinmanniana (Conran 2038, ADU); C M. arborea x M. preissiana (Rippey 169, PERTH); D M. arborea (Conran 1527, ADU). Scale bars 10 mm. See front cover of journal for colour reproductions of these photographs.

1. Malva preissiana Miq. [Lavatera plebeia auct. non Sims: Benth., Fl. Austral. 1: 185 Mig. in Lehm., Pl. Preiss. 1(2): 238 (1845). — Type (1863), p.p; F.Muell., Pl. Indigenous Colony Victoria 1; citation: “L. Preiss 1893, 14 Nov. 1839, Crescit in 166 (1860), p.p.; J.LM.Black, Fl. S. Austral. 3: 373 (1926), insula Penguin-Island” [S of Cape Peron, Rockingham, D-p; J.M.Black, FI. S. Austral. ed. 2, 3: 554 (1952), p.p.; WA, fide Marchant (1990)]. Syntypes: LD 1065806!*: W.R.Barker in Jessop & Toelken, FI. S. Austral. 2: 833 TCD 0010674!*, labelled “No. 654”; MEL 1528422! (ex (1986), p.p.]

Herb. Steetz); MEL 2282397! [Lavatera australis auct. non Schrad. ex Colla: J.M. Black Lavatera plebeia Sims var. tomentosa Hook.f., J. Bot. Fil. S. Austral. 3: 373 (1926).| (Hooker): 412 (1840). — Type citation: “Near Wool- [Lavatera plebeia Sims var. eremaea auct. non J.M.Black: north. Mr Gunn (n. 655), (also in New Holland, J.M.Black, Fl. S. Austral. 3: 373 (1926), p.p. (only as Cunningham).” Lectotype (here designated): R. Gunn to 7.G.B. Osborn s.n., Jan 1922, Franklin Islands, AD 655/18 [3]7, 24 Nov. 1836, Van Diemens Land, near 96303018!). | Woolnorth (K000659311!*, p.p., ex Herb. Hook.; Fig. [Malva behriana auct. non Schlitdl.: W.R.Barker et al., 2). Possible syntype or isosyntype: 4. Cunningham 87, Census S. Austral. Vasc. Pl. ed. 5 (= J. Adelaide Bot. [Jan 7] 1818, King George Sound (K000659315!*), p.p. Gard. Suppl. 1), 93 (2005), p.p. | L. australis Schrad. ex Colla, Hortus Ripul. 1: 134 (1824), [?Lavatera hispida auct. non. Desf.: Benth., Fl. Austral. 1: nom. nud. 186 (1863) (see below)]

18

J. Adelaide Bot. Gard. 25 (2012)

Table 1. Comparison of Malva preissiana, M. weinmanniana, M. arborea and the hybrid Malva arborea x M. preissiana.

Character

Habit

Stems

Leaf arrangement

Leaf indumentum

Malva preissiana

Usually sympodial and multi-stemmed at the base

Densely tomentose

Clustered apically,

internodes short

More or less equally densely tomentose on both surfaces

Malva weinmanniana

+ Monopodial, branches mostly arising along a main central stem

Densely tomentose

Widely spaced along

stems, internodes long

Less tomentose above; moderately to densely tomentose below

Malva weinmanniana (Malvaceae)

Malva arborea x Malva preissiana

+ Monopodial

Densely tomentose

Clustered apically,

internodes short

‘Tomentose on both surfaces

Malva arborea

+ Monopodial

Sparsely hairy to sub-

glabrous

Clustered apically,

internodes short

‘Tomentose on both surfaces

Leaf upper surface Dark grey-green Greyish yellow-green Dark grey-green Dark green colour Pedicels Usually shorter than the Usually longer than the Equalling calyx Exceeding calyx petals petals Epicalyx (bracteoles) Not exceeding calyx Not exceeding calyx Equalling calyx Exceeding calyx Calyx in fruit Enclosing mericarps Enclosing Enclosing Not enclosing Petal shape Obovate Obovate Broadly reniform Truncate-obovate Petal apex Deeply notched Deeply notched Shallowly notched ‘Truncate Petal colour White; unmarked to _— Pale pink; pink-striped Pink; deeper pink- Cerise; maroon-striped faintly mauve-striped striped Petal base and claw Pale greenish Pale yellowish Pink Deep blackish-maroon colour Mericarps 11-12 11-12 10-11 6-7 Habitat Coastal and offshore Generally inland Coastal and offshore Coastal and offshore islands islands islands Soil/nutrient preference Coprophilic Disturbed soil, not Coprophilic Coprophilic, nutrient-

coprophilic

Typification of Malva preissiana Maiq.

Preiss’s collection of Malva preissiana in the Lund herbarium is available for viewing on the web as is another duplicate in the herbarium of Trinity College Dublin. However the authors have yet to see the Preiss collection from Miquel’s own herbarium in Utrecht and so lectotypification of the name has not been undertaken. Other species named by Miquel in Plantae Preissiana have almost invariably been lectotypified by a specimen from Utrecht; e.g. see citations for a number of Casuarina L. and Leptomeria R.Br. species in the Australian Plant Name Index (APNI 2011-2012).

Placement of Bentham’s Lavatera hispida Desf.

Bentham (1863) included Lavatera hispida Dest. (now generally considered to be L. olbia L. within Lavatera s.str., see Molero Briones & Montserrat Marti 2007) in his account of Malvaceae in Flora Australiensis, describing it as:

a hirsute species with nearly sessile flowers forming a long terminal raceme or interrupted spike, and with broad hirsute involucres, a native of the Mediterranean region,

appears to be naturalized in some islands of Bass’s Straits (FE) Mueller).

19

rich, or disturbed soils

From Bentham’s description, the specimens might belong to M. preissiana, or possibly represent naturalised material of Malva arborea (L.) Webb & Berthel, both of which are characteristically tomentose. However, at this stage no Mueller specimens from the Bass Strait Islands have been located in MEL and a search of Bentham’s herbarium at Kew is required. Until specimens are located, the identity of this material and the application of the name remain unknown.

Typification of Lavatera plebeia var. tomentosa Hook.f.

In describing var. tomentosa, Hooker (1840) states:

This seems to be the species alluded to under L. plebeja, in the Bot. Mag. as having been collected on the south coast of New Holland. The var. 6 [= tomentosa] differs from the described state of L. plebeja, in having the upper side of the leaf equally tomentose as the underside. A sheet in Hooker’s herbarium in K (K000659311 ex Herb. Hook.), reproduced here in Fig. 2, has a typical Gunn label with the number 655/18[3]7. It has been annotated as “L. plebeia Sims B” and also bears the collecting locality Woolnorth for the 24 November 1836. The label is clearly in agreement with the protologue. An additional locality on the label, Trefoil

J.G. Conran, R.M. Barker & E. Rippey J. Adelaide Bot. Gard. 25 (2012)

ROYAL BOTANIC GARDENS KEW

OWN

K000659310

WV aeons pte bea SIS var. boner, bora WOOK. dét. / rev. Gerald Krebs ¥% 6 6.1986

Herbarium Universitatis Lipsiensis

|

paniasal JyGHUAdOS

+ \= . *. + ff - - = sy. &2 * - al {)| Ltt a y i aw “ 7 - ee ne at re on

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Neen lex, plebea SIMS. Var. tome to7a 06K. tooked CA te typus of the

taxon. A! nen Mrs. LES 19 FS

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~~ ri _ Ad] snerirpenre cal 3 shy DBP cover BE eee ied MS Gt ' = “Ynag a atl bee Be ee Ee : ROYAL BOTANIC GARDENS KEW * ay : - > wi Zio, subpart | : " eae oe z, - 3 ig Oeil | 4 KOOO6S9311 é ; Cie ea : : iat os

Fig. 2. Lectotype (bottom left and top right branches) of Lavatera plebeia var. tomentosa (K 000659311). Available at apps.kew.org/herbcat/ getimage.do?imageBarcode=K000659310 and used with the permission of the Trustees of the Royal Botanic Gardens, Kew.

20

J. Adelaide Bot. Gard. 25 (2012)

Island, may or may not relate to the same date but it does suggest that there are two collections present. At the time of collection Gunn was the Police Magistrate at Circular Head in NW Tasmania and he made numerous trips to Woolnorth by the Van Diemen’s Land schooner (Buchanan 1990); presumably one of these trips may have been extended further north to Trefoil Island off the NW coast of Tasmania.

Three of the specimens on the sheet have been identified with this label by a pencil line — the two on the bottom left and the one on the right hand side. At an earlier stage the label was presumably attached to one or all of these specimens. All of the specimens comply with Hooker’s statement about the hairiness of the leaf, and there is a note by G. Krebs (Herbarium, University of Leipzig), dated 28" May 1985, that they represent type material.

The Cunningham specimen mounted on the same sheet as the lectotype (K 000659310!*, Fig. 2) is from Peel’s Range (now Cocoparra Range), New South Wales and is labelled as Lavatera australis. It is VM. weinmanniana (see below). Even though it is a Cunningham collection in Hooker’s own herbarium, it bears a different name and is not in agreement with the protologue with respect to the hair covering on the leaves and so is presumed not to be the Cunningham specimen alluded to by Hooker in the protologue.

A sheet from Bentham’s herbarium also has a Cunningham specimen (4. Cunningham _ 87; K 000659313*), collected in 1818 and mounted with a Horticultural Society of London specimen dated 6.9.[18]29 (K 000659315*). The Horticultural Society specimen is M. weinmanniana, while the Cunningham collection is M. preissiana. The latter possibly represents the Cunningham material alluded to in the protologue and therefore may be syntype or isosyntype material of Hooker’s var. tomentosa.

Note

A specimen (Andrews 254; K 000659318*) in Ben- tham’s herbarium, collected in the vicinity of Lake Eyre in Nov. 1875, also appears to be M. preissiana. Although far outside the normal coastal and offshore island range of this taxon, Lake Eyre is a major seabird rookery at times of inundation and the saline, guano-enriched rookery soils there could easily provide a suitable habitat for seeds carried by coastal seabirds migrating to these new, ephemeral breeding grounds.

2. Malva weinmanniana (Besser ex Rchb.) Conran, comb. nov.

Lavatera weinmanniana Besser ex Rchb., Iconogr. Bot. Exot. 1: t. 60 (1824), plate with analysis (Art. 42.3); p. 45 (1825), description; pp. xiv, 68 (1827), notes & addendum. — Type citation (from the 1825 description): “in Nova Hollandia, teste cl. Besser I.c. floret in hortis sub dio ad a Julio ad auctumnum”. Lectotype (here designated): Reichenbach’s plate t. 60! (Fig. 3). Lavatera plebeia Sims, Bot. Mag. 48: pl. 2269 (1821), nom. illeg., non M. plebeja Stev., Bull. Soc. Imp. Naturalistes

Bal

Malva weinmanniana (Malvaceae)

Moscou xxix. (1856) I. 325. Benth., Fl. Austral. 1: 185 (1863), p.p.; F.Muell., Pl. Indigenous Colony Victoria |: 166 (1860), p.p.; J.LM.Black, Fl. S. Austral. 3: 373 (1926), p.p.; J.M.Black, FI. S. Austral. ed. 2, 3: 554 (1952), p.p.; W.R.Barker, Fl. S. Austral. 2: 833 (1986), p.p. — Althaea plebeia (Sims) Schult. ex Steud., Nomencl. Bot. ed. 2, 1: 210 (1840). — M. australiana M.F.Ray, Novon 8: 291 (1998). — Type: Sim’s plate 2269! (see discussion in Ray 1998, p. 292. The implication is that this might be considered to be the holotype, but this designation 1s not used).

Lavatera australis A.Cunn. ex Hook.f., J. Bot. (Hooker) 2: 412 (1840), nom. nud. (based on a Cunningham collection from Peel’s Range, now Cocoparra Range, New South Wales, and labelled as Lavatera australis; see K 000659310* and lectotypification of L. plebeia var. tomentosa, above).

Malva_ behriana Schlitdl., Linnaea 20: 633 (1847). — Lavatera behriana (Schltdl.) Schltdl., Linnaea 24: 699 (1852). — Type citation: “In Thale des Murray-Flusses. Juli.” [Behr]. Lectotype (here designated): H.H. Behr s.n., July [1845], In Thale des Murray-Flusses (HAL 0098406!*). Possible syntype or isosyntype: F) Mueller s.n., Australia (IK 000659320*).

Lavatera plebeia Sims var. eremaea J.M.Black, Fl. S. Austral. 3: 373 (1926), p.p. — Type citation: “Ardrossan, Y.P.; Caroona, E.P. Ooldea; Franklin Islands; along the Great Bight’. Lectotype (here designated): 7: Richards s.n., Nov. 1879, Euria (AD 96303017! p.p.). Syntypes: O. Tepper s.n., 1879, Ardrossan (AD 96303017! p.p.; MEL 1528294 n.v.); R. Tate s.n., 8 Feb. 1879, Gully Bunda Cliffs E from Wilson Bluff (AD 96303017! p.p.) — all from J.M. Black’s herbarium at AD and annotated as var. eremaea. Possible syntype: Anon. s.n., in Herb. J.M. Black, 24 Sep. 1920, Ooldea (AD 97612120B!), not annotated as var. eremaea by Black. [Excluded syntype: 1.G.B. Osborn s.n., Jan 1922, Franklin Islands (AD 96303018!) — see M. preissiana.|

The identity and validity of Lavatera weinmanniana Besser ex Rchb.

As with previous workers we had thought that this name was illegitimate until the proof stage of this paper, when it was discovered that the protologue of L. weinmanniana was merely the illustration (Reichenbach 1824) and not the associated description (printed a year later, in 1825), which included earlier names in synonymy. While the description no longer forms part of the protologue it is needed to check against earlier publications using this same name and also to provide an identification of the taxon concerned and ensure that it is associated with the correct Australian species.

In describing Lavatera weinmanniana, Reichenbach attributed the name to von Besser and referred to von Besser’s (1823a) Cat. Hort. Cremen., presumably a list of plants growing in the gardens administered by von Besser at Kremenets in western Ukraine. We have not been able to find a copy of this publication, but since it seems to be a list of names and since other names listed in this same publication, such as Melilotus pallidus Besser and Rumex reticulatus Besser, have been legitimised in later publications, we have treated von Besser’s name as a nomen nudum. It may be that the publication referred to was von Besser’s (1823b) seed list of the same year

J.G. Conran, R.M. Barker & E. Rippey

for the same gardens and here the name was certainly a nomen nudum.

Reichenbach’s name might still have been a later homonym because earlier in the same year de Candolle published Lavatera weinmanniana Trev. ex DC. However further investigation revealed that de Candolle (1824) had merely listed L. weinmanniana as a queried synonym of L. sylvestris Brot. (= Malva linnaei M.F.Ray); the author “Trev.” who provided the original name is assumed to be L.C. Treviranus, Professor of Botany at the University of Bonn; no publication by him using this name has been traced.

Based on the protologue illustration and the later published description, especially the reference to pink flowers and widely-spaced sub-glabrous leaves, the species 1s clearly associated with the pink-flowered mainland taxon, and the name has been adopted here.

Publication and typification of L. welnmanniana Besser ex Rchb.

The publication of L. weinmanniana by Reichenbach was a protracted affair and our interpretation and acceptance of this name is reliant on the publication dates given in 7Z-2 (Stafleu & Cowan 1983) being correct.

Reichenbach’s illustration of Lavatera weinmanniana (Fig. 3) was published as plate 60 in 1824, the year before the publication of the detailed description of the species on p. 45. It has always been assumed that the publications were at the same time and so the citation of earlier names in the synonymy accompanying the description and notes on L. weinmanniana led to it being deemed an illegitimate name. However if it is just the illustration that forms the protologue, and this meets all of the criteria for valid publication, then the name has to be considered valid. Since the illustration includes a flowering branch with leaves as well as dissections of the various parts of the plant, viz. external and internal views of the calyx, the young fruit, the columella with a single carpel attached and a dorsal view of the carpel, it meets the requirements of the botanical code (Art. 42.3 & 42.4; McNeill et al. 2006), for an illustration with analysis, even though the caption was published with the later description, and the description is needed in order to determine some of the finer detail.

Subsequent mentions of L. weinmanniana in the same work include p. xiv of a section titled ‘Erstes hundert’ (The first one hundred, viz. of plates), which is in German and includes some horticultural notes as well as the comment that the plant 1s also called L. australis and L. plebeja, and that the name L. sylvestris was applied erroneously; this was published in 1827. In the same year a further section entitled ‘Addenda et emendanda ad Centuram primam’ included the statement on p. 68 that L. weinmanniana was the same as L. plebeia, but not L. silvestris:

L. weinmanniana convenit omnino L. plebeia Sims Bot.

Mag. 2269, cui pariter adscribiter patria Nova Hollandiae, est minime L. silvestris

22

J. Adelaide Bot. Gard. 25 (2012)

Since the illustration is the basis for the name, it has here been designated as the lectotype. According to Stafleu & Cowan (1976-1988), the plates were hand-coloured copper engravings, this particular one by Reichenbach himself, although others were by Humm. There is no colour associated with the copy of the illustration reproduced here from the volume held by the library of the National Herbarium of Victoria (Fig. 3) and nor is there any colour associated with the copy of this publication available on the web.

Whether there are any extant specimens in what remains of Reichenbach’s herbarium in W has not been established, but the name was clearly used in horticultural circles for some time before its publication and so there may well be some specimens with this name in European herbaria.

Etymology

Johannes Anton Weinmann (1782-1858), whose name presumably provides the epithet for this species, was a German born Russian botanist and gardener who worked in Dorpat (Tartu) in Estonia from 1804—1813 and then for Empress Maria Feodorowna (1759-1828), “Mother of Czars” (Grant 1905), at her palace in Pavlovsk, St Petersburg, Russia. The association of his name with the species in horticultural circles predated the protologue as indicated above, but there is no indication of why his name was used and what connection he had with Australian plants. This was however the peak of interest in Europe in plants from Australia and there was considerable exchange between the various gardens.

W.S.J.G von Besser (1784-1842), who is credited with naming the species after Weinmann, was an Austrian-Polish botanist at Kremenets, western Ukraine (then Russia), while Ludwig Reichenbach (1793-1879) was a German botanist and artist, and director of the Dresden botanical garden (Stafleu & Cowan 1976-— 1988).

Lectotypification of Malva behriana Schltdl.

Although there are no original labels by Behr on the specimen in HAL it bears the same information as 1s given in the protologue and 1s clearly the specimen used by Schlechtendal, then editor of Linnaea and resident in Halle, in describing M. behriana. It is therefore the obvious choice for lectotype of this name. A specimen at K in Herb. Bentham (K 000659320*) received from Mueller and previously in Sonder’s herbarium has also been annotated with this name and may represent syntype or isosyntype material, although it was annotated on the 16" May 1985 by G. Krebs: “In my opinion not once [sic] of the 3 examplars is a Holo-, Iso- or Syn typus.” While the leaves appear similar to those of the lectotype and flowers are present on both collections, the material in K does seem to be at a somewhat later stage since, unlike the lectotype, there appear to be young fruits present.

J, Adelaide Bot. Gard. 25 (2012) Malva weinmanniana (Malvaceae)

Fig. 3. Reichenbach’s (1824) illustration and lectotype of Lavatera weinmanniana. Reproduced with permission from the State Botanical Collection, Royal Botanic Gardens Melbourne.

J.G. Conran, R.M. Barker & E. Rippey

Placement of Lavatera plebeia Sims

Ray (1998) designated Sim’s (1821) plate as the type, and the depiction of widely spaced leaves and pink flowers with a yellowish centre supports its placement here. Furthermore, the accompanying description implies that the upper leaf surface is less hairy than the lower and also indicates that the original specimen was from an inland expedition beyond the recently crossed Blue Mountains. This evidence was also no doubt utilised by previous workers in establishing a typical variety in relation to var. eremaea J.M.Black and var. tomentosa Hook.f. (e.g. Black 1926, Barker 1986), or by its exclusion from the taxon representing the tomentose form (Hooker 1840).

Of interest is the further comment that this species was different from Robert Brown’s collection of a “nearly related, but distinct species” from the south coast, presumably a reference to Brown’s collections from Seal Island in King Georges Sound, Western Australia, and from Mt Brown in South Australia. Both of these specimens are entered in the Brown database (Chapman et al. 2001) as L. plebeia and need to be re- examined so that they can be correctly named.

Lectotypification of Lavatera plebeia var. eremaea J.M. Black.

The Richards collection from Euria in the Penong area of South Australia was chosen as lectotype since it 1s most representative of the taxon and has been annotated with the varietal name by J.M. Black. It consists of the two specimens on the left hand side of the sheet, both of them with flowers and fruits. It is mounted on the same sheet as two of the syntypes: Tate’s collection from Gully Bunda Cliffs on the Bight near the Western Australian border, consisting of two small branches with flowers and fruits, and Tepper’s collection from Ardrossan on Yorke Peninsula, which consists of four small branches. Both of these syntypes have also been annotated by J.M. Black as var. eremaea. The Tepper collection is somewhat atypical for MZ weinmanniana being particularly hirsute and with very small leaves. Nevertheless, it still falls within the range of variation seen in this species, resembling the occasional salt- stressed plants seen from from Bass Strait and offshore islands in Western Australia.

The other possible syntype in the Black herbarium, collected from Ooldea, has not been annotated as var. eremaea by Black and consists merely of a long piece of stem with a single leaf.

Osborn’s collection from Franklin Islands near Cowell is an excluded syntype since it represents typical M. preissiana; it has been annotated as var. eremaea by Black. A specimen from Caroona on Eyre Peninsula has not been found in AD at this time.

3. Malva arborea X M. preissiana

Occasional sterile hybrids between these two species have been recorded in Western Australia (Rippey 2004). It has been known for some time that the introduced M.

J. Adelaide Bot. Gard. 25 (2012)

arborea 1s replacing the native M. preissiana on off- shore islands in South Australia, Victoria and Western Australia (Rippey 2004) and so there is a clear need to be able to distinguish between the hybrid and the two parents for those involved in monitoring such changes.

The hybrid can be distinguished easily from both M. preissiana and M. weinmanniana by having the epicalyx (bracteoles) and calyx lobes of similar length, whereas the epicalyx lobes of the native taxa are shorter than those of the calyx (Table 1). The hybrid also tends to have broader, less apically notched petals with only very faint striping and a darker pink claw, whereas both native species have prominently notched petals and pale ereenish-yellow claws (Fig. 1). While the hybrid and . weinmanniana both share pink flowers, the former is much more tomentose overall than M. weinmanniana, has an upper leaf surface which is less hairy than the lower and usually bears c. 12 mericarps per fruit.

Acknowledgements

Val Stajsic (MEL) is thanked for comments on Malva, for confirmation of the presence or absence of particular specimens in MEL, and for arranging a copy of the Reichenbach plate of L. weinmanniana from the holdings of the National Herbarium of Victoria. Graham Bell (AD) 1s thanked for help in confirming J.M. Black’s handwriting in the lectotypification of L. plebeia var. eremaea and Bill Barker (AD) is thanked for helpful comments on the manuscript. We are indebted to the Editor, Jurgen Kellermann, for drawing the publication dates of Reichenbach’s [conographia botanica exotica to our attention. The Keeper of the Herbarium, RBG Kew, is thanked for permission to use the image of the lectotype for L. plebeia var. tomentosa.

References

Australian Plant Name Index (APNI) (2011-2012). IBIS database. (Centre for Australian National Biodiversity Research, Australian Government: Canberra). www. cpbr.gov.au/cgi-bin/apni [accessed 13 Jan 2011 & 2 May 2012).

Barker, R.M. & Conran, J.G. (2007). Malva_ preissiana Mig., an overlooked name for Lavatera plebeia Sims (Malvaceae), with a note on variation within the species. Journal of the Adelaide Botanic Gardens 21: 771-72.

Barker, W.R. (1986). Lavatera. In: Jessop, J.P. & Toelken, H.R. (Eds). Flora of South Australia 2: 833-835. (Government Printer: Adelaide).

Bentham, G. (1863). Flora Australiensis: a description of the plants of the Australian territory, Vol. 1 (Ranunculaceae to Anacardiaceae). (Lovell Reeve and Co.: London, UK).

Besser, W.S.J.G. von (1823a). Catalogus Plantarum in Horto botanico gymnasii Volhyniensis Cremeneci cultarum. (Kremenets).

Besser, W.S.J.G. von (1823b). Enumeratio seminum, quae e messi anni MDCCCXXIII e horto botanico lycaei Volhyn. Cremeneci offeruntur in mutuam commutationem. (Kremenets). Available at: www.nationaalherbarium.nl/ seedlists/images/Krzemieniec/1823/All%20descriptions. htm [accessed 17 May 2012].

Black, J.M. (1926). Flora of South Australia, Part III (Melia- ceae—Scrophulariaceae). (Government Printer: Adelaide).

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J. Adelaide Bot. Gard. 25 (2012)

Buchanan, A. (1990). Ronald Campbell Gunn (1808-1881). In: Short, P.S. (ed), History of systematic botany in Australasia, pp. 179-192. (Australian Systematic Botany Society Inc.: Collingwood).

Candolle, A.P. de (1824). Prodromus systematis naturalis regni vegetabilis, sive, Enumeratio contracta ordinum generum specierumque plantarum huc usque cognitarium, Juxta methodi naturalis, normas digesta, Vol. 1. (Treuttel & Wiirtz: Paris).

Chapman, A.R., Moore, D.T., Rees, R.G. & Groves, E.W. (2001). Robert Brown’ Australian botanical specimens, 1801-1805 at the BM. Database. florabase.dec.wa.gov.au/ brown/search [accessed 2 May 2012].

Grant, C. (1905). A Mother of Czars. A sketch of the life of Marie Feodorowna, wife of Paul I and mother of Alexander I and Nicholas I. (John Murray: London).

Hooker, J.D. (1840). Contributions towards a Flora of Van Diemen’s Land, chiefly from the collections of Ronald Gunn, Esq. and the late Mr Lawrence. Hooker s Journal of Botany 2: 399-421.

Lewis, A. (1999). Spatial distribution of Malva dendromorpha and Malva australiana on West Island, South Australia. Unpublished B.Sc. (Hons) thesis, The University of Adelaide.

Marchant, N.G. (1990). The Western Australian collecting localities of J.A.L. Preiss. In: Short, P'S. (Ed). History of systematic botany in Australasia, pp. 131—135. (Australian Systematic Botany Society Inc.: Collingwood).

McNeill, J., Barrie, F.R., Burdet, H.M., Demoulin, V., Hawksworth, D.L., Marhold, K., Nicholson, D.H., Prado, J., Silva, P.C., Skog, J.E., Wiersema, J.H. & Turland, N.J. (2006). International Code of Botanical Nomenclature (Vienna Code). (A.R.G. Gantner: Ruggell, Lichtenstein). [Regnum Vegetabile 146].

Molero Briones, J. & Montserrat Marti, J.M. (2007). A new species of Lavatera Sect. Olbia (Medik.) DC. (Malvaceae) from north-east Morocco. Botanical Journal of the Linnean Society 153: 445-454.

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Malva weinmanniana (Malvaceae)

Moore, B. (1994). Variation in Lavatera plebeia Sims (Malvaceae) in South Australia, and the taxonomic status of var. tomentosa Hook. f. and var. eremaea Black. Unpublished B.Sc. (Hons) thesis, The University of Adelaide.

Ray, M.F. (1998). New combinations in Malva (Malvaceae: Malveae). Novon 8: 288-295.

Reichenbach, H.G.L. (1824-1827). Jconographia botanica exotica, sive Hortus botanicus, imagines plantarum imprimis extra Europam inventuram colligens, cum commentario succincto editus. Prima Centuria Tabularum. (Fredericum Hoffmeister: Leipzig). Available at: books. gsoogle.com.au/books?id=-E] AAAAcAAJ&dq= iconogra phia%20botanica%20exotica&pg=PP l#v=onepage&g&f =true0 [accessed 24 Apr. 2012].

Rippey, E., Rippey, J. & Dunlop, N. (2002). Management of indigenous and alien Malvaceae on islands near Perth, Western Australia. In: Veitch, C.R. & Clout, M.N. (Eds), Turning the Tide: The Eradication of Invasive Species [Proceedings of the International Conference on Eradication of Island Invasives/], pp. 254-259. (UCN SSC Invasive Species Group, IUCN: Gland, Switzerland & Cambridge, UK).

Rippey, E. (2004). Malva dendromorpha / Malva australiana hybrid. Western Australian Naturalist 24: 198-200.

Stafleu, F.A. & Cowan, R.S. (1976-1988). Taxonomic Literature: a selective guide to botanical publications and collections with dates, commentaries and types. 7 vols. (Bohn, Scheltama & Holkema: Utrecht).

Sims, J. (1821). Lavatera plebeia. Mallow-like Lavatera. Curtis s’ Botanical Magazine 48: Plate 2269. Available at: bhl.ala.org.au/item/14332#page/218/mode/lup [accessed 28 May 2012].

Zed, T., Conran, J.G. & Lewis, A. (2006). Vegetation patterns in relation to bird nesting preferences on West Island, South Australia. Zransactions of the Royal Society of South Australia 130: 211-226.

A | OQOURNAL of the ADELAIDE

BOTANIC GARDENS

AN OPEN ACCESS JOURNAL FOR AUSTRALIAN SYSTEMATIC BOTANY

flora.sa.gov.au/jabg

Published by the STATE HERBARIUM OF SOUTH AUSTRALIA on behalf of the BOARD OF THE BOTANIC GARDENS AND STATE HERBARIUM

State Herbarium of South Australia

PO Box 2732

Kent Town SA 5071

Australia

Board of the

Botanic Gardens and

State Herbarium

a)

Government — of South Australia

Department of Environment, Water and Natural Resources

J. Adelaide Bot. Gard. 25 (2012) 27-54

A Revision of the genus Stellaria (Caryophyllaceae) in Australia

Cathy H. Miller & Judy G. West

Australian National Herbarium, Centre for Australian National Biodiversity Research, G.P.O. Box 1600, Canberra, Australian Capital Territory 2601 E-mail: Cathy.Miller@csiro.au; Judy. West@environment.gov.au

Abstract

A review of the species of Stel/aria L. occurring in Australia is provided. The genus 1s represented by 10 species, of which S. papillata C.H.Mill. & J.G.West is described as new, as are 3 subspecies: S. angustifolia subsp. rotundisepala C.H.Mill. & J.G.West, S. multiflora subsp. collaris C.H.Mill. & J.G.West and S. multiflora subsp. nebulosa C.H.Mull. & J.G.West. Two new combinations are made, namely S. angustifolia subsp. tenella (Benth.) C.H.Mill. & J.G.West, based on S. glauca Hook. var. (?) tenella Benth. and S. /eptoclada (Benth.) C.H.Mill. & J.G.West, based on Stellaria glauca var. (?) leptoclada Benth. Lectotypes are selected for S. angustifolia Hook., S. caespitosa Hook.f., S. glauca Hook. var. (?) tenella Benth, S. filiformis (Benth.) Mattt., S. flaccida Hook., S. multiflora Hook., S. pungens Brongn. and S. squarrosa Hook. Notes, maps, illustrations and an identification key are provided for all taxa occurring 1n Australia.

Keywords: Caryophyllaceae, Ste//aria, Australia, revision, taxonomy, nomenclature.

Introduction

This paper is a precursor to the treatment of Stellaria L. (Caryophyllaceae) for the Flora of Australia series and provides names for four new taxa, as well as typification and nomenclatural information. Stellaria consists of about 120 species (Morton 2005) and is found throughout temperate areas worldwide and at higher altitudes in tropical areas. There are a few weedy species, such as S. media (L.) Vill. and S. graminea L., which are found almost worldwide.

Stellaria belongs to the family Caryophyllaceae Juss., subfamily Alsinoideae Beilschm. Members of this subfamily are distinguished from subfamily Caryophylloideae Arn. by their free sepals and from the subfamily Paronychoideae Meisn. by their exstipitate leaves (Conn 1983; Chater & Heywood 1993; McNeill 1962). The genus is characterised by the presence of five sepals and five petals which are usually bifid; however in some species the petals are markedly reduced or absent. The closest related genera are currently thought to be Cerastium L. and Holosteum L. (Fior et al. 2006; Harbaugh et al. 2010).

There are currently six endemic species and three introduced species recognised with one further new endemic species and three endemic subspecies being described here. Two additional new combinations have also been made.

In Australia some of the endemic species appear to be closely related and form complexes. The ‘angustifolia group’ usually occurs in wetter areas and 1s morphologically similar to northern hemisphere species, particularly S. palustris L. This complex is distinguished

2/

by the presence of sessile leaves, five fully-formed bifid petals, ten stamens, and by the absence of staminodes. It currently consists of two species, namely S. leptoclada (Benth.) C.H.Mill. & J.G.West, and S. angustifolia Hook., the latter with three subspecies. It can be difficult to separate these taxa without good flowering or fruiting material. Two additional species, S. flaccida Hook. and S. pungens Brongn., have some affinities with the ‘angustifolia group’ but are quite distinct, particularly in habit and vegetative characters. A second ‘multiflora eroup’ occurs in dryer or harsher regions such as coastal or inland dunes, on islands or in alpine areas. It is distinguished by the presence of leaves with narrowed and elongated bases so that they often appear petiolate (although they are technically still sessile) and by the reduction in size, form and number of floral structures. This ‘multiflora group’ consists of three species: S. filiformis (Benth.) Mattt., S. papillata C.H.Mill. & J.G.West and S. multiflora Hook., the last with three subspecies.

This occurrence of such complexes seems to be common in the genus in the northern hemisphere taxa as well. There have been a number of studies carried out on the North American S. /ongipes Goldie complex using both morphological and molecular analyses. The results from these studies (Chinnappa & Morton 1984; Chinnappa et al. 2005) showed that environmental factors influence key characters such as inflorescence type and the occurrence of scarious bracts. Chinnappa & Morton (1984) stated that “it proved possible to ‘change’ several of the ‘species’ into other ’species‘ simply by altering the environmental conditions under which they were growing” (Chinnappa & Morton 1984;

C.H. Miller & J.G. West

Table 1. Summary of subgeneric classification schemes for Stellaria. Fenzel (1840) b. “Eustellaria’

ao. Petiolares

Section Subsection Species listed

Subsection 6. Larbreae

Species listed S. graminea

Subsection Eg. Spinescentes

Species listed

Unplaced species

Chinnappa et al. 2005). Due to this plasticity Morton (2005) decided to adopt a broad concept of S. /ongipes and only recognized one infraspecific taxon. By contrast S. media and S. pallida are two members of the ‘media eroup’ that, together with S. neglecta Weihe, have been variously interpreted as varieties, subspecies or, as here, species following most recent authors such as Chen & Rabeler (2001) and Morton (2005).

Taxonomic History

Stellaria was first described by Linnaeus (1753) with several European taxa listed, including S. nemorum L., S. dichotoma L., S. radians L., S. graminea, S. cerastoides L., S. biflora L. and S. holostea L. This last taxon was chosen by Hitchcock & Green (1929) as the type species of the genus. To date there have been no major worldwide revisions; the closest is the synopsis published in Die Natiirlichen Pflanzenfamilien by Pax & Hoffmann (1934). Most publications have been regional floras such as for the British Isles (Clapham et al. 1952), Europe (Chater & Heywood 1993), China (Chen & Rabeler 2001) and North America (Morton 2005).

Species. The first Australian species of Stellaria, S. pungens was illustrated and named by Brongniart in 1834, for which no description was provided and currently no specimen has been located. W.J. Hooker (1834) published two species, S. angustifolia and S. squarrosa Hook., in the same year, two more species, S. flaccida and S. multiflora, in 1836 (Hooker 1836) and a further species, S. caespitosa Hook. in 1840 (Hooker 1840).

The first treatment of the genus for Australia was by Bentham (1863) in which he listed five species. This comprised the introduced species S. media, and three endemic species, S. multiflora, S. pungens and S. flaccida. The fifth species listed was the northern hemisphere S. glauca With. Bentham included S. angustifolia under S: glauca. Additionally two infraspecific taxa, S. glauca var. (?) leptoclada Benth. and S. glauca var (?) tenella Benth. were published by Bentham. He also described another species, Drymaria filiformis Benth. which was later transferred to Stellaria (Mattfeld 1938).

S. media, S. flaccida

S. squarrosa, 5. pungens

J. Adelaide Bot. Gard. 25 (2011) (=)

Pax & Hoffmann (1934) I. ‘Eustellaria 1. Petiolares

S. media, S. pallida 4. Larbreae

S. graminea, S. palustris 5. Spinescentes

S.pungens

S. multiflora

During the late 1800’s the use of the name S. glauca was replaced by the name S. palustris Ehrh. ex Retz (Mueller 1887) and this name was in common usage until the later part of the 1900’s. During the 1980’s until the current day the consensus has been that this taxon is truly Australian and the name S. angustifolia has been applied (Doust 1990, Miller & West 1996).

The introduced species S. graminea (Curtis 1956) and S. pallida (Dumort.) Crep. (Eichler 1965) were first reported as occurring in Australia in the second half of the twentieth century.

No new names for Australian taxa have been published since the first half of the twentieth century; however some informal names have been used in various state floras in the second half of the twentieth century. In this treatment S. media, S. multiflora, S. pungens, S. flaccida, S. filiformis (Benth.) Mattt., S. graminea and S. pallida are recognised. Stellaria multiflora 1s divided into three subspecies, two of which are new. S. angustifolia is resurrected and divided into three subspecies, one of which is new and one is a new combination based on Bentham’s S. glauca var (‘?) tenella. Bentham’s S. glauca var. (?) leptoclada is raised to species level. S. papillata is newly described for Australia.

Suprageneric classification. Fenzl (1840) published the first suprageneric classification for Caryophyllaceae, which was enlarged upon by Pax & Hoffmann (1934). Many of their tribal and subtribal names have not been used in recent literature (Rabeler & Bittrich 1993) as these classifications were based upon “arbitrary interpretations of morphology” (Bittrich 1993). It has been suggested that convergent evolution of morphological characters, such as reduction or loss of floral parts, makes it difficult to define a clear cut classification (Smissen et al. 2002). Some work has subsequently been done on the higher level classification of Caryophyllaceae using molecular techniques. These studies have shown that while the family Caryophyllaceae is monophyletic, none of the subfamilies as currently defined are and more work needs to be done to determine a phylogenetically robust classification (Smissen et al 2002, Fior et al. 2006). Harbaugh et. al. (2010) have proposed to abandon subfamilies within the Caryophyllaceae and recognize

28

J. Adelaide Bot. Gard. 25 (2011)

at least 11 tribes to deal with the polyphyletic nature of the family. They propose to include Ste//aria in the tribe Alsineae, which agrees with Pax & Hoftmann’s (1934) classification.

Subgeneric classification. The infrageneric classification of Stellaria was published by Fenzl (1840). This was then expanded by Pax & Hoffmann (1934). All of the Australian species were placed in the section Stellaria (as ‘Eustellaria’ Fenzl). They included the introduced species S. graminea, S. media and S. pallida as well as the endemic species S. flaccida and S. pungens in these classifications (Table 1). Pax & Hoffmann (1934) mentioned S. multiflora in the biogeographic discussion but did not place the species in their classification. As the majority of the Stellaria species were not included, the classification has not been taken up in Australian literature.

Morphology

Leaves. In the family Caryophyllaceae the leaves are most commonly opposite, decussate, usually sessile or subsessile or sometimes petiolate (Bittrich 1993). In Australia, the Ste//laria species have leaves that are either sessile, as in the majority of the ‘augustifolia group’, S. pungens and the introduced species S. graminea or appearing petiolate, as in the majority of the ‘multiflora eroup’ and S. flaccida, similar to what occurs in the introduced species S. media and S. pallida. On the narrow petiolate portion of the leaf there is always some tissue so it is not truly a petiole. There is a tendency that the petiolate nature of the leaf 1s more pronounced towards the base of the plant and this shortens and the leaf becomes sessile towards the apex and particularly in the flowering sections of the plant.

Leaf shapes in the family range from linear, needle shaped or grass-like to broadly ovate; some are rigid and acute or even spiny at the apex (Bittrich 1993). In the Australian Stellaria species the leaf shape has an even greater variation, ranging from filiform, linear, elliptic, ovate, obovate, as well as variation in width from narrow to broad. This character needs to be used with caution as the leaves can become progressively longer and narrower as the plant develops (Chinnappa & Morton 1984). In Australia the only species that may become rigid and has a spiny apex is S. pungens.

In the transition from the vegetative to the flowering parts of the plant either of two leaf states occurs. In the first state which 1s listed in the keys and descriptions as ‘Leaves a continuous series’, the structures subtending the flowers, which are technically bracts, appear the same as the lower leaves and one cannot distinguish between them as they remain the same texture, colour and shape but they gradually become smaller and, if petiolate, this becomes shorter and even sessile towards the extremities of the plant. The alternative character state which is listed in the keys and descriptions as ‘Leaves not a continuous series’ is characterized by

29

Stellaria in Australia (Caryophyllaceae)

the abrupt change from leaves to bracts where the size is markedly reduced, they are always sessile and the texture usually changes from green and leafy to white and scarious. However, one must be cautious in applying absolute values to leaf shape and size as it has been observed that these characters can be altered by altering environmental factors (Chinnappa & Morton 1984).

Inflorescence structure. In the Caryophyllaceae the inflorescence can be few to many flowered determinate (monotelic) thyrses, partial inflorescences that are mostly dichasia, or more rarely monochasia (Bittrich 1993). In the Austalian Stellaria species the most common inflorescence states are a monochasium, a few flowered partial monochasium or one to two flowered. Only larger specimens of S. angustifolia and all specimens of S. filiformis have the more common family character of dichasia, while all three introduced species, S. graminea, S. media and S. pallida, also have a dichasial inflorescence.

This type of inflorescence can cause problems in interpretation since the type of inflorescence can change as the plant matures, particularly from a single or few- flowered state to the partial monochasium to a many flowered monochasium. In the North American species, S. longipes, research has found that environmental factors can influence the number of flowers in the inflorescence and the development of the scarious floral leaves or bracts (Chinnappa & Morton 1984). The second character, the development of the scarious bracts, has not been observed to change in the only Australian species, S. filiformis, that has this character state. This study found that flower size was not affected by changing environmental conditions and is a good character to use when determining species.

Floral structures. In the Caryophyllaceae the flowers are generally actinomorphic and normally consist of three, four or five tetra- or pentamerous whorls (Bittrich 1993). In Stellaria there are five sepals with most commonly five petals which are usually deeply bifid. The stamens are usually in one or two whorls of five, and while the number may be reduced or supplemented with staminodes the total number is never more than ten.

In Australia there are two flower types. The first flower type is correlated with the ‘angustifolia group’, S. flaccida, S. pungens and the introduced species S. graminea. These have five sepals, five deeply bifid petals about the same length as the sepals and ten stamens in two whorls and no staminodes. This is the same pattern found in many of the Northern hemisphere taxa. The second type is correlated with the ‘multiflora group’ and the introduced species S. media and S. pallida. These have five sepals, five or less bifid petals that are small, usually less than half the length of the sepals, or reduced to small fragments or totally absent. There are usually two to five stamens and zero to five staminodes, except for subsp. multiflora and subsp. nebulosa which have three to ten stamens. When there are five or less stamens

C.H. Miller & J.G. West

J. Adelaide Bot. Gard. 25 (2011)

== Fa ; J ns > ee ie } ) J , . f =. Ripe "wa Og e - By Oe fa th Sa os gris

FOE a I>

Fig. 1. SEM of seeds. A-C Sfellaria angustifolia. A subsp. angustfolia; B subsp. rotundisepala; C subsp tenella. D S. filiformis. E S. graminea. F S. leotoclada. G S. media. H S. flaccida. | S. pallida. J S. papillata. K S. pungens. L-N S. multiflora: L subsp. multiflora; M subsp. collaris; N subsp. nebulosa. Scale bar: 100um. — A A.C. Beauglehole 82200 (CANB 364207); B JH. Willis s.n. (MEL 501962); C R.A. Black s.n. (MEL 1579640); D A. Cooper s.n. (NSW 145525); E A.H.S. Lucas s.n. (NSW 29863); F R. Coveny et. al. 8954 (NSW 298626); G C.H. Miller & J. Palmer 590 (CANB 409321): H WM. Curtis s.n. (HO 29492): | C.H. Miller & J. Palmer 603 (CANB 409333); J J. Carrick 3135 (AD 97207083); K J. McKean WL 5189 (CANB 327583); LL.G. Adams 1644 (CANB 166831); M H.S. McKee 11668 (CANB 319264): N JS. Whinray 1150 (CANB 442426).

present they are positioned opposite the sepals and have a slightly enlarged base.

Fruits and Seeds. There are two to five or rarely more styles, free to the base and clearly distinct from the ovary, a character that occurs throughout the subfamily

30

Alsinoideae (Bittrich 1993). In Stellaria the style number is usually three, rarely two, four or five. This also holds for the Australian species where it is extremely rare to find more than three styles.

For the tribe Alsineae the fruit is a capsule that normally opens with valves or teeth equal to or twice

J. Adelaide Bot. Gard. 25 (2011)

the number of carpels, or is rarely indehiscent (Bittrich 1993). In Stellaria the number of valves is twice the number of carpels and in the Australian species this is six, very rarely more. The surface texture of the capsule valves is usually translucent to opaque and relatively smooth. There is one species in Australia, S. papillata that has the surface covered in distinct papillae. On maturity the valves split, usually for more than half their length or almost to the base and the apices of the valves displace, either moving only slightly outward so the valves remain almost straight, the apices curve out so they are recurved or the valves curl completely around to be revolute. Also the valve alignment can either remain in place, retaining the capsule shape or displace outwards, giving a spreading appearance. Capsule characters such as colour and angle of the valves have been shown to be defining characters as they are not affected by environmental factors (Chinnappa & Morton 1984).

In Stellaria the seeds are small (0.4—3 mm long), numerous, rarely few or one, roundish to reniform, usually laterally compressed and the testa 1s variously sculptured by more or less papilliform cells, rarely completely smooth (Bittrich 1993). In Australia, Stellaria has these seed characters and further study is needed to determine reliable characters to separate the taxa. The cell shape is always stellate, although this can be sometimes hard to see, particularly in S. filiformis (Fig. 1D), which has a very small, light-coloured seed, c. 0.5 mm long, that has a semi translucent testa and 1s smooth with small raised pits in the middle of the cells. The rest of the taxa have well defined stellate cells that are often inflated in the centre to form either a ridge when the cell is long and thin, such as S. papillata (Fig. 1J) and S. angustifolia subsp. angustifolia (Fig. 1A), or hills when the cell shape is more rounded, such as S. multiflora subsp. nebulosa (Fig. IN). The cells are much more raised on the outer dorsal surface of the seeds. In some cases these inflated cells become almost finger- like, such as in S. flaccida (Fig. 1H) and S. pungens (Fig. IK), and in an extreme case the finger-like projections cover the entire seed so that the stellate cell shape can no longer be seen, such as in S. angustifolia subsp. rotundisepala (Fig. 1B), which is not seen in any other taxa in Australian Stellaria. The introduced species, S. media (Fig. 1G) and S. pallida (Fig. 11), also have the addition of small papillae on the arms of the stellate cells, a character that 1s not present in any of the Australian species of Stellaria.

From the North American S. /ongipes study none of the morphological characters was shown to be associated with polyploidy; nor was there any consistent association between chromosome number and geographic distribution (Chinnapa et al. 2005). From this study it was shown that the Arctic populations were generally low, single flowered plants lacking scarious bracts, while progressive development in internode length, flower number and presence of scarious bracts were observed in more southerly populations. In

5]

Stellaria in Australia (Caryophyllaceae)

Australia Ste//aria 1s restricted to the southern part of the continent and no obvious association can be observed between morphological traits and latitude. There may be reason to think that the predominance of the monochasial inflorescence 1s due more to the harsher, drier environmental conditions found in Australia rather than a difference in latitude. The presence of scarious bracts is only found in the Australian species S. filiformis which has a distribution across the southern part of Australia from Wyalong in New South Wales to Cowcowing in Western Australia in sandy soils with high summer temperatures and low rainfall. There are more of the monchasial species, such as those in the ‘angustifolia group’ and the ‘multiflora group’ that grow both north and south of S. filiformis.

Typifications

Ronald C. Gunn and Robert W. Lawrence collections from Tasmania. Some of the type material for the Australian Stellaria species is from collections made by Gunn and Lawrence in Tasmania. Due to a number of peculiarities of their collecting and recording methods some difficulties have arisen in determining type material. The numbers on Gunn and Lawrence labels refer to their species numbers and not collecting numbers. There is often more than one collection that has the same number but different collection dates. Where there is only a year written on the label this indicates the year of dispatch of the parcel to London and not the actual collection date of the collection. There are cases where there is also a full collection date on the same label and this is later than the dispatch date as the parcel was sent later than expected. Some of these collections were collected by other people and Gunn arranged these collections within his own numbering system (Buchanan 1988).

Collections from England. All material from K and BM was searched for types. Many of the names published by Bentham are typified with material from Herb. Hookerianum (K), often annotated by Bentham. No type specimens were found in Bentham’s own Herbarium (Herb. Benthamianum, K).

Materials & methods

This review has been based on dried material from the following herbaria: AD, BM, BRI, CANB (incl. CBG), CHR, DNA, HO, K, LD, MEL, NSW, OXF and PERTH. Limited field work was undertaken in New South Wales, the Australian Capital Territory, Victoria and New Zealand.

For Scanning Electron Microscopy (SEM), Fig. 1, seeds were mounted onto 10 mm diameter metal stubs using double sided tape. All samples were coated with gold before observation in a Jeol 6400 Scanning Electron Microscope at an accelerating voltage of 15kV. Images taken using B&W polaroid camera. Facilities provided by the CSIRO Black Mountain Microscopy Centre, A.C.T.

C.H. Miller & J.G. West

Taxonomic Treatment

Stellaria L.

Sp. Pl. 1: 421 (1753). — Type species: S. holostea L.., fide Hitchcock & Green, Prop. Brit. Bot.: 155 (1929).

Annual or perennial herbs, glabrous or sparsely hairy. Leaves opposite; stipules absent; floral leaves herbaceous to scarious. Flowers in spreading dichasia to partial monochasia, or solitary or 2-flowered, 5-merous, bisexual. Sepals 5, free, usually narrow to broadly ovate. Petals usually 5, free, bifid, shorter to longer than sepals, in some taxa reduced or absent. Stamens 1-10. Staminodes 0-5. Ovary 1-celled; styles 3, free to base; placentation free-central. Fruit a conical capsule, splitting to middle or below, 6-valved. Seeds 2 to many, subdiscoid to nautiloid to obloid; testa minutely to prominently tuberculate.

Distribution & habitat. A genus of about 120 species (Morton 2005) found throughout temperate areas worldwide and at higher altitudes in tropical areas. There are a few weedy species, such as S. media and S. graminea that occur almost worldwide.

In Australia Stellaria is represented by 10 species including both endemic and introduced elements. The genus is distributed in temperate areas from southern Queensland, throughout New South Wales, Victoria, Tasmania, the southern parts of South Australia and Western Australia and central parts of the Northern Territory. There are three introduced species of which two, S. media and S. pallida, have become naturalised and distributed throughout the native range of the genus in Australia and extend northwards to the tropics in Queensland, the Northern Territory and Western Australia. A third introduced species, S. graminea, has a very localised distribution in alpine areas of New South Wales and Victoria, subalpine areas in Tasmania, and the hills of southern Fleurieu Peninsula, South Australia.

Stellaria 1s found in a wide range of habitats in Australia from coastal dunes, heathlands, grasslands, herblands, swamps, fern communities in gully forests, underlying open eucalypt or Callitris woodlands, rainforests, alpine bogs, mallees and Chenopodiaceae or Leptospermum scrublands. The endemic species are often associated with wetter areas such as watercourses, areas that seasonally flood, drainage areas or dams. The weedy species are often associated with disturbed areas such as roadside drains, dams or cultivated areas. They can grow in a wide variety of soil types from sands to clays and on almost any rock type.

The native Australian species of Stellaria are endemic and do not occur in nearby countries such as New Zealand and Papua New Guinea. New Zealand has five native species and four introduced species (Webb et al. 1988). Two of their introduced taxa, S. media and S. graminea, also occur in Australia. One of the Australian species, S. flaccida, shows affinities with the New Zealand endemic S. parviflora Hook.f. Much less is known about the flora of Papua New Guinea, with

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J. Adelaide Bot. Gard. 25 (2011)

only one introduced species, S. media, and one native species, S. stellatopilosa Hayata, present (van Royen 1982). S. stellatopilosa appears to have affinities with S. decipiens Hook.f. from New Zealand.

Etymology. The name derives from the Latin stella (a star) and aria (connected with), an allusion to the radiating, deeply bifid petals.

Note. Species are treated here in alphabetical order as there is no satisfactory infrageneric treatment available at this time.

1. Stellaria angustifolia Hook.

J. Bot. (Hooker) 1: 250 (1834). — Type citation: ”Mr. Lawrence, Formosa, (n. 241,) Mr. Gunn, (n. 238.)”. Lectotype (here designated): Formosa, Lawrence n. 241, Aquatic (K, Herb. Hooker p.p. [3 pieces to the right hand side of the pencil line]). Isolectotype: RWL |awrence] 241, Aquatic. (NSW). Residual Syntypes: [Gunn] 238, 1833 (NSW p.p.); Gunn’s Herbarium of Tasmanian Plants. (n. 238). (NSW p.p.).

Stellaria caespitosa Hook.f., J. Bot. (Hooker) 2: 411(1840). — Stellaria glauca var. (?) caespitosa (Hook.f.) Benth. FI. Austral. 1: 158 (1863). — Type citation: “In a marsh at Circular Head. Mr. Gunn (n. 652 and 653°)’. Lectotype (here designated): In a marsh at Circular Head, Gunn 652?, 1837 (K, Herb. Hooker p.p. [2 pieces on the top right hand side of the pencil line]).

Stellaria palustris auct. non Ehrh. ex Retz.: F.Muell., Key Syst. Victorian Pl. 1: 166 (1887); C.Moore, Handb. FI. N.S.W. 99 (1893); W.A.Dixon, Pl. N.S.W. 77 (1906); J.M.Black, Fl. S. Austral. 2: 231 (1924); Ewart, FI. Victoria: 491 (1931); J.H.Willis, Handb. Pl. Victoria 2: 136 (1972); W.M.Curtis, Stud. Fl. Tasman. |: 70 (1975); K.Chorney in Jessop & Toelken, Fl. S. Austral. 1: 235 (1986).

Stellaria glauca auct. non With.: Benth., Fl. Austral. 1:158 (1863); Tate, Handb. Fl. Extratrop. S. Austral. 43 (1890); F.M.Bailey, Queensl. Fl. 1: 87 (1899).

Annual or perennial, stems tall and slender to spreading or prostrate, with slender rhizome which may root from nodes; stems (5—) 11—60 (—90) cm long, usually single stemmed or branching from base, glabrous to scabrous. Stem and inflorescence leaves forming continuous series, sessile, not clasping, narrowly ovate, linear, narrowly elliptic or narrowly obovate, (3.5—) 10-40 (—60) mm long, 0.9-3 (—3.9) mm wide, apex acute to subacute, rarely obtuse, margin entire, rarely recurved, glabrous to scabrous all over, sometimes with 2—5-celled hairs on margins. /nflorescence axillary and solitary, interrupted monochasia of 2—7 flowers, monochasia of 5—7 flowers or rarely dichasia of 7—23 flowers. Pedicels (15.5—) 20—72 (—90) mm long, slender, erect in fruit, glabrous to scabrous, often quadrangular, ridged or grooved. Sepals 1.5—8 (—9) mm long, acute to acuminate, obtuse, or folded over to form hood, sometimes with hood thickened, the margin entire, rarely minutely ragged. Petals 2.4—7.5 (—9) mm long, usually about equal to sepals to twice sepal length; bifid to deeply bifid. Stamens 10. Staminodes 0. Styles 0.8-4 (—5.3) mm long. Capsule ellipsoid to ovoid, (2.8—) 3.5— 6.8 (—8.5) mm long, (1.8—) 2.44 (—5) mm wide, from

J. Adelaide Bot. Gard. 25 (2011) Stellaria in Australia (Caryophyllaceae)

Key to Species of Ste/laria in Australia

1. Leaves sessile but lower leaves at least, appearing petiolate 2. Internodes with single line of hairs for entire length 3. Petals present; fertile stamens 3—5, rarely 2; all seeds dark in colour, usually > 1 mm long ............. 6. * S. media 3: Petals absent; fertile stamens 2, rarely more; at least some seeds pale to midbrown,< 1mm long....... 8. * S. pallida 2: Internodes without single line of hairs 4. Petals present, > '2 sepal length, > 2 mm long 5. Plant erect, fine-stemmed, to 20 (—30) cm long; sepals glabrous; fruit < 5 mm long, < 2.5 mm wide .. 5.8. leptoclada

5: Plant lax, to 50 cm long; sepals with hairs or ciliate along some margins, often hairs also on sepal back: orinidrib; trait 3 minv long; > 2. SM WIE i, be ecw eee ee ns ee be ge eee ee ee 3.8. flaccida

4: Petals absentor 1f present, < 4'sépal leneth; <2 mm lone 0c nae ee ie RA OS eee AMR as 7.8. multiflora 1: Leaves sessile 6. Petals absent or 1f present < '/2 sepal length, < 2 mm long 7. Inflorescence a dichasium; stem leaves in basal rosette, filiform, herbaceous to fleshy; floral leaves linear to ovate to lanceolate, usually scarious; seeds with minute tubercles, appearing smooth (Fig. IE Ie A 24 evr Ear we oad a al LTC TA NEN fet, bed Bat ar aN aT NUP ICTR AT ONE bea A Ean wr oe AC SELLE ICER ae SES eH Ae OA We fyi cater HNL? RE AT8 2. 8. filiformis 7: Inflorescence axillary and solitary or a monochasium; leaves in continuous series, stem leaves not in basal rosette and not filiform; floral leaves herbaceous, never scarious; seeds with prominent tubercles, appearing reticulate (Fig. 3h—n) 8. Capsule valves thick, opaque, covered with angular papillae; seeds obloid to ellipsoid, 1—2 per

egosule: 1S) Oo MONG, bt snes gee Sad sopnck tet oe ok dessa ene aed iad, tet hy ed dew eb apeare ied eh he ec 9. S. papillata 8: Capsule valves thin, semitransparent, smooth; seeds suborbicular or kidney-shaped or broad- ellipsoid, more than 2 per capsule,< 1.3mm long ......... 0... ee teen ees 7.8. multiflora

6: Petals present, > ’2 sepal length > 2 mm long 9, Flowers 5 or more, usually either a monochasium or dichasium 10. Stems, leaves and sepals sparsely to densely covered with long curly hairs; leaf apex usually plinsent: Sepal apex. vishally pingent, mEeVver HOOdEd! ssc... 5 oe ee EEE EADS 9G 4 EH FREES 10. S. pungens 10: Stems, leaves and sepals glabrous, scabrous or if hairs present then these not long or curly; leaf apex acute to subacute, never pungent; sepal apex acute to obtuse, never pungent, hooded or not 11. Leaves not a continuous series; floral leaves much smaller, scarious, sparsely to densely ciliate along entire length; outer sepals often ciliate along entire length; sepal apex not hooded; INTIOCESCENCE SICH AS ILM 4. le ent ss ace eee 2 yal tees le nti Soles Sea's etal ial oh le loin’: o¥ce sens ata talaiyl« 4. * S. graminea 11: Leaves a continuous series; floral leaves herbaceous, glabrous, scabrous or with short hairs basally; sepals glabrous or scabrous; sepal apex hooded or not; inflorescence a monochasium, an interrupted monochasium or solitary, rarely a dichasium 12. Stems usually glabrous to scabrous; leaves never in basal rosette; sepal apex straight, rarely

hooded: seed usually > Wmartelone .. 24.) 4-4 oie selec ay ee He 1. S. angustifolia (mostly subsp. angustifolia) 12: Stems sparsely covered with short hairs or scabrous; leaves often arising from loose basal rosette; sepal apex straight, not hooded; seeds usually < 1mm long ....................... 5. 8. leptoclada

9: Flowers usually solitary, rarely 2—3-flowered

13. Stems, leaves and sepals sparsely to densely covered with long curly hairs; leaf apex acute to plnsenl= sehal apex Acute te purrcent,. ever HOOdE 6-5-6. dcace sce eins gotee Seeiele aletpce ee ace use ernee Soatele ols 10. S. pungens

13: Stems, leaves and sepals glabrous, scabrous or rarely some margins with short hairs; leaf apex acute to subacute, not pungent; sepal apex acute to obtuse, not pungent, hooded or straight............ ESE EES tar eles ela de EE TR OE LO Sees eeliee ee eR SS Wd LO 1. S. angustifolia (mostly subspp. rotundisepala and tenella)

1/3 shorter to 4 times longer than sepals; valves straight | from the European specimens, but retained the taxon in to spreading, apex straight, recurved to revolute, rarely S. glauca and placed S. angustifolia in synonymy. He twisted sideways. Seeds 2—33, suborbicular to ellipsoid, also reduced S. caespitosa to a variety under S. glauca. flattened top and bottom, (0.5—) 0.8—1.2 (-1.4) mm long, Other authors have used S. palustris and have also used light brown to grey brown to reddish brown or purplish, |§ Bentham’s infraspecific taxa, but at different ranks.

tubercles ridged, wrinkled to rounded and narrow, spiny Stellaria palustris specimens from England and in appearance. northern Europe have erect, linear leaves on the stems and floral leaves that are markedly shorter, ovate to narrowly ovate and often scarious and the inflorescence is usually a terminal dichasial cyme. S. angustifolia specimens have linear to lanceolate leaves, floral leaves which are In a continuous series, and the inflorescence can be Notes. Further study of this taxon 1s much needed. axillary and solitary, or an interrupted monochasium, Historically, this taxon has been referred to the European a monochasium or occasionally a dichasium in larger species either S. palustris or S. glauca (asynonym of S. _ plants. As the species can usually be distinguished from palustris) in the Australian literature. Bentham (1863) — each other, S. angustifolia is treated here as a separate noted instances where the Australian plants differed — entity from the widespread European taxon S. palustris.

Distribution. A widespread endemic species in temperate Australia, from southern Queensland, throughout New South Wales, Victoria, southern South Australia (including Kangaroo Island) and Tasmania.

53

C.H. Miller & J.G. West J. Adelaide Bot. Gard. 25 (2011)

Sat ll

| Ny f

i

del: da sors

Fig. 2. A-J Stellaria angustifolia subsp. angustifolia: A habit; B flower; C flower with petals & sepals removed; D gynoecium; E part of flower showing sepal, petal and stamen arrangement; F petal; G capsule showing position in relation to sepals; H mature capsule; | seed, side view; J seed coat cell detail. K-P S. angustifolia subsp. tenella: K habit; L part of tower showing sepal, petal and stamen arrangement; M petal; N capsule; O seed, side view; P seed coat cell detail. Q-W S. angustifolia subsp. rotundisepala: Q habit; R part of flower showing sepal, petal and stamen

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J. Adelaide Bot. Gard. 25 (2011)

It 1s evident that the taxa in the ‘angustifolia group’ are closely related, particularly in their vegetative characteristics. The taxon which has been consistently referred to as var. tenella can be separated from other plants by its smaller size, mat forming habit with short internodes, sepals that end in a hooded structure, petals that are usually about twice the sepal length and flowers that are almost always solitary and axillary. However, under favourable growing conditions the plants can become larger, more erect and with longer internodes and more flowers. The plants at the smaller end of the spectrum can for the most part be readily separated and match well with the type specimens of var. tenella. Bentham (1863) compared var. (?) tenella to var. caespitosa, but commented that tenella is smaller and much more slender, with crowded, very small leaves.

Applying the epithet var. caespitosa to plants 1s more problematic as most of the characters used are size related and this is very dependent on the environmental conditions in which the plants grow. Examination of the type specimen of S. caespitosa has revealed that the plants have more characters in common with the type specimen of S. angustifolia than the type specimen of var. tenella. They are closer in size and there are more flowers per plant, the leaves are similar in shape, and while the growth is sometimes mat-forming, the leaves never seem to be as fine as those of var. tenella. The sepals are often subacute and slightly hooded especially in bud, but they can usually be differentiated from var. tenella plants, particularly when in fruit. We feel that var. caespitosa is toward the smaller end of the S. angustifolia spectrum, but there are not enough consistent characters to warrant the separation of the taxon from S. angustifolia. Bentham (1863) considered S. glauca var. caespitosa distinct in its inflorescence characters but commented that specimens showed “a very gradual passage” towards S. glauca in leaves as well as sepals.

Bentham’s taxon var. (?) leptoclada is always recognisable from the other taxa in its growth form and inflorescence structure as well as its limited distribution, and has been raised to species level here.

Nomenclatural notes.

Stellaria angustifolia Hook. One sheet from Herbarium Hookerianum held at K contains type material. The sheet has two collections. The first label on the right hand side of the pencil line is ‘Aquatic Formosa. Van D’s Land. n. 241. Lawrence’; it matches the protologue and locality information and is thus designated as the lecotype. A separate label with ‘Arenaria L.R.’ is a Cunningham collection from the Lachlan River in NSW and 1s not type material.

Stellaria in Australia (Caryophyllaceae)

Three sheets from NSW contain type material. The first sheet has three collections with the following labels; ‘238/1833’, ‘238/1842 Marsh Formosa 7/12/42’ and °238/1842 Marlborough 5/1/41’. Only the specimen with the label ‘238/1833’ has been designated as a residual syntype as the other specimens post-date publication. The second sheet has a single plant with the label “RWL]awrence| 241 aquatic’ on it and it has been designated as an isolectotype. The third sheet has the label ‘Gunn’s Herbarium of Tasmanian Plants 238’ and has been designated as a residual syntype.

Stellaria caespitosa Hook.f. One sheet from Herb. Hookerianum held at K contains type material. The sheet has three collections. Only the collection on the top of the sheet on the right hand side of the pencil line with the label ‘ In a marsh at Circular Head, Gen 652°, 11 Jan 1837’ matches the protologue and is here designated as the lectotype. The collection with the label ‘ Oatlands, [Gunn] 652, Nov 1835 also contains line drawings of floral and seed characters drawn by Hooker. The sepal illustrated 1s a typical S. angustifolia shape with an acute apex which contradicts Hooker’s description of S. caespitosa which 1s “calycibus ovato- lanceolatis subacutis”’.

Key to subspecies of S. angustifolia

1. Sepals (3—) 4-8 (—9) mm long, apex acute to acuminate; petals (3.5—) 4.5—7.5 (—9) mm long, about equal to sepals; fertile stamens > 3 mm long...... a. subsp. angustifolia

1: Sepals 1.5—2.5 mm long, apex folded over to form hood; petals 2—3.2 (—3.9) mm long, longer than to twice sepal length; fertile stamens < 3 mm long

2. Sepals narrow ovate to elliptic, apex obtuse or acute, hood never thickened; pedicel usually < 20 mm long; seed tubercles sem1-inflated ridges... b. subsp. tenella

Sepals round or broad elliptic, apex obtuse, hood

sometimes thickened; pedicel > 20 mm long, seed spiny,

with narrow, pointed prominent tubercles. ..........

c. subsp. rotundisepala

la. Stellaria angustifolia Hook. subsp. angustifolia

Illustration. C.H.Mill. & J.G.West in N.G.Walsh & Entwisle, Fl. Victoria 3: 236, Fig. 45e-f (1996), as S. angustifolia.

Annual or perennial, weak to tall and spreading to 75 (—90) cm long, glabrous to scabrous. Stem and inflorescence leaves as a continuous series, sessile, linear, narrowly ovate to lanceolate, rarely obovate (9—) 10—40 (—60) mm long, 0.9—3 (—3.9) mm wide, acute or rarely subacute, margin rarely recurved, occasionally with short hairs. /nflorescence solitary, an interrupted monochasium, monochasium, or rarely a dichasium. Pedicels (15.5—) 20—72 (—90) mm long, slender, erect in fruit. Sepals narrowly to broadly ovate or elliptic, (3.5—) 4-8 (-9) mm long, acute to acuminate. Petals (3.5—-)

arrangement; S petal; T-U flower with immature (T) and mature (U) capsule; V seed, side view; W seed coat cell detail. X-DD S. leptoclada: X habit; Y part of flower showing sepal, petal and stamen arrangement; Z petal; AA flower showing capsule; BB mature capsule; CC seed, side view; DD seed coat cell detail. Scale bars: habits (A, K, Q, X) 5 mm; flowers & fruits 1mm; seeds (I, O, V, CC) 0.5 mm. — A, E-H R. Schodde 1224 (AD 96227132); B—D D. Hunt 2876 (AD 96913165); I-J Bushman 214 (AD 98232306); K-P R. Bates 21927 (AD 99010076); Q-W R.J. Bates 27162

(AD 99206284): X-DD JR. Hosking s.n. (CANB 00710190).

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C.H. Miller & J.G. West

4—7.5 (-9) mm long, usually about equal to sepals; bifid. Stamens 10, filaments 3.3-6 (—6.5) mm long. Staminodes 0. Styles 2.2-4 (—5.3) mm long. Capsule ellipsoid to ovoid, (2.8—) 3.5—6.8 (—8.5) mm long, (2—) 2.5—4 (—5) mm wide, shorter than to equal to sepals or sometimes slightly longer; valves straight to spreading, apex straight, recurved to revolute, rarely twisted sideways. Seeds 8-33, suborbicular to ellipsoid, (0.6—) 0.8—1.2 (—1.4) mm long, light brown to grey brown to reddish brown; tubercles semi-inflated ridges. Fig. 1A, 2A—J. Flowering: Sep.—Mar.

Distribution and habitat. The taxon is widespread in southeastern Australia. It occurs south from Stanthorpe in Queensland, as well as west in the Warrego and Darling River systems. In New South Wales, it occurs along the Great Dividing Range from Armidale, through the Blue Mountains, along the Southern Highlands and alpine areas of Kosciuszko National Park. In Victoria, it also occurs in alpine areas along the Great Dividing Range and as far south as Port Phillip as well as to the west across mountain areas and in the Delatite, Delegate and Dry River systems. It also occurs along entire length of the Murray River system. It is rare in Tasmania where it occurs in the Ouse River system. In South Australia, it occurs in higher altitudes, in the Mt Gambier area, as well as the entire length of the Mt Lofty Ranges to Fleurieu Peninsula and Kangaroo Island (Fig. 6A).

Locally common, widespread herb growing in wet areas amongst grasslands, herblands, sedgelands, lignum thickets in swamps, along watercourses particularly after flooding or underlying in open woodlands or Acacia or eucalypt forest; rarely in disturbed areas such as roadside drains or dams. Most commonly found in higher altitude areas of ranges above 500 m, rarely found at sea level, where it 1s associated with large rivers. Grows in rich soils of basalt origin or deep cracking clays through to light sandy soils derived from granites.

Selected specimens (of c. 300 seen)

QUEENSLAND: Darling Downs, 6 km N of Goondiwindi, 8 Sep. 2001, 4.R. Bean 17796 (CANB); Darling Downs Wyberba, 30 Dec. 1962, S.T’ Blake 21994 (MEL, NSW); Warrego District, Currawinya National Park, S end of Corni Paroo Waterhole, 17 Sep. 1992, R.W. Purdie 4151 (CANB).

New SoutH WaALEs: North Western Plains, 7.5 km N of Bruxner Highway at Yetman on road to Yelarbon, 7 Oct. 1990, R.G. Coveny & R.O. Makinson 14480 (CANB, MEL, NSW); Ca 10 miles S of old Jindabyne, 8 Jan. 1963, C.W.E. Moore 3594 (CANB, NSW); °Winbar’ about 40 miles S of Louth (Tundulya block area near mill), 28 Sep. 1976, C.W.E. Moore 7390 (CANB); Black Swamp, 55 km N of Deniliquin, Oct. 1974, WE. Mulham 791 (NSW).

AUSTRALIAN CAPITAL TERRITORY: Cotter River district, Between Jack’s Creek and the Cotter River, 14 Dec. 1960, R. Schodde 1224 (AD, CANB, NSW).

VicToRIA: Barmah State Park, Murray Valley Study area Sector F, subblock 40B, 18 Nov. 1985, A.C. Beauglehole 82200 (CANB, MEL, NSW); E. Highlands, Wonnangatta Stn., upstream from Dry Rv. S17, 7 Jan. 1993, E.A Chesterfield 3570 (CANB, MEL); Dargo High Plains, 17 Jan. 1990, J. Strudwick 804 (MEL).

36

J. Adelaide Bot. Gard. 25 (2011)

TASMANIA: Central Highlands, Wihareja Lagoon 9 km NE of Waddamana, Feb. 1984, A. Moscal 6571 (HO); Stone Hut, between Ouse River and Great Lake, 25 Jan. 1980, J./. Yates s.n. (HO).

SOUTH AUSTRALIA: Kangaroo Island, Ravine des Casoars, s.dat.. RJ. Bates 30356 (AD, CANB); Murray, Upper Saunders Creek Gorge, s.dat., R./J. Bates 35608 (AD, CANB); Southern Mount Lofty Range, ca 2 km S of Spring Mount Trig point, which is ca 8 km SE of Myponga (Myponga is near coast, ca 55 km SSW of Adelaide), 23 Nov. 1966, 4.4. Shaw 734 (AD, CANB, NSW).

lb. Stellaria angustifolia subsp. tenella (Benth.) C.H.Mill. & J.G.West, comb. nov. Stellaria glauca With. var. (?) tenella Benth., Fl. Austral. 1: 158 (1863). — Stellaria palustris Retz. var. tenella (Benth.) J.M.Black, Fl. S. Austral. 2: 231 (1924). — Type citation: “Victoria. Near Melbourne, Adamson; Glenelg river, Robertson. Tasmania. Derwent river and Kitt’s Group in Bass’s Straits, R. Brown; granite rocks in St. Patrick’s river, Gunn’. Lectotype (here designated): Victoria, Glenelg river, Robertson (K, Herb. Hooker). Residual syntype: Victoria. Melbourne, -? Adamson, 17/4/[18]53 Us, Herb. Hooker). Excluded syntypes: R. Brown, Derwent River (BM, K p.p.); V. D. Land, Gunn 652, St Patrick’s River 16/11/[18]44 (kK, Herb. Hooker). — see S. multiflora. Arenaria axillaris Luehm. ex Ewart, Victorian Naturalist 23: 42—43 (1906). — Stellaria glauca var. axillaris (Luehm. ex Ewart) Ewart, Proc. Roy. Soc. Victoria 19(2): 34 (1907). — Type Citation: “Mr. St. Eloy D’ Alton, C.E.; from a peat swamp near Dimboola”’. Holotype: MEL, n.v. Isotype: NSW 117582. Stellaria caespitosa auct. non Hook.f.: C.H.Mill. & J.G. West in N.G. Walsh & Entwisle, Fl. Victoria 3: 235 (1996).

Illustration. C.H.Mill. & J.G.West in N.G.Walsh & Entwisle, Fl. Victoria 3: 236, Fig. 45h-1 (1996), as S. caespitosa.

Annual, spreading mat, stragegly or shortly erect herb to 14 (-20) cm long, with many short, vegetative side shoots, rooting at nodes, glabrous or rarely scabrous. Stem and inflorescence leaves a continuous series, sessile, lanceolate, narrowly ovate or linear, often falcate, (1.5—) 2-10 (-17.5) mm long, (0.4—) 0.6—1.5 (—2.3) mm wide, acute. /nflorescence axillary and solitary or rarely 2 flowers on same stem. Pedicels (2.7—) 6—14.5 (—20) mm long, usually deflexed in mature fruit. Sepals elliptic to ovate, (1.3—) 1.6—2 (—2.4) mm long, obtuse or acute, apex folded over to form a hood, hood 0.1-— 0.45 mm long, sometimes margins minutely serrate or ragged. Petals 2—3.2 (—3.9) mm long, slightly longer to twice sepal length, deeply bifid. Stamens 10, rarely 1 or 2 missing. Staminodes 0. Styles (1.5—) 1.9—2.6 mm long. Capsule ellipsoid or ovoid, (2.5—) 2.9-3.7 mm long, (1.4—) 1.6—-2 mm wide, equal to up to twice as long as sepals; valves straight or spreading, apex only straight or slightly recurved. Seeds 4—9, ellipsoid or suborbicular, 0.9-1.2 mm long, red to dark ruby red, tubercles semi- inflated ridges. Fig. 1C, 2K—P. Flowering: Sep.—May.

Distribution and habitat. The taxon occurs in New South Wales from Rules Point on the Murrumbidgee

J. Adelaide Bot. Gard. 25 (2011)

River, south to Lake Eucumbene, Hume Weir then west along the entire length of the Murray River system and in surrounding drainage areas. In Victoria it occurs in the east around the Goulburn River and coastal areas near Sale and in the west, it is scattered through the Grampians and Dimboola areas. In South Australia it occurs in the Naracoorte to Mount Gambier area and in the Mount Lofty Hills. In Tasmania it occurs on the north and west coasts and is recorded from one locality in the highlands at Oatlands (Fig. 6B).

Locally common in moist areas around swamps, rivers, lakes or dams. Often found growing in muddy or grassy areas after water has receded under open eucalypt woodlands. Found growing in rich damp soils.

Nomenclatural Notes.

Stellaria glauca var. tenella Benth. Four sheets from K contain type material. One sheet from the Herbarium Hookerianum has a label ‘1/20 44 Glenelg river 401’ and written on the sheet is ‘Victoria Robertson’. Additionally the name ‘Stellaria glauca var tenella’ 1s written in what is thought to be Bentham’s handwriting. This is a match to the locality in the protologue and the specimens match the description with very small leaves, few small flowers and sepals being rather obtuse. As these specimens are the best match to the protologue, this sheet is here designated as the lectotype.

The second sheet from the Herbarium Hookerianum has a label with ’65 Melbourne 17/4/53’ and written on the sheet is ‘F. Adamson’ and again ‘var tenella’ 1s written in what is thought to be Bentham’s handwriting. The collection matches the locality and specimen details in the protologue but has fewer plants so it has been designated as a residual syntype.

The third sheet has two collections. The collection above the pencil line has a blue label with ‘R. Brown, Iter Australiense, 1802-5 No. 5210 Stellaria glauca With. var. tenella Derwent River’. The collection below the pencil line has a blue label with “R. Brown, Iter Australiense, 1802—5 No. 5210 Stellaria glauca With. var. tenella Kent’s Group Bass Straits’. The top collection matches Bentham’s locality data but the bottom collection has Kent’s Group rather than Kitt’s Group as in the protologue. Additionally, the specimens do not match the description, as the sepals are acute rather than blunt. The collections are both S. multiflora and there is no indication that this sheet belonged to Herbarium Hookerianum, so the top collection 1s considered here to be excluded syntype material.

The fourth sheet is from the Herbarium Hookerianum and has the following labels: ‘652 St Patricks Rv 16/11/44 V. D Land Gunn;’ and ‘652 Collected on Granite rocks at St Patricks River at an elevation of about 2000 feet’. Again while the locality information matches the protologue, the specimens of this collection do not match the description with the sepals being acute rather than obtuse. The specimens are all S. multiflora and so these are considered here to be excluded syntype material.

Stellaria in Australia (Caryophyllaceae)

One sheet from BM contains type material. It has a blue label with ‘R. Brown Iter Australiense 1802—5 No. 5210 Genus Caryophyll prop Polycarpiem & Alsenem Derwent V D Land 1804’. This collection matches the collection on the top half of the third Kew sheet. The specimens of this collection are S. multiflora and do not match the description, and so are considered here to be excluded syntype material.

Arenaria axillaris Luehm. ex Ewart. From examination of the isotype in NSW this taxon belongs to S. angustifolia subsp. tenella. At present the specimen from MEL has not been seen. This specimen is referred to being in the Melbourne Herbarium by Ewart in 1906. A search of MEL needs to be undertaken to ensure it still exists.

Selected specimens (of c. 105 seen)

New SoutH WALES: Barmah adjacent to Murray River upstream from Echuca, 22 Feb. 1979, E.A. Chesterfield s.n. (NSW); Hume Weir, Albury District, 29 Dec. 1965, R.J. Flynn s.n. (NSW); Southern Tablelands, Kosciuszko National Park, southern end of Tantangara Reservoir, 26 Jan. 2007, R.W. Purdie 6325 (CANB); Lake Victoria, Far SW of NSW, June 1945, Tolley s.n. (AD).

VicTorIA: Kulkyne Forest, Far NE corner, Chalka Creek, 24 km (15m) NE of Hattah PO, 14 Oct. 1972, A.C. Beauglehole 40570 (MEL); Wallpolla Island, 26 Oct. 1972, A.C. Beauglehole 40649 (MEL); Barmah Regional Park Grid 152: Murray Valley Study Area, | Jan. 1985, A.C. Beauglehole $3536 (MEL); Corangamite Study area, between Lake Terangpom and Lake Gnarpurt, 26 km WSW of Cressy PO, 24 km NE of Camperdown, 15 Oct. 1977, G.J/. Hirth s-.n. (MEL); Nyah State Forest about 4 km N of Nyah, 6 Feb. 1977, N. Macfarlane 985 (MEL); Cairn Curran Reservoir, near Pyrenees Highway Bridge, Grid J9, North Central Study Area, 11 Jan. 1983, E.E. Perkins s.n. (MEL); Lake Hume, just east of old Tallangatta, 0.5 km east of Tallangatta Vallet turnoff on Murray Valley Highway, 22 Jan. 1981, N.H. Scarlett 81-22 (MEL); Wangaratta to Milawa road, c. 9-10 km SE of Wangaratta. Oxley Flats, 8 May 1980, /. Tankard s.n. (MEL).

TASMANIA: Oatlands, 4 Jan. 1913, R.A. Black s.n. (MEL); Pennerowne Point, 26 Jan. 1984, A.M. Buchanan 2822 (HO); Mouth of Lagoon River, Jan. 1954, W.D. Jackson 318 (HO).

SOUTH AUSTRALIA: Near Eden Valley, 13 Jan. 1990, R./. Bates 21927 (AD); Southern Lofty, Upper M[ount] Bold Reservoir, 21 Mar. 1993, RJ Bates 31800 (AD, CANB); Hundred of Robertson, ca 10 km S of Naracoorte, 24 Sep. 1962, D. Hunt 1423 (AD); Cox Scrub, 16 Feb. 1990, D.E. Murfet 908 (AD); Shores of Millbrook Reservoir, 11 Mar. 1962, D.E. Symon 2072 (AD, BRI, CANB, NE).

Ic. Stellaria angustifolia subsp. rotundisepala C.H.Mill. & J.G.West, subsp. nov.

A subsp. angustifolia et subsp. tenella (Benth.) C_H. Mill. & J.G.West sepalis rotundatis vel late ellipticis, apice obtuso et plicato cucullato formenti; seminibus tuberculiis angustis rotundatis —prominentibus, distinguenda. Holotypus: New South Wales, At Source of Murray Rfiver], 6 Jan. 1992, RJ. Bates 27162 (AD 99206284). Isotypi: CANB, NSW, MEL, NY. Stellaria angustifolia subsp. Timbarra River (N.A.Wakefield 4853) C.H.Muill. in Austral. Pl. Name Index database (APNI), http://www.anbg.gov.au/cgi-bin/apni [accessed

C.H. Miller & J.G. West J. Adelaide Bot. Gard. 25 (2011)

i ~ oe d al = - oe —_ a 7 — ~ = a —"

i er rei aa ws >

,

\) \ 1 . | Y >

Fig. 3. A-G Stellaria flaccida: A habit; B part of flower showing sepal, petal and stamen arrangement; C outer view of sepals; D petal; E capsule showing position in relation to sepals; F seed, side view; G seed coat cell detail. H-M S. media: H habit; | part of flower showing sepal, petal and stamen arrangement; J petal; K flower showing capsule; L seed, side view; M seed coat cell detail. N-R S. pallida: N habit; O part of flower showing ovary, stamens and style arrangement; P flower showing sepals and mature capsule; Q seed, side view; R seed coat cell detail. Scale bars: habits (A, H,

38

J. Adelaide Bot. Gard. 25 (2011)

6 May 2010] & Austral. Pl. Cens. database (APC), http:// www.anbg.gov.au/cgi-bin/apc [accessed 6 May 2010].

Annual, slender, erect, usually single stemmed, (6—) 11—25 cm long, glabrous. Stem and inflorescence leaves a continuous series, sessile, obovate or linear or ovate or narrower, (3.5—) 6—9.5 (—13) mm long, 0.9—-1.8 mm wide, acute to subacute, rarely obtuse. /nflorescence axillary and solitary or an interrupted monochasium of 2—3 flowers. Pedicels (22—) 27-40 mm long, slender, erect in fruit. Sepals round to broad-elliptic, 1.5—2.1 mm long, obtuse, apex folded over to form a hood, sometimes with hood thickened. Petals 2.4—3 mm long, longer than sepals, bifid. Stamens 10. Staminodes 0. Styles 0.8—1.5 mm long. Capsule ellipsoid to ovoid, (2.9—) 3.44.6 mm long, (1.8—) 2.4-3.1 mm wide, from 2 to 4 times longer than sepals; valves straight, apex straight. Seeds 2—5 (—7), suborbicular, 0.9-1.4 mm long, mid brown to reddish brown to purplish, tubercles very narrow rounded projections totally covering the seed, giving a spiny appearance. Fig. 1B, 2Q—W. Flowering: Jan.

Distribution and habitat. This taxon 1s restricted to high altitudes around the source of the Murray and Tuross Rivers in Kosciuszko National Park in New South Wales and Timbarra River in East Gippsland, Victoria (Fig. 6C).

Found in alpine areas in wet bogs with mixed herbfields.

Notes. Subsp. rotundisepala is distinguishable from the other subspecies by its distinct seed and floral characters. Currently only three specimens are known and further collecting 1s needed in the areas where it occurs to obtain more information about it.

Etymology. Subsp. rotundisepala has been named to reflect the unique shape of the sepals which is not found in any other species in Australia; from the Latin rotundus for round and sepalum for a sepal.

Specimens examined

New SoutH WALtEs: S. Tablelands, Dillundoo Valley, 11 Jan. 1970, J.-A. Willis s.n. (MEL, NSW).

VicTorIA: East Gippsland, Timbarra River, 13 Jan. 1959, N.A. Wakefield 4553 (MEL).

2. Stellaria filiformis (Benth.) Mattt.

Repert. Spec. Nov. Regni Veg. Beith. 100: 148, tab. VII (1938). — Drymaria filiformis Benth., Fl. Austral. 1: 162 (1863). — Type Citation: “W. Australia, Drummond, n. 694.” Lectotype (here designated): Swan River Drummond n. 694 (K, Herb. Hooker p.p,. 2 plants on right hand side of sheet). Isolectotypes: (K, Herb. Hooker; MEL 723012).

Euthales (?) filiformis de Vriese, Pl. Preiss. 1(3): 414 (1845). — Type citation: “In solo sublimoso fertili prope praedium rusticum Dom. Marell, York, d. 30m. Marti 1840. Herb. Preiss. No. 1889.” Holotype: L (Herb. Lugd. Bat. sh. 909.62-546), n.v, fide J.H. Kern, Blumea 13(1): 116 (1965).

Stellaria in Australia (Caryophyllaceae)

Illustration. C.H.Mill. & J.G.West in N.G.Walsh & Entwisle, Fl. Victoria 3: 236, Fig. 45n—o (1996).

Annual with erect to spreading stems to 15 (—22) cm long, glabrous. Leaves forming a basal rosette, sessile, filiform, (4—-) 7-20 (—26.5) mm long, (0.2—) 0.4—1.2) mm wide, subacute, herbaceous to fleshy; inflorescence leaves sessile, linear to ovate to lanceolate, (0.7-) 1—6.4 (—13) mm long, (0.25—) 0.4—-1.1 mm wide, acute, scarious or sometimes herbaceous. /nflorescence a dichasium. Pedicels 2—11 (—16) mm long, wiry, erect in fruit. Sepals 2—4 mm long, acute to obtuse. Petals (0.5—) 1-1.5 (-2) mm long, shorter than half sepal length, deeply bifid. Stamens (2—) 3-5. Staminodes 0-5. Styles 0.2-0.6 mm long. Capsule narrow, ovoid to ellipsoid, (2—) 2.5-4.5 (—5.5) mm long, 1—-1.5 mm wide, equal to twice sepal length; valves straight, rarely spreading, apex straight. Seeds 5—13, suborbicular, 0.5—0.75 mm long, cream, light to mid brown, smooth, with minute conical tubercles. Fig. 1D, 4A—H. Flowering: (June—) Aug.—Nov.

Distribution and habitat. This taxon has a disjunct distribution across southern Australia. There are isolated records in the Wyalong district in New South Wales. In Victoria it is confined to the north-west region around Wyperfeld National Park, south to the Grampians area and then westwards across to the south-east of South Australia as well as the Eyre Peninsula. In Western Australia it occurs from the Fraser Range in the east, Lake King, Cowcowing and west across to the Irwin River (Fig. 6D).

Locally common in mallee’ scrub, _ eucalypt woodlands and heathlands often in sandy soils and well drained shallow sandy loams, sometimes in the swales between dunes.

Common Name. Thread starwort.

Notes. Stellaria filiformis 1s distinct as it is the only species in Australia with a basal rosette of filiform leaves and it lacks prominently tuberculate seeds. Instead the seeds are smooth with minute conical tubercles, usually with pits in the top (Fig. 1D) and while the cell shape is similar to those of the subspecies of S. multiflora, the seeds are much smaller than those of the subspecies of S. multiflora.

Stellaria filiformis shows affinities with S. multiflora as they share similar habitats and reduced petals and stamens. The collections made in New South Wales are all from the 1920s. It is unusual that there are no other known collections, as the habitat in which the taxon occurs in other states is found in the central west to south west of New South Wales.

Nomenclatural Notes.

Drymaria_ filiformis Benth. Two sheets from Herbarium Hookerianum held at K contain type material. One sheet has two collections on it. The collection on

N) 5 mm; flowers & fruits 1 mm; ovary (O) & seeds (F, I, P) 0.5 mm. — A-G B. Copley 5175 (AD 97810081); H-J C.R. Alcock 605 (AD 966220095);

N-R C.R. Alcock 10738a (AD 99021068).

C.H. Miller & J.G. West

the left hand side (LHS) of the pencil line has the label ‘West Australian Goldfields Spencer Moore 1895’ and is not considered part of the type collection. The collection on the right hand side (RHS) of the pencil line has ‘694 Swan river Drummond’ written on the sheet and matches the protologue and has thus been designated here as the lectotype. Also written in pencil on the sheet is a description of Drymaria filiformis which 1s thought to be in Bentham’s handwriting.

A second K sheet has the label ‘Swan R[iver]. Drummond No 694 Pres? By W.W. Sanders, Esq. FLS’. This sheet matches the other type material and thus has been designated here as an isolectotype.

One sheet from MEL, (MEL 723012) has the label ‘W.A. J Dr’ with a tag with the number 694 located on the specimen. As the collecting information and the specimen match the type material this has also been designated here as an isolectotype.

Euthales (?) filiformis de Vriese. This is a legitimate taxon, as Kern (1965) discusses. It 1s the oldest name available for the species but, as the epithet /iliformis is already occupied in the genus Sfe/laria, it cannot be transferred to that genus, and as such is unavailable for use.

Selected specimens (of c. 70 seen)

New SouTtH WALEs: Pine Ridge, Wyalong, Nov. 1920, A. Cooper s.n. (BRI, CANB, NSW); Central W. Slopes, Upper slope of Wamboyne Mountain, W of Lake Cowal, Dec. 1993, A.N. Rodd & A. Clements 9245 (NSW).

VicToRIA: Wyperfeld National Park, junction of track with Dattuck Track, ca 3 mls ENE of Eastern Lookout, | Oct. 1968, A.C. Beauglehole 28770 (CANB, MEL); Sunset Country, E to W track, c. 35 km S of Meringur, 23 Sep. 1981, /.H. Browne 52 (MEL, NSW); Wandown Fauna Reserve, 10 miles ENE of Annuello, 5 Sep. 1970, N. Macfarlane 537 (MEL).

SOUTH AUSTRALIA: Northern Flinders Ranges; Baratta Hill, 11 Sep. 1986, RJ. Bates 7164 (AD); Eyre Peninsula; Gawler Ranges, 24 Sep. 1989, RJ. Bates 21078 (AD); Gammon Ranges ca. 65 km E of Leigh Creek, Near mouth of gorge of Arcoona Creek S of Arcoona Bluff Range, 17 Sep. 1956, H7. Kichler 12685 (AD); Murray; Lowan Conservation Park, | Oct. 1981, A.G. Spooner 7595 (AD).

WESTERN AUSTRALIA: Goldfield Ranges survey, Flat north of range, c. 9km NE of Bungalbin Hill, 28 Sep. 1995, M. Gibson & M. Lyons 3356 p.p. (CANB); Cowcowing, Aug. 1904, M. Koch 1122 (HO, PERTH); Central South, Eyre Hwy ca 23 miles (37km) E of Fraser Range, 6 Sep. 1963, J.-H. Willis s.n. (MEL); 23 km S of Lake Grace township, 8 Sep. 1967, P.G. Wilson 6225 (PERTH); 26 km W of Warriedar HS, 26 Sep. 1986, PG. Wilson 12267a (PERTH).

3. Stellaria flaccida Hook.

Companion Bot. Mag. 1(9): 275 (1836). — S. media var. flaccida (Hook.) Hook.f., Bot. Antarct. Voy. III. (FI. Tasman.) 1: 43 (1855). — Type Citation: ‘Mr. Gunn (n. 450.)’. Lectotype (here designated): Common near Launceston amongst rocks where shaded, Gunn 450 (K, Herb. Hooker p.p. [LHS of the pencil line], Kew Loan Number 0589-86-5). Isolectotype: 450 very common in dense shady thickets growing under logs, etc., [Gunn] (K, Herb. Hooker p.p. [LHS of the pencil line], Kew Loan Number 0589-86-7).

40

J. Adelaide Bot. Gard. 25 (2011)

Stellaria flaccida f Hook., J. Bot. (Hooker) 2: 411 (1840), nom. inval.

Illustration. C.H.Mill. & J.G.West in N.G.Walsh & Entwisle, Fl. Victoria 3: 236, Fig. 45j-k (1996).

Perennial, stems lax to 50 cm long, which may root at nodes, with creeping rhizome, almost glabrous to sparsely hairy. Stem and inflorescence leaves a continuous series, subsessile to appearing petiolate; narrow ovate, elliptic, broad ovate or obovate, (5-—) 7—20 (—23) mm long, (1—) 2—10 (—17) mm wide, acute, margin entire or undulate with hairs few to almost ciliate, flaccid. Inflorescence axillary and solitary to a monochasium of up to 6 (—10) flowers. Pedicels (13—) 20—73 (—80) mm long, erect to pendant in fruit. Sepals 3.5-6.5 (—7.5) mm long, margins glabrous to ciliate. Petals (3.7—) 4—7.5 (—8.2) mm long, equal to or longer than sepals, deeply bifid. Stamens 10. Staminodes 0. Styles 2.2—3.7 mm long. Capsule usually ovoid, 5.2—7.7 mm long, (2.9—) 3.4—-5 (—5.5) mm wide, equal to twice sepal length; valves spreading, rarely straight, apex recurved or twisted sideways, rarely straight. Seeds 4—14, suborbicular to obloid, 1.3—2.2 (—2.6) mm long, light grey-brown to almost black, tubercles rounded to shortly ridged, inflated. Fig. 1H, 3A—G. Flowering: Sep.—Mar.

Distribution and habitat. This taxon occurs on the tablelands and coastal plains of south-eastern Australia from southern Queensland to Tasmania, excluding the Bass Strait Islands. It also occurs westward from the Otway Ranges, and Dandenongs in Victoria as well as in the Southern Lofty Ranges and on Kangaroo Island in South Australia (Fig. 6E).

The species grows in damp sites associated with rivers and creeks in fern communities, rainforests, and wet sclerophyll to tall open eucalypt forests, often with erassy understorey; often found in disturbed areas such as along roadsides and tracks. It occurs on a range of soil types from sands to silts and clays and on granite, basalt, limestone and sandstone.

Notes. The indumentum of the sepal margins varies considerably with the most common state consisting of two sepals with glabrous margins, one sepal with one margin ciliate and one margin glabrous, and two sepals with both margins ciliate. The degree of hairiness of the margins varies from almost glabrous with a few hairs on the base of the margins to densely ciliate along the entire margin; almost every state between the two has been observed. This pattern of distribution of the hairs 1s also found in the introduced taxon S. graminea. Specimens on the edges of the distribution range, particularly in Queensland and South Australia, tend toward S. angustifolia in being more glabrous and having leaves closer to linear.

Nomenclatural Notes. Stellaria flaccida Hook. Two sheets from Herbarium Hookerianum held at K contain type material. The

J. Adelaide Bot. Gard. 25 (2011)

first sheet has two collections, the first on the LHS of the pencil line has a label with ’[Gunn] 450 Stellaria flaccid Common near Launceston, amongst rocks where shaded’ and hand written on the sheet is ‘No. 450. Mr. Gunn Van D’s Land St. flaccida Hook’ in what is thought to be Hooker’s handwriting. As this information and the specimen match the protologue it is here designated as the lectotype. On the RHS of the sheet is another collection from Kiama NSW and it is not type material.

The second sheet has two collections on it. The first label attached to the specimen on the LHS of the pencil line, has *450 Stellaria flaccida Hook Very common in dense shady thickets growing under logs, etc’ on it. This information matches the protologue and thus is here designated as the isolectotype. The second label has ‘450/1842 V D Land Gunn Circular Head 4/11/37’ written on it, which post-dates publication and is thus not considered to be type material. This collection also has a handwritten note in Hooker’s hand; ‘Stellaria flaccida B 450 Gunn V D land’. This name was invalidly published by Hooker (1840).

Selected specimens (of c. 235 seen)

QUEENSLAND: Mount Merino, Beereenbano Lookout, 2 Dec. 1970, L.R. Telford 2625 (BRI, CANB).

New SoutH WALES: 12 miles S of Nowedoc, Pigeon Top and track, 7 Nov. 1972, J. Carrick 3287 (AD); SE Coast, Mt Dromedary near central Tilba, which is ca 190 km N of Vic border, 26 Apr. 1973, R.J. Chinnock 266 (AD); NT, Eastern side of current Barrington trail c. 100 m S of TO to Little Murray camping area, Barrington Tops National Park, 9 Apr. 2003, JR. Hosking & B.J. Neilly 2301 (CANB).

Victoria: On track to summit of Mt Ellery, between Big River Track and summit, 22 Feb. 1984, D.E. Albrecht & B.J. Conn 211 (BRI, MEL); Near Delegate [Mount], 8 Jan. 1992, R.J. Bates 27246 (AD); D26 Grampians, Victoria Range, Victoria Range Track (W of Sawmill Track), on top of range, 14 Jan. 1969, A.C. Beauglehole 30262 (MEL); Gippsland region, Wilson’s Promontory National Park. Along road from carpark to Mt Oberon, 6 Dec. 1975, Hj. Eichler 21793 (CANB).

TASMANIA: Four Mile Stream, 6 km S of Falmouth, 23 Nov. 1974, R.J. Chinnock 2180 (AD); Ferntree, Mt Wellington, | Jan. 1972, J. H. Hemsley 6658 (HO, NSW); Midlands, Kubla Khan Cave State Reserve ca 10 km W of Mole Creek township, 16 May 1983, A. Moscal 2392 (HO); Central Highlands, East slope of Archers Sugarloaf, Feb. 1986, A. Moscal 12501 (HO); Ben Lomond Tower Hill, East Tower, Feb. 1980, 1G. Noble 29078 (HO).

SOUTH AUSTRALIA: Kangaroo Island, Ravine des Casoars, s.dat., R./J. Bates 30356 (AD, CANB); Ravine des Casoars, s.dat., R.J. Bates 30425 (AD, CANB); Waterfall Gully, near Adelaide, 16 Nov. 1887, /.H. Maiden s.n. (NSW).

4. * Stellaria graminea L.

Sp. Pl. 1: 422 (1753). — Type Citation: ‘Habitat in siccis juniperetis sepibus Europea’. Lectotype: Herb. Burser XI: 111 (UPS n.v.; LINN 584, photo), fide Jonsell & Jarvis in Nordic J. Bot. 14: 159 (1994).

Illustration. C.H.Mill. & J.G.West in N.G.Walsh & Entwisle, Fl. Victoria 3: 236, Fig. 45g (1996).

Perennial, usually erect single or several stemmed to 60 (—89) cm long, sometimes with short vegetative side

4]

Stellaria in Australia (Caryophyllaceae)

shoots, with a slender rhizome and frequently rooting at nodes, glabrous except for hairs on margins of leaves and some sepals. Stem leaves sessile, narrow ovate, linear or obovate, (7.3—) 10—30 (—35) mm long, (1.6—) 2—5 (—5.3) mm wide, acute to subacute, herbaceous; inflorescence leaves sessile, ovate to broadly ovate, triangular, (1.8—) 2.5-6 (-9.2) mm long, (0.6—) 1-2 (—2.6) mm wide, scarious, rarely green. /nflorescence usually a dichasium. Pedicels (9—) 15-50 (—75) mm long, slender, erect in fruit, usually ridged. Sepals (2.3—) 3-6 (—6.7) mm long, margins glabrous to ciliate, acute. Petals (1.5—) 2.2—5.2 (—5.9) mm long, shorter than to longer than sepals, very deeply bifid. Stamens 10, often innermost whorl of 5 reduced but functional. Staminodes 0. Styles (1—) 1.5—3.5 (4) mm long. Capsule ovoid to ellipsoid, 3.3-4.8 mm long, 1.8—2.8 mm wide, equal to or longer than sepals; valves straight, apex straight or sometimes upper margin recurved. Seeds 6—10 (—26), suborbicular to ellipsoid, (0.5—) 0.8—1.2 mm long, reddish brown to ruby red, tubercles narrow ridges, semi-inflated. Fig. 1E, 5A—G. Flowering: Nov.—Feb.

Distribution and habitat. This species has been collected only a few times from five localised areas in Australia. It appears to have been introduced into Tasmania first in the early 1900s at Tyenna and Junee River. The last collection on the main island was at Junee River in 1942. One specimen has been collected from King Island in 1979. In Victoria it has only been collected once from the Cann River in 1946. In New South Wales several collections dating from 1951 until 2007 have been made in the Kosciuszko region. In South Australia One specimen has been collected from Parawa in 1984 (Fig. 6F). The native range is thought to be most of Europe but the species has also spread and is now found throughout Europe and North America and is considered to be a weed (Chater & Heywood 1993).

This is a rare weed that is found in very damp areas on the edges of rivers, swamps, lakes in grassland or Leptospermum scrub. It grows in soils that retain moisture such as black mud or granites.

Notes. Stellaria graminea can be distinguished from the endemic S. angustifolia by its dichasial inflorescence that has differentiated floral leaves and by its distinct pattern of hairs on the sepal margins. This pattern of hairs is also found in the Australian species S. flaccida. This character is also useful to distinguish S. graminea from the European species S. palustris. S. graminea can be distinguished from S. flaccida by its dichasial inflorescence that has dimorphic floral leaves.

Specimens seen

NEw SoutH WALtEs: S. Tablelands, [Kosciuskzo], Diggers Ck, 25 Jan. 1951, L.4.S. Johnson & E.F Constable s-.n. (NSW); Southern Tablelands, Kosciuszko National Park, on track from Charlottes Pass to Blue Lake on right hand side in drainage line on last hillside before, 22 Feb. 2007, J. McAuliffe et al. 732 (CANB); S. Tablelands, [Kosciuskzo], White River area, 24 Jan. 1967, TY Stead s.n. (NSW); S. Tablelands, [Kosciuskzo], Diggers Ck Lake, National Park, 2 Feb. 1978, J. Thompson 2777 (NSW).

C.H. Miller & J.G. West J. Adelaide Bot. Gard. 25 (2011)

~~

.e oe ed PAP Tne 2A Sak ST eit. - a ' —Soos

a” - a", . J + es >

ri aa Wy \ 1a Nu Alem \\ CUNY a | My

2 ih 2 UY aay P., Wi)

La (Sag

ye

ee ° a t owe > * _~ = : i

2 Sore te ser Ht (Thaet tyne <4

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Fig. 4. A-H Stellaria filiformis: A habit; B flower; C part of flower showing sepal, petal and stamen arrangement; D petal; E flower showing mature capsule; F flower with opened capsule; G seed, side view; H seed coat cell detail. -O S. multiflora subsp. multiflora: | habit; J flower; K part of flower showing ovary, stamen and style arrangement; L flower showing mature capsule; M flower with immature capsule; N seed, side view; O seed coat cell detail. P-V S. multiflora subsp. collaris: P habit; Q flower; R part of flower showing sepal, petal and stamen arrangement;

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J. Adelaide Bot. Gard. 25 (2011)

VicTorIA: East Gippsland, Cann River Grid Z31, 1946, N.A. Wakefield 3691 (MEL).

TASMANIA: Junee River, 19 Dec. 1942, H. & J. Gordon s.n. (HO); Waratah, Dec. 1924, 4.H.S. Lucas s.n. (NSW); King Island, Sea Elephant River, 9 Jan. 1979, DI. Morris 7964 (HO); East Coast Tyenna, Nov. 1903, L. Rodway s.n. (HO).

SOUTH AUSTRALIA: 2 km W of Parawa, 28 Jan. 1984, R./. Bates 3652 (AD).

5. Stellaria leptoclada (Benth.) C.H.Miull. & J.G.West, comb. nov. Stellaria glauca var. (?) leptoclada Benth., Fl. Austral. 1: 158 (1836). — Type citation: “New England, C. Stuart’. Holotype: New England, C. Stuart (K, Herb. Hooker). Stellaria sp. leptoclada (B.G.Briggs 2194) C.H.Mill. In: Austral. Pl. Name Index database (APNI), http://www. anbg.gov.au/cgi-bin/apni [accessed 6 May 2010] & Austral. Pl. Cens. database (APC), http://www.anbg. gov. au/cgi-bin/ape [accessed 6 May 2010].

Annual, fine, small, weak to erect, single or several stemmed to 20 (—30) cm long, sometimes arising from (loose) basal rosette, glabrous, scabrous or with hairs sparsely scattered on stems. Stem and inflorescence leaves a continuous series, often attenuate, appearing petiolate; narrowly obovate, linear to narrowly ovate, (2—) 3-17 (—20) mm long, (0.6—) 0.8—2 (—3.4) mm wide, acute to subacute, entire, margin scabrous, toothed or with short hairs on lower parts, rarely scabrous on midrib. /nflorescence a monochasium. Pedicels (9—) 14— 34 (-45) mm long, slender, erect, often reflexed in fruit, glabrous or slightly scabrous. Sepals (3—) 3.5—5.2 (—5.6) mm long, acute to acuminate. Petals (3—) 3.4-5 mm long, shorter than to slightly longer than sepals; deeply bifid. Stamens 10. Staminodes 0. Styles (1—) 1.5—2.2 mm long. Capsule narrowly ovoid to ellipsoid, rarely broader, 3.2-4.8 (—5.5) mm long, 1.5—2.4 mm wide, usually equal to slightly longer than sepals, rarely shorter than sepals, valves spreading or straight, apex straight or recurved. Seeds (5—) 10-14 (—22), suborbicular, rarely broadly reniform, (0.55—) 0.6—0.85 (—1.1) mm long, light to mid or reddish-brown, tubercles narrow ridges, semi-inflated. Fig. 1F,2X—DD. Flowering: Aug.—Jan.

Distribution and habitat. There is a single specimen known from south-east Queensland in the Silverwood area. The remaining specimens have been collected from New South Wales mostly from the Great Dividing Range from Boggabri in the north to Gloucester in the south (Fig. 6G).

Scattered to locally common in moist sites in undisturbed grassland under open woodlands or forests. It grows on granites or soils derived from granite, or rarely on serpentine or basaltic soils.

Notes. S. leptoclada 1s a relatively rare herb that has frequently been overlooked. S. leptoclada can be

Stellaria in Australia (Caryophyllaceae)

distinguished from S. angustifolia using habit, leaf shape, indumentum and floral characters. It differs from S. graminea in its habit, leaf shape and lack of differentiated floral leaves. It also has a much more restricted geographic distribution. See also notes under S. angustifolia.

Selected specimens (of c. 40 seen)

QUEENSLAND: Silverwood, Sep. 1922, C.7) White 1713 (BRI).

New SouTH WALES: Near Cobar, Sep. 1910, L. Abrahams s.n. (NSW); Watchimbark Creek, NW of Gloucester, 21 Sep. 1968, D.F! Blaxell & R.G. Coveny 6 (NSW); Graman, Jan. 1959, TV. Bourke s.n. (NSW); Coulson’s Creek, foot of Liverpool Range, N of Merriwa, 28 Sep. 1968, B.G. Briggs 2194 (NSW); Warialda, Aug. 1933, F) Browne s.n. (NSW); North Western Slopes, Warrabah National Park, semi-cleared area, 2.5 km E of camping area, 5 Oct. 2001, L.Z Copeland 3198 (CANB); NW Slopes, Ashford Road via Bonshaw Road, 17 Oct. 1993, R.G. Coveny & A.J. Whalen 16649 (CANB); Attunga State Forest, 27 Sep. 1985, J.R. Hosking 323 (CANB, NE); North-west Slopes, eastern side of Woodsreef mine, 27 Aug. 1992, J.R. Hosking 516 (CANB, MEL, NE); [Dumaresq] dam, above walking track, 18 Oct. 1990, S. McIntyre s.n. (NSW); Wallangra, 28 Sep. 1929, Ex Herb Rodway 603 (NSW); Glenn Innes, Mar. 1913, H. Wenholz s.n. (NSW).

6. * Stellaria media (L.) Vill.

Hist. Pl. Dauphine (Villars) 3(1): 615. (1789). — Alsine media L., Sp. Pl. 1: 272 (1753). — Type citation: ‘Habitat in Europea cultis*. Lectotype: Herb. Linn. No. 388.1 (LINN, Photo), fide Turrill in Turrill & Muilne-Redhead, Fl. Trop. E. Africa, Caryophyllaceae: 24 (1956); image also available at http://www.nhm.ac.uk/jdsml/research- curation/research/projects/linnaean-typification

Stellaria media (L.) Cirillo, Essent. Pl. Char. Comment. 36

(1784), nom. inval.

Illustration. C.H.Mill. & J.G.West in N.G.Walsh & Entwisle, Fl. Victoria 3: 236, Fig. 45a—b (1996).

Annual, semi-erect to erect herb to 40 (—56) cm long with long straggly branches, may root from nodes, with single line of hairs down internodes and pedicels and hairs on the back of sepals and leaf margins. Stem and inflorescence leaves a continuous series, the lower leaves are often petiolate, narrowed, very attenuate, 1— 15 (—18) mm long, upper part broad ovate to ovate to elliptic, rarely linear, (3.5—) 6-18 (—24.5) mm long, (1—-) 2-10 (-14) mm wide, acute, often undulate. Inflorescence a condensed, leafy dichasium. Pedicels (2.1—) 4-16 (—22) mm long, erect in fruit. Sepals 3—5.5 (—6) mm long, obtuse, often forming small hood, often with purple mark, outer sepal backs sparsely to densely hairy, frequently glandular, rarely glabrous. Petals (1.6—) 2.5—3.6 mm long, half to almost sepal length, deeply bifid. Stamens 3—5. Staminodes QO, rarely 1-3. Styles 0.5—0.8 (—1) mm long. Capsule ovoid to ellipsoid, (3—) 4-6 (-6.5) mm long, 2.2— 3.5 (—3.8) mm wide, just

S petal; T flower showing mature capsule and thickened collar on pedicel; U seed, side view; V seed coat cell detail. W-BB S. multiflora subsp. nebulosa: W habit; X flower; Y part of flower showing sepals, ovary, stamen and style arrangement; Z flower showing mature capsule; AA seed, side view; BB seed coat cell detail. Scale bars: habits (A, I, P, W) 5 mm; flowers & fruits 1 mm; petals (D, $), ovary (K) & seeds (G, N, U, AA) 0.5 mm. — A-H R.J. Chinnock 7348 (CANB 00523485); IEN R.J. Bates 30390 (AD 99328276); O-U Hy. Eichler 12833 (AD 95727048); V-AA R.J.

Bates 25994 (AD 99146306).

C.H. Miller & J.G. West

shorter than to longer than sepals; valves spreading, apex recurved or straight. Seeds (5—) 7—15, discoid to flattened-ellipsoid, (0.8—) 0.9—-1.2 (—1.4) mm long, red to black to dark purplish-brown, tubercles rounded hills, semi-inflated, cell walls with minute papillae. Fig. 1G, 3H-—M. Flowering: all year, but predominantly July—Dec.

Distribution and habitat. Occurs in the North from Bundaberg, Queensland along the east coast and Great Dividing Range through New South Wales to the Victorian coast and mountains, along the Murray floodplain and west to South Australia from Adelaide and surrounding ranges as far west as Port Lincoln. Also occurs in eastern Tasmania and the Bass Strait Islands. There are a few records for the Northern Territory from Alice Springs and in Western Australia from coastal areas from Albany to just north of Perth. Also a single specimen is known from Kununurra in northern Western Australia. There are also scattered occurences outside these areas in gardens (Fig. 6H). The native range is thought to be throughout Europe except in the extreme north (Chater & Heywood 1993), but it is currently found on every continent, absent only from the polar regions and very dry areas. It is an important economic weed of crops.

Stellaria media 1s a common introduced weed, especially in areas of disturbance such as roadsides, waterways and areas of cultivation. Usually found in shady or moist places in coastal dunes, grass or herbfields, gully forests, open forests, heathlands, scrubs or swamps. Grows on most substrates from rocky soils to sands.

Common name. Chickweed, Common Chickweed.

Notes. S. media is frequently confused with S. pallida. From close examination of Australian material and limited overseas material it has become apparent that there are problems in the delimitation of these two taxa. The main character that distinguishes them is the presence or absence of petals. In the Australian specimens of S. pallida petals are always absent, and the character state of reduced or minute petals does not occur. However, non-Australian literature indicates that this is not always the case overseas (Clapham et al. 1952; Morton 1972; Rabeler 1988; Chater & Heywood 1993; Morton 2005).

In Australian specimens of S. media the petals are always present and easily visible. In occasional specimens the petals may be very short but they are always bifid. However, there are several other characters where there is a slight overlap between the two taxa. The most useful character to distinguish the two species is the number of stamens. S. media usually has 3 — 5 stamens present or very rarely only two. S. pallida has 2 stamens or very rarely 3 or 4. Seed colour and size can also be useful. In less than 10% of S. pallida specimens seeds measure more than 0.8 mm in diameter. S. pallida has either pale seeds only or a mixture of dark and light seeds in one capsule. The seeds in S. media are usually

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J. Adelaide Bot. Gard. 25 (2011)

1 mm or larger in diameter, rarely less. The colour of S: media seed is brown to black or occasionally medium brown but they are never pale when mature. S. pallida has shorter stigmas (0.5 mm or shorter) and these tend to be spreading and reflexed. By contrast, S. media has longer stigmas (0.5 mm or longer) and these are mostly erect with only the apex revolute.

Characters traditionally used in keys to separate these taxa such as sepal length, the presence of glandular hairs on backs of sepals and the attachment of the upper leaves show too much variation and overlap to be useful.

In non-Australian literature these two taxa have usually been treated separately, particularly in recent years (Chater & Heywood 1993; Chen & Rabeler 2001; Morton 2005). This is also the case in the Australian literature (Curtis 1975; Chorney 1986; Doust 1990; Miller & West 1996). Chromosome numbers also support this separation with S. media having 2n=40, 42, 44 and S. pallida having 2n=22 (Chen & Rabeler 2001; Morton 2005).

Nomenclatural Notes. In some Australian literature the authorship of S. media has been given as “(L.) Cirillo”. However Cirillo (1784) did not validly publish the combination, as he did not definitely associate the generic name and the specific epithet.

Selected specimens (of c. 295 seen)

QUEENSLAND: Leslie Dam, 10km W of Warwick, 3 Oct. 2000, G.N. Batianoff & C. Appelman 2010375 (CANB); Athol Hall, near Westbrook, about 11 km SW of Toowoomba, 05 Sep. 1947, S.L. Everist 3156 (BRI); Jolly’s Falls, c. 2.5km WNW of the Summit railway siding, 12 Sep. 2003, B.L. Lepschi et al. 4955 (CANB); In grounds of CSIRO Long Pocket Laboratories, Indooroopilly, Brisbane, 19 July 1973, V.K. Moriarity 1330 (CANB).

New SouTtH WALEs: S. Tablelands, Bendethera Caves, 20 miles [32 km] c. W of Moruya, 10 May 1966, E.F’ Constable 6863 (NSW); Murrumbidgee River, in Euroley State Forest (1 km N of entrance), 20 Sep. 1992, C_\H. Miller & J. Palmer 590 (CANB); Broken Hull, 24 Oct. 1927, A. Morris s.n. (AD); North Coast Clouds Creek, via South Grafton, | Nov. 1949, HM. Provisional School 31 (NSW); C. Tablelands, Coxs R, 6 miles [9.6 km] W of Little Hartley, 30 Aug. 1970, J. Thompson 628 (NSW).

AUSTRALIAN CAPITAL TERRITORY: S. Tablelands, Turner, Canberra, 5 Sep. 1964, H.S. McKee 11590 (NSW)

VicToRIA: Gippsland region, Raymond Island, SW corner, 14 km SE of Bairnsdale, 27 Sep. 1992, I. Crawford 1885 (CANB, MEL, NSW); Just inside entrance of Johnson Swamp Game Reserve, 4.6 km off Kerang to Leitchville Road, 24 Sep. 1992, C_H. Miller & J. Palmer 601 (CANB); Port Phillip Bay, Mud Island, Oct. 1983, J. Yogovic s.n. (MEL).

TASMANIA: King Island, Councillor Island, east coast of King Island, 8 July 1966, W. Bartlett s.n. (HO); Central Highlands, Wild Dog Tier, Mar. 1984, 4A. Moscal 6787 (HO); Mt. Field, beside Sitzmark Lodge, Mt. Field National Park, 6 Jan. 1978, MB. Smith 320 (HO); Lenah Valley, 2 Aug. 1963, P.A. Tyler s.n. (HO); East Coast East of Bicheno township, on seaward side, Jan. 1983, JG. West 4533 (HO); Furneaux Group, Mount Chappell Island, 12 Aug. 1973, JS. Whinray 1141 (MEL); Kents group, Deal Island, Lighthouse Gully, 29 Nov. 1970, JS. Whinray 1251 (CANB).

J. Adelaide Bot. Gard. 25 (2011)

NorTHERN TERRITORY: AZRI [Arid Zone Research Institute, Alice Springs], 18 Oct. 1985, PK. Latz 10260 (DNA); Alice Springs, 19 Sep. 1977, A.S. Mitchell 419 (AD, DNA).

SOUTH AUSTRALIA: Southern Eyre Peninsula, 12 Ellen St, Port Lincoln, 19 Aug. 1965, C.R. Alcock 605 (AD, NSW); Southern Lofty, Onkaparinga Gorge, Oct. 1991, R.W/. Bates 25970 (AD, CANB); South-eastern, 71 Crouch St, Mt Gambier, 11 Oct. 1969, B. Copley 2826 (AD, NSW); Koonamore Station, ca. 400 km NNE of Adelaide, near Bindyi, Mustering Pdk no. 3, 7 Sep. 1973, M.D. Crisp 547 (CANB); Nixon-Skinner Conservation Park, Hundred of Myponga, Section 245, 25 Sep. 1974, T.-M. Heddle & R.B. Heddle WSK27 (AD); Bool Lagoon, ca 25 km S of Naracoorte, 18 Sep. 1961, D. Hunt 436 (AD); Northern Flinders Range, Leigh Creek township, (ca 280 km NNE of Pt Augusta), East Park, 27 Sep. 1971, T.:R.N. Lothian 5162 (AD); Southern Flinders Range, Mambray Creek (Mt Remarkable National Park), ca 40 km SSE of Port Augusta, Lower Alligator Creek area, 7 July 1974, D.J.E Whibley 4379 (AD).

WESTERN AUSTRALIA: Lake William, West Cape Howe, 30 km W of Albany, 10 Sep. 1987, G./. Keighery & J.J. Alford 1819 (PERTH); Oria Orchards, Packsaddle Plain, 8.3 km from Kununurra PO bearing 218 degrees, 2 Aug. 2002, A.S. Mitchell 7341 (CANB); Pig Saleyard, Mt Barker, 1 Sep. 1979, G. Perry 948 (PERTH).

7. Stellaria multiflora Hook.

Companion Bot. Mag. 1: 275 (1836). — Type Citation: ‘Mr. Gunn (n. 451)’. Lectotype (here designated): V D’s Land, Mr Gunn No. 451 |maybe written in Hookers hand] (K, herb. Hooker p.p., two plants in centre of sheet inside pencil line box).

Stellaria sp. Cooyar (A.R.Bean 10622) Qld Herbarium, Austral. Pl. Name Index database (APNI), http://www. anbg.gov.au/cgi-bin/apni [accessed 6 May 2010] & Austral. Pl. Cens. database (APC), http://www.anbg. gov. au/cgi-bin/apce [accessed 6 May 2010].

Stellaria glauca var. tenella auct. non Benth.: Benth., FI. Austral. 1:158 (1863), p.p., only with respect to Robert Brown’s and Gunn’s Tasmanian collections.

Annual with slender taproot, sometimes rooting at nodes; glabrous or with sparse hairs on stems and leat margins; stems prostrate and mat-forming or spreading to erect, to 25 (-45) cm long. Stem and inflorescence leaves in a continuous series, sessile to appearing petiolate; obovate to elliptic to ovate to lanceolate or linear, (1.5—) 2.5—11 (—17.5) mm long, (0.3—) 0.7—2.5 (—3.5) mm wide, acute to obtuse, glabrous or with hairs sparse on lower half of margin. /nflorescence of (2—) 3-15 (—27) flowers, interrupted monchasium to a well-developed monochasium. Pedicels 0.5—20 (—26) mm long, erect in fruit, quadrangular, smooth, collar beneath fruit present or absent. Sepals 2—5.5 (—6) mm long, acute to acuminate, straight or bent. Petals 0 or 2—5, rarely 6, (0.2—) 0.4—1 (—1.2) mm long, shorter than half sepal length, shortly bifid or entire. Stamens (2-) 3—10. Staminodes 0-5. Styles 0.2—0.8 mm long. Capsule narrow to broad, ovoid to ellipsoid, (2—) 2.5—5.5 (—6) mm long, 1—2.5 (—3.5) mm wide, from 0.75 to 1.5 times sepal length; va/ves spreading or straight, apex straight, recurved to revolute in upper half. Seeds (3-—) 5-21 (—25), suborbicular to broadly ellipsoid to reniform, 0.5—-1 (—1.3) mm long, yellow, light to mid brown,

45

Stellaria in Australia (Caryophyllaceae)

sometimes reddish, tubercles rounded to narrow ridges, often inflated. Flowering (Jul.—) Aug.—Dec. (—Feb.).

Distribution. A widespread endemic species in temperate Australia, from south-eastern Queensland, eastern New South Wales, Victoria, southern South Australia, south- western Western Australia, Tasmania and the Bass Strait Islands.

Notes. S. multiflora 1s a taxon of considerable variability and requires further study. We have decided to recognise some of the more obvious variation at subspecies level as there 1s sufficient evidence to warrant this but more work needs to be done.

The members of the S. multiflora can be distinguished by the reduction of the floral structures, particularly the petals and stamens. These characters are variable, which is problematic for the identification of these taxa. However by using a combination of characters, or indeed a shared lack of characters, the following taxa can be determined. It is interesting that one of the key characters for delimiting the genus Stel/aria, the presence of bifid petals is not always observed in this complex. The petals when present are usually minute, less than 1 mm long and can be reduced to fragments. The stamens are also reduced, usually less than 1 mm long and staminodes are also sometimes present.

The specimen A.R. Bean 10622 (Stellaria sp. Cooyar) has been examined and belongs to S. multiflora. The specimen 1s very immature and the floral characters place the specimen in either subsp. multiflora or subsp. nebulosa. Mature fruit 1s required to place it definitively.

Nomenclatural Notes.

Stellaria multiflora Hook. One sheet from Herbarium Hookerianum held at K contains type material. The first label attached to the top LHS of the sheet with one plant is *451/1842 Launceston 15/10/41’. As the collecting date post-dates publication, it is not considered to be type material. The second label in the middle of the sheet, defined by a pencil line, has two plants with the following written on the sheet; ‘No. 451 Stell. multiflora Hook. V. Ds L and Mr Gunn Petals 0’. Gunn 451 matches the protologue and thus is here designated as the lectotype. The third label at the bottom of the sheet associated with several plants has * 1077 V. D[iemens| Land JDH][ooker]’ written on it and as this does not match the protologue, it is not considered to be type material.

Key to subspecies of 8. multiflora

1. Mature fruit broadly ellipsoid, usually > 2 mm wide; valve

apex strongly revolute. ........... a. subsp. multiflora

1: Mature fruit narrowly ovoid to narrowly ellipsoid, < 2

mm wide; valve apex straight or slightly recurved, never revolute

2. Fruiting pedicels with distinct collar, pedicel at collar >

0.5 mm wide; petals 3—5 (—6), minute but usually bifid.

b. subsp. collaris

2: Fruiting pedicels lacking collar, pedicel c. 0.5 mm wide

or less; petals absent. ............ c. subsp. nebulosa

C.H. Miller & J.G. West J. Adelaide Bot. Gard. 25 (2011)

= \ —

= ek >

—

~ r- a=

ap | ot | } Y del: das rlorsk 4

Fig. 9. A-G Stellaria graminea: A habit; B cross section of flower showing sepal, petal and stamen arrangement; C part of flower showing detail of sepal, petal and stamen; D petal; E capsule showing position in relation to sepals; F seed, side view; G seed coat cell detail. H-O S. papillata: H habit; | part of flower showing sepal, petal and stamen arrangement; J-K petal; L flower showing capsule; M mature capsule showing detail of surface papillae; N seed, side view; O seed coat cell detail. P-U S. pungens: P habit; Q part of flower showing sepals, petals and stamen arrangement; R petal:

46

J. Adelaide Bot. Gard. 25 (2011)

7a. Stellaria multiflora Hook. subsp. multiflora

Illustration. C.H.Mill. & J.G.West in N.G.Walsh & Entwisle, Fl. Victoria 3: 236, Fig. 45r—t (1996).

Annual, often mat-forming or erect to 16 (—36) cm long, sometimes rooting at nodes, glabrous. Stem and inflorescence leaves in a continuous series, sessile to appearing petiolate; obovate, elliptic, ovate or linear, (3—) 4—9.8 (—11.5) mm long, (0.6—) 1—2.5 (—3.5) mm wide, acute to subacute. Inflorescence amonochasium. Pedicels 0.5—10 (—20) mm long, collar absent. Sepals (2—) 2.5—4.5 (—5) mm long. Petals 0, sometimes 2-4, (0.2—) 0.5—0.8 (—1) mm long, shortly bifid or reduced to a single arm. Stamens 5—10. Staminodes usually 0, sometimes 1-4. Styles 0.3—-0.6 mm long. Capsule broad ellipsoid, (2—) 2.5—5 (—5.5) mm long, (1.5—) 2—3 (—3.5) mm wide, 0.75 to 1.25 times sepal length; valves straight or spreading, apex strongly revolute. Seeds (5—) 7—21, suborbicular to broadly ellipsoid, 0.5—1 (—1.3) mm long, yellowish, light to mid to reddish brown, tubercles rounded ridges, inflated. Fig. 1L, 4I—O. Flowering: Aug.—Nov.

Distribution and habitat. Subsp. multiflora is widespread but uncommon. There is one record from south-eastern Queensland from the Bunya Mountains collected in 1919, but the majority of collections are found in south- eastern Australia from the southern tablelands and alpine areas of New South Wales, eastern Victoria and central to eastern Tasmania. There are a few scattered collections in south-central and south-western Victoria. It also occurs in the Mt Lofty Ranges and on Kangaroo Island in southern South Australia. There are also a few collections from south-west Western Australia, one collected in 1867 from Porongurup, one collected in 1986 from Hamersley Inlet and a couple of early collections by J. Drummond with a locality of Swan River (Fig. 61).

It grows in various habitats from damp eucalypt forests to woodlands and grasslands, alpine to subalpine areas or rarely found in coastal situations. Grows in rocky sites or shallow soils overlying granite or dolerite.

Selected specimens (of c. 110 seen)

QUEENSLAND: Bunya [Mountains] RAOU Exn, Oct. 1919, C.T. White s.n. (BRI).

NEw SouTH WALEs: South Coast, 2 km N of Nungatta (47 km SW of Eden), 14 Oct. 1974, R.G. Coveny & J. Armstrong 5777 (NSW); Southern Tablelands, ‘Royalla’ S boundary, E of Jerrabomberra Creek, 3.1 km ENE of Lobb Hill, 11 Dec. 1998, I. Crawford 5074 (CANB); Tumbarumba, The Glen, 7 Nov. 1949, E.J. McBarron 3956 (NSW); S. Tablelands, Myanba Gorge Lookout, 25 km E of Bombala, Coolangubura National Park, 10 Sep. 1998, J. Miles s.n. (NSW); Creek W of the Perisher, 8 Apr. 1979, J. Thompson 3091 (NSW); 1 mile E of Gaerlock Homestead, Countegany area, 4 Aug. 1969, T. & J. Whaite 3251A (NSW).

AUSTRALIAN CAPITAL TERRITORY: Mt. Gingera, Cotter River District, 29 Nov. 1966, L.G. Adams 1644 (CANB, NSW); Upper Cotter Valley, 24 Feb. 1959, N.7: Burbidge 6361

Stellaria in Australia (Caryophyllaceae)

(CANB); Kowen area near NSW/ACT border, 30 Sep. 1959, N.T. Burbidge & M. Gray 6549 (CANB); Western base of Black Mountain, 5 Sep. 1964, H.S. McKee 11587 (CANB, NSW).

VictoriA: Melbourne study area, Brisbane Ranges National Park, W of Nelson Lookout, 1.5 km E of Switchback Road, 3 km N of Anakie, 1 Oct. 1977, A.C. Beauglehole & E.G. Errey 56717 (MEL); Glenmaggie Regional Park, Gippsland Lakes Hinterland Study area, 21 Oct. 1984, 4.C. Beauglehole 78670 (MEL); Snowfileds, Snowy Range, beside headwaters of Shaws Creek, 2 km NNE from the Gorge; 5 km NNW from Mt Arbuckle, 2 Apr. 1994, N.G. Walsh 3638 (MEL).

TASMANIA: South West Hibbs Bay, 27 Jan. 1984, 4.M Buchanan 2840 (HO); East Coast, Broad River near Cluny Lagoon, 8 Oct. 1989, P Collier 4270 (HO); North East Wedgetail Peak, 22 Oct. 1983, A. Moscal 3769 (HO); Central Highlands Little Split Rock, 8 Mar. 1984, A. Moscal 6738 (HO).

SOUTH AUSTRALIA: Southern Lofty, Upper Hermitage, R../. Bates 29684 (AD); Kangaroo Island, Ravine des Casoars, 21 Dec. 1992, RJ. Bates 30423 (AD, CANB); Rivoli Bay, 1888, F! Mueller s.n. (MEL); SE Wirrega, ca 25 km NW of Bordertown, ca 32 km SE of Keith, 2 Oct. 1916, R. Tate s.n. (AD); Blanchetown (ca. 115 km NE of Adelaide), s.dat., R. Tate s.n. (AD 97602434).

WESTERN AUSTRALIA: Hamersley Inlet, 3.6 km WNW of Edwards Point (IRNP), 28 Sep. 1986, K. Newbey 11162 (PERTH); Porongurup Range, Castle Arch, Sep. 1958, G.G. Smith s.n. (PERTH).

7b. Stellaria multiflora subsp. collaris C.H.Miull. & J.G.West subsp. nov.

A subsp. multiflora et subsp. nebulosa pedicelo collo Juxtim fructu, distinguenda; quoque a subsp. multiflora fructu maturo angusto—ovoideo < 2 mm lato, et valva apice recto nunquam revoluto, differt; et a subsp. nebulosa petalis semper praesentibus differt. Holotypus: Australian Capital Territory: Western slope of Black Mountain, 11 Oct. 1964 HS. McKee 11668 (CANB 145524). Isotypi: CANB 319264, NSW. Stellaria aff. filiformis, N.Burb. & M.Gray, FI. Austral. Capital Terr. 166 (1970). Stellaria sp. D sensu Doust in G.J.Harden, Fl. N.S.W. 1: 276 (1990). Stellaria sp. 1 sensu C.H.Mill. & J.G.West in N.G.Walsh & Entwisle, Fl. Victoria 3: 238, Fig. 45p-q (1996). Stellaria sp. 1 Flora of Victoria (H.S.McKee 11668) C.H.Mill., Austral. Pl. Cens. database (APC), http:// www.anbg.gov.au/cgi-bin/apc [accessed 6 May 2010].

Glabrous herb with erect to spreading stems, to 20 (-23) cm long. Stem and inflorescence leaves in a continuous series, sessile to appearing petiolate; obovate, linear or lanceolate, (2—) 3-11 (—13.5) mm long, 0.5—2 (—2.5) mm wide, acute to subacute. Inflorescence a monochasium of (2—) 5—15 (—27) flowers. Pedicels to 8 (—10) mm long, rarely longer with distinctive collar-like thickening present in fruit. Sepals 3—5.5 (—6) mm long. Petals 3—5 (—6), (0.2—) 0.4—1 (—1.2) mm long, usually bifid. Stamens (2—) 3—5. Staminodes 0. Styles 0.2—0.6 mm long. Capsule narrow ovoid, (2.5—) 3.5—5.5 (—6) mm long, 1—1.5 (—1.8) mm wide, | to 1.5 times sepal length; valves spreading, rarely straight, apex straight,

S flower showing sepals and mature capsule; T seed, side view; U seed coat cell detail. Scale bars: habits (A, H, P) 5 mm; flowers & fruits 1 mm; seeds (G, O, U) 0.5 mm. — A-G RJ. Bates 3652 (AD 98421323); H-O J. McKean 5789 (CANB 327583); P-U R.J. Bates 3577 (AD 98433224),

47

C.H. Miller & J.G. West

never revolute. Seeds (9—) 12-20 (—25), reniform to suborbicular, 0.5—0.7 (—0.8) mm long, light to reddish- brown, tubercles narrow rounded ridges, semi-inflated. Fig. IM, 4P-V. Flowering: (June—) Aug.—Oct. (—Dec.)

Distribution and habitat. Subsp. collaris occurs in south- eastern Queensland, with a sporadic inland distribution from Chinchilla and Warwick. It also occurs on the slopes and plains of northern New South Wales from Cobar in the west to Breeza in the east and Temora in the south, with one collection from Black Mountain in the Australian Capital Territory. In Victoria it is found in the central districts from Bendigo west to Wyperfeld National Park. It occurs in South Australia, just across the border in Chowilla, north to Wilpena and the Gammon Ranges (Fig. 6J).

Locally common in understorey of eucalypt open forest and woodlands, Callitris woodlands or mallee shrublands. Grows on sandy to loam soils.

Notes. Subsp. collaris tends to be a larger, more robust plant than the other subspecies of S. multiflora. It 1s found at medium altitudes on the slopes of ranges or further west on the plains. It does not appear to grow in more extreme habitats such as alpine or coastal areas where the other subspecies of S. multiflora are found. This taxon has a disjunct distribution and more collecting is needed to determine the full extent of its range.

Etymology. The name is derived from the Latin collare which means collared, in reference to the collar-like structure that is present on the top of the pedicel, just under the mature fruit. This character is not found in any other Ste//aria occuring in Australia.

Selected specimens (of c. 30 seen)

QUEENSLAND: Near Warwick, Jun. 1892, F-M. Bailey s.n. (NSW); Charleys Creek near Chinchilla, Sep. 1978, G. Lithgow 21 (BRI).

New SoutH WALEs: Breeza, Between Gunnedah & Werris Creek, Oct. 1899, W. Court s.n. (NSW); Western plains, 40 km NNW of Cobar, Bundella Station, Elura mining lease, Emu Tank, 8 Sep. 1978, MD. Crisp 4209 (CANB); Temora, Oct. 1915, Rev. .W. Dwyer s.n. (CANB); SW, Munagai Stn, 26 Jul. 1968, 7: Henshall 1062 (DNA); Trangie, Sep. 1968, 7: Hunter s.n. (NSW); SW Plains, Monaro Vale, Sep. 1983, D.E. Jones 1498 (NSW).

Victoria: Wyperfeld National Park, N of Black Flat, 19 Sep. 1968, A.C. Beauglehole 28268 (MEL); Wimmera Study Area, Ellam Flora reserve, 18 Sep. 1986, A.C. Beauglehole 84702 (MEL); 143rd Meridian Rd. 2—3 miles S of Murray Valley Highway SE of Robinvale, 4 Aug. 1973, N. Macfarlane 636 (MEL).

SOUTH AUSTRALIA: Chowilla Survey 1988, On road to Hypurna (still on Chowilla Station), 5 Aug. 1988, R./. Bates 14866 (AD); Lochness Well Gammon Range National Park, far Northern Flinders Range, 15 Oct. 1993, R.J. Bates 34268 (CANB); Northern Flinders Range, Gammon Ranges (ca 65 km E of Leigh Creek = Telford), Ridge N of North Tusk (ca 12 km E of Owieandana Hut), 19 Sep. 1956, H7. Eichler 12819 (AD); Wilpena hill slope opposite chalet, 16 Sep. 1960, D.E. Symon 656 (AD, BRI).

48

J. Adelaide Bot. Gard. 25 (2011)

7c. Stellaria multiflora subsp. nebulosa C.H.Miull. &

J.G.West subsp. nov.

A subsp. multiflora fructu maturo anguste ovoideo vel anguste ellipsoideo minus quam 2 mm lato, et valva apice recto nunquam revoluto, distinguenda; et a subsp. collaris CL\H.Mill. & J.G.West pedicelo collo juxtim fructu absente, et petalis absentibus, differt. Holotypus: South Australia: Region 13 South-eastern: Marshes Swamp, 14 Oct. 1991 RJ Bates 25994 (CANB 470161). Isotypus: AD 99146306.

Herb, glabrous or occasionally with sparse hairs. Stems prostrate to erect, to 16 (—23) cm long. Stem and inflorescence leaves in a continuous series, sessile to appearing petiolate; obovate, linear, elliptic or occasionally narrowly ovate, (1.5—) 2.5—11 (—17.5) mm long, (0.3—) 0.7—2 (—3) mm wide, acute to subacute, glabrous or with hairs on lower half of margin. Inflorescence a monochasium of 2—15 (—25) flowers. Pedicels (1.5—) 2-20 (—26) mm long, sometimes quadrangular, collar absent. Sepals 2—5.5 (—6) mm long. Petals 0. Stamens 3-10. Staminodes 0-5. Styles 0.3-0.8 mm long. Capsule narrow, ovoid to ellipsoid, (1.9-) 3—5.4 (-6.1) mm long, 1—2 mm wide, equal to 1.3 times sepal length; va/ves spreading or straight, apex straight, sometimes slightly recurved. Seeds (3—) 5-17 (—21), suborbicular, sometimes ellipsoid, (0.5—) 0.7-1 (—1.2) mm long, yellowish, light to mid-brown or rarely reddish-brown, tubercles broad hills, semi-inflated. Fig. IN, 4W-BB. Flowering: (Jul.—) Aug.—Dec. (—Feb.).

Distribution and habitat. This taxon occurs at Yetman on the northern slopes of New South Wales, in coastal areas of Victoria, from the Gippsland area to Wilson’s Promontory and from the eastern Bass Strait islands to north-eastern Tasmania. It is also found in south-western areas of Victoria around Dimboola, and south-eastern South Australia around the Mount Gambier area as well as on Kangaroo Island (Fig. 6K).

Common to locally abundant in a wide range of habitats including Eucalyptus baxteri coastal dune scrub, heathlands, low alpine herbfields and open eucalypt forests, often in damp areas on sandy to rocky soils.

Notes. Currently, subsp. nebulosa consists of those individuals of S. multiflora that lack the characters of the other two subspecies and does not have any unique defining characters such as the revolute capsule valves of subsp. multiflora and the thickened collar-like structure on the fruiting pedicel of subsp. col/aris. It is found in a diverse range of habitats and locations. There seems to be three groups within the distribution: one 1s a southern band from the north of Tasmania across the Bass Strait Islands with an outlier on the central Victorian coast; the second is centred around Dimboola in western Victoria across to eastern South Australia in the Mount Gambier region with an outlier on Kangaroo Island; and the third is from the central west of New South Wales north possibly into central Queensland. This taxon is poorly represented, particularly in the north and often these

J. Adelaide Bot. Gard. 25 (2011)

specimens do not contain fully mature fruit to allow for a definitive identification.

Etymology. This subspecies has been named from the Latin nebulosus which means ‘clouded, cloudy, misty or foggy’ which reflects the definition of this taxon which is currently those individuals that remain after other elements of the species have been defined.

Selected specimens (of c. 45 seen)

New SoutH WA Es: Mt Jagungal, Kosciuszko region, s.dat., D.N. McVean s.n. (CANB); Nicholls Rd, Bebo State forest, 1.3 km S of Wood Bend (10.2 km N of Yetman), 23 Aug. 1987, R.G. Coveny et al. 12701 (CANB, NSW).

VicToRIA: Grampians, Mt. Arapiles, 18 Feb. 1959, A.C. Beauglehole 5338 (MEL); Nurcoung Flora Reserve. Wimmera Study area, Sector A, subblock 4A, 13 Nov. 1986, A.C. Beauglehole 86892 (CANB, MEL); East Gippsland: Tildsley Forest Block. 200 m E of walking track from Gibbs Beach along 90 mile Beach. 20 m N in lee of primary dune, Nov. 1992, A.B. Pollock s.n. (MEL); South-west, Wilson’s Promontory Pillar Point track above mouth of Tidal River, 5 Oct. 1973, J.H. Willis s.n. (MEL).

TASMANIA: Furneaux, Behind Planter Beach, East Flinders Island, 26 Sep. 1989, P. Collier 4200 (HO); North East Reeves Creek Picnic Rocks, 13 Sep. 1983, A. Moscal 2675 (HO); Kents Group; North East Island, South Hull, 29 Sep. 1971, JS. Whinray 1150 (CANB); Furneaux Group, Flinders Island. High dunes c. 150 yards E of entrance of East River, s.dat., JS. Whinray 8318 (CANB).

SOUTH AUSTRALIA: South eastern, Marshes Swamp, 14 Oct. 1991, RJ. Bates 25992 (AD, CANB); Kangaroo Island, Ravine des Casoars, 21 Sep. 1992, R./. Bates 30244 (AD, CANB); South-eastern, Near Naracoorte South Primary School, c. 95 km N of Mt Gambier, 11 Sep. 1964, D. Hunt 2132 (NSW); Ca 30 km SE of Mt Gambier, Sep. 1966, /.B. Wilson 536 (AD, CANB).

8 * Stellaria pallida (Dumort.) Crep.

Man. Fl. Belgique, ed. 2, 19 (1866). — Alsine pallida Dumort., Fl. Belg. (Dumortier) 109 (1827). — Type citation: ‘In cultis humidis solo arenoso’. Type: BR?, n.v. S. pallida (Dumort) Pire, Bull. Soc. Roy. Bot. Belgique 2: 49 (1863), nom. inval.

Illustration. C.H. Mill. & J.G.West in N.G.Walsh & Entwisle, Fl. Victoria 3: 236, Fig. 45c—d (1996).

Annual, almost prostrate, semi-erect to erect and leafy to 30 (-41) cm long, with a single line of hairs down stem internodes and pedicels. Stem and inflorescence leaves a continuous series, sessile to appearing petiolate, lower part narrowed, very attenuate, (O—) 2.5—10 (—16) mm long, upper part broad elliptic to elliptic or ovate, (2.5—) 4-15 (-29) mm long, (0.9—) 1.5-10 (-17) mm wide, acute, entire, often undulate, hairs sparse to almost ciliate on narrowed leaf base margin. /nflorescence a leafy dichasium often condensed. Pedicels (2—) 3-15 (—18) mm long, erect or reflexed in fruit. Sepals (2.2—) 2.5—-4 (—5.1) mm long, obtuse, often forming small hood, rarely acute, sometimes with purple mark at apex , hairs on midrib only to densely covering backs, rarely glandular, sometimes glabrous. Petals 0. Stamens 2, rarely 3 or 4. Staminodes 0-2 (-4). Styles 0.4—0.5 (—0.6) mm long. Capsule ovoid to ellipsoid, (2.3—) 3-4

Stellaria in Australia (Caryophyllaceae)

(4.9) mm long, (1.5—) 1.7—2.2 (—2.5) mm wide, equal to slightly longer than sepals, rarely just shorter than sepals; valves spreading, apex only recurved, rarely revolute. Seeds (6—) 8-15 (—19), discoid to flattened ellipsoid, (0.55—) 0.6—0.8 (—1) mm long, yellowish to mid-brown, rarely darker, tubercles rounded hills, semi-inflated, cell walls with minute papillae. Fig. 11, 3N—-R. Flowering: all year around, but predominantly July—Nov.

Common name. Lesser Chickweed.

Distribution and habitat. Found at scattered localities within New South Wales, along the Great Dividing Range from Tenterfield to Canberra, with one collection inland at Broken Hill. It also occurs along the Murray River drainage system in New South Wales and Victoria. It is found throughout most areas of Victoria except alpine areas as well as throughout coastal and coastal ranges of South Australia including Kangaroo Island; the northern most collections extend into the Flinders Ranges and the western most collection is at Fowlers Bay near Ceduna. This species is uncommon in Tasmania being mainly found on offshore islands and in coastal regions but occasionally further inland. It is also uncommon in Western Australia with coastal collections from Rottnest Island and Busselton in the west, Bremer Bay in the south and Cocklebiddy in the east near the South Australian border (Fig. 6L).

The native range of S. pallida extends throughout most of central to western and southern Europe (Chater & Heywood 1993). It has also spread as a weed into the Americas and Asia.

In Australia, S. pallida is an introduced weed, often locally common, especially in areas of disturbance such as roadsides, waterways and areas of cultivation. Usually found in shady or moist places. It grows in coastal dunes, grass or herbfields, gully forests, open forests, heathlands or scrubs. It usually grows in sandy soils, sometimes in shallow soils over rock or loam to clay soils.

Notes. See notes under S. media. This weed probably has a wider distribution in Australia than listed here but is often overlooked or often misidentified as S. media. From examining the herbarium collections it appears S. pallida is a relatively recent introduction to Australia. There are very few collections prior to the 1950’s with the earliest collection recorded in 1905 at Tallandoon in Victoria.

Selected specimens (of c. 200 seen)

New SoutH WALEs: Murrurundi, park on Western side of New England Highway, 12 Oct. 1993, R.G. Coveny & A.J. Whalen 16555 (CANB); Southern Tablelands, Mirrunga, 8km S of ACT border, Murrumbridgee River at confluence with Gossoon Creek, 12 Oct. 1995, I. Crawford 3168 (CANB); Central Coast Salesyards, Flemington, 28 Sep. 1968, E.J/. McBarron 15864 bis (NSW); On road to Zara station, 0.7 km from entrance on edge of Billabong Creek, 21 Sep. 1992, CH. Miller & J. Palmer 595 (CANB); NW Plains, Iolanthe, c. 26 km SW of Garah, 28 Sep. 1978, K.L. Wilson 1913 (NSW).

49

C.H. Miller & J.G. West

M

J. Adelaide Bot. Gard. 25 (2011)

1600 km

Fig. 6. Distribution maps of Sfellaria in Australia. A S. angustifolia subsp. angustifolia; B S. angustifolia subsp. tenella; C S. angustifolia subsp. rotundisepala; D S. filiformis; E S. flaccida; F S. graminea; G S. leptoclada; H S. media; | S. multiflora subsp. multiflora; J S. multiflora subsp.

collaris; K S. multiflora subsp. nebulosa; L S. pallida; M S. papillata; N S. pungens.

J. Adelaide Bot. Gard. 25 (2011)

AUSTRALIAN CAPITAL TERRITORY: Hill opposite Cotter Pumping Station, Murrumbidgee River, 21 Aug. 1964, J. Edyvane s.n. (CANB, NSW).

Victoria: Mt Arapiles, ca 20 miles W of Horsham, the NE portion of the mount near the lookout, 3 Oct. 1963, H.[Aston 1064 (MEL); Grampians National Park, 30 Aug. 1983, 4.C. Beauglehole 74484 (MEL); Wilsons Promontory National Park, 15 Dec. 1983, A.C. Beauglehole 75969 (MEL); Port Campbell National Park, W of Port Campbell, W side of Port Campbell Creek, 5 Sep. 1966, A.C. Beauglehole & E.W. Finck 21053 (MEL); Ca 1 km inland from the mouth of the Aire River, 5 km SW of Horden river, 28 July 1980, P.C. Heyligers 80020 (CANB); Thompsons Beach, Murray River at Cobram, 24 Sep. 1992, C_.H. Miller & J. Palmer 603 (CANB); Little Desert Salt Lake, Coynallan Parish, 15 Oct. 1966, J.H. Willis s.n. (MEL).

TASMANIA: Midlands Port Sorell, 20 Sep. 1986, P Collier 1630 (HO); East Coast Betsey Island, 15 Oct. 1983, K. Harris s.n. (HO); Furneaux Group, Prime Seal Island, off West coast of Flinders Island, 11 Dec. 1986, S. Harris s.n. (HO); North East, Picnic Rocks, 19 Sep. 1983, A. Moscal 2836 (HO, MEL); Waddles, Hunting Grounds, Jordan river, 20 Aug. 1965, M. Ridpath WL420 (CANB).

SOUTH AUSTRALIA: Muray Mallee, Reserve at Chauncey’s Line S of Monarto South (Ca 60 km SE of Adelaide), 23 Sep. 1959, Hj. Eichler 16201 (AD); Kangaroo Island, Block 101, Kingscote North, 24 May 1983, G. Jackson 1590 (AD); Piccaninnie Blue Lake, ca 20 km E of Port MacDonnell (near the border of Vic & SA), 29 Aug. 1964, 71R.N. Lothian 2935 (AD); Adelaide, Marlowe Rd, Keswick, 09 Sep. 1987, J. Roberts 289 (CANB); Yorke Peninsula, Port Davenport, 23 Sep. 1978, D.E. Symon 11072 (AD); 3 km W of Blinman on rd to Parachilna Gorge, 29 Sep. 1985, J.G. West 5067 (CANB); Fowler’s Bay, about 6 km W along roadside, (Fowler’s Bay ca 115 km W of Ceduna), 12 Sep. 1960, D.J.E. Whibley 608 (AD, CANB); Southern Flinders Range, Mambray Creek (Mt Remarkable National Park), ca 40 km SSE of Port Augusta, Lower Alligator Creek area, 7 July 1974, D.J.E. Whibley 4404 (AD).

WESTERN AUSTRALIA: Eyre, SW Cocklebiddy, 8 Oct. 1985, G.J. Keighery & J.J. Alford 434 (PERTH); Rottnest Island, Bickley Swamp, 14 Aug. 1987, G./. Keighery 9063 (PERTH); Busselton, 20 July 1987, GJ. Keighery 9115 (PERTH); Kooljerrenup Nature reserve; 400 m S of Herron Point, 12 Oct. 2005, G.J. Keighery 16732