Swainsona 33: 11–29 (2020) © 2020 Board of the Botanic Gardens & State Herbarium (Adelaide, South Australia)

Grammosolen (Solanaceae - Anthocercideae) revisited

L. Haegi
State Herbarium of South Australia, GPO Box 1047, Adelaide, South Australia 5001
Email: laurence.haegi@sa.gov.au

Abstract: Study of newly collected specimens from beyond the former known geographical range
of Grammosolen, together with consideration of newly published results of molecular phylogenetic
studies in tribe Anthocercideae have led to an expanded concept of the genus. The rare G. archeri Haegi
is newly described from far south-eastern Western Australia; G. odgersii is transferred from Cyphanthera;
and the known range of G. truncatus, formerly regarded as an endemic South Australian species, is
extended to adjacent far eastern Western Australia.

Keywords: Grammosolen, Cyphanthera, Anthocercideae, Solanaceae, taxonomy, new species,

intergeneric hybrids

Introduction Materials and Methods

Grammosolen Haegi was first described as a new genus This study is based on investigations carried out over
in the context of a monographic study leading to the several decades, commencing in the first instance as
re-instatement of the distinctively Australasian tribe part of the subject of a PhD thesis (Haegi 1983). It
Anthocercideae G.Don (Haegi 1981, 1983). The study is based on detailed morphological examination of all
led to the recognition of 31 species in seven genera– available herbarium material in Australian and relevant
Anthocercis Labill., Anthotroche Endl., Crenidium Haegi, overseas herbaria as well as many specimens collected
Cyphanthera Miers, Duboisia R.Br., Grammosolen and in the course of the study. Measurements of each
Symonanthus Haegi (Haegi 1981, 1983). Grammosolen character used in the descriptions were, where possible,
was established to accommodate two closely related, taken from a sample of between 20 and 30 specimens
distinctive species: G. dixonii (F.Muell. & R.Tate) selected to represent the whole geographic range.
Haegi (formerly mistakenly assigned to Newcastelia Characters which showed variability were measured for
F.Muell. in family Verbenaceae) and G. truncatus (Ising) at least two organs on each specimen in the sample and
Haegi, originally described as a species of Anthotroche. in the case of leaves, the dimensions of the largest as
One new species has since been described in each of well as of two or three of those judged to be close to
Anthocercis (Macfarlane & Wardell-Johnson 1996) and the mode were recorded. Dried material was used for
Duboisia (Craven et al. 1995), bringing the total to 33 all measurements, except the flowers which were either
softened in a weak solution of detergent and hot water
species in the tribe.
or for which spirit material collected in the course of
the study was used. Leaves were measured to the nearest
Grammosolen has been known only from the state of millimetre with a ruler, while most other organs were
South Australia. Additional collecting in more remote measured to the nearest ½ millimetre using a low-power
parts of Australia has since not only extended the known stereo-microscope fitted with an ocular micrometer, at
distribution in South Australia but has also brought to 10× magnification. Larger hairs (1.0–2.5 mm long)
light specimens from Western Australia. Some of these were measured in this way to the nearest 0.05 mm;
represent a range extension of G. truncatus a relatively smaller hairs (0.025–0.075 mm) were measured to the
short distance across the border from South Australian nearest 0.025 mm at 40× magnification. Indumentum
populations nearby. Others, confined to a small area terminology follows Haegi (1991).
much further west, inland from Cape Arid on the
southern coast, have proved to be not readily assignable Chromosome numbers were investigated for three
to either existing species, or to any other species in species from the study of meiosis in pollen mother cells.
the tribe. These specimens are consistently distinct To date it has not been possible to obtain living material
in several features, including their small, orbicular- of the fourth species, G. archeri. Flower buds collected
reniform to obcordate leaves and in their smaller in the field were placed immediately in a mixture of 3
flowers and fruits. The new species Grammosolen parts of ethanol to 1 part of glacial acetic acid (mixed
archeri, endemic to south-eastern Western Australia, is in situ). After 24 hours the buds were transferred to
described here on the basis of these specimens. 100% ethanol, kept in cool conditions and finally

Published online: 31 Jan. 2020 • flora.sa.gov.au/swainsona ISSN 2206-1649 (Print) • ISSN 2206-1657 (Online)
L. Haegi Swainsona 33 (2020)

stored in temperatures below 0°C. Standard anther More recently, molecular phylogenetic studies of the
squash techniques (Darlington & La Cour 1970) Anthocercideae and near relatives, analysing sequence
were used to obtain microscope slide preparations for data from four chloroplast DNA regions — ndhF and
observing chromosomes at meiosis. The preparation trnL/F (Garcia & Olmstead 2003), as well as trnS-G
was examined with a compound light microscope at and matK (Clarkson et al. 2004) — have provided new
up to 1,000× magnification, sometimes with the aid of evidence for assessing the affinities and relationships
phase contrast illumination. The results were recorded of these genera. It is now generally accepted that
using a camera lucida device fitted to the microscope. together with the genus Nicotiana, these genera make
up a distinctive grouping within the family, for the
Although it can be caused by other factors (such as time being treated as subfamily Nicotianoideae. This is
environmental ones) sterility of pollen is a useful guide confirmed and placed within the context of an overview
to the occurrence of hybridisation, especially when of the molecular phylogeny of the Solanaceae as a whole
pollen from possible parents sampled at the same time by Olmstead et al. (2008). There is strong support
proves highly viable. Stainability of pollen was tested as for continued recognition of Tribe Anthocercideae
an indicator of viability using 0.5% lactophenol cotton (Clarkson et al. 2004), perhaps with some uncertainty
blue, which stains cytoplasm. Pollen from a single about the placement of the genus Symonanthus, whose
mature anther from each of several different flowers relationships with the remainder of the Anthocercideae
from recently collected herbarium specimens was on the one hand and Nicotiana on the other, remain
teased out into a drop of stain on a microscope slide, unresolved (Olmstead et al. 2008). Within the
a coverslip was applied and the preparation examined Anthocercideae these molecular phylogenetic studies
with a compound microscope, with all grains present provide partial support for the morphologically-based
scored. generic classification (including continued generic
recognition of Grammosolen) but also provide evidence
The composite distribution map (Fig. 3) was prepared for a different approach to the circumscription of
utilising point distributional data from the AVH, some genera. On the basis of that evidence C. odgersii
viewed in the ALA portal (accessed 3 July 2019), is demonstrated, as part of a highly supported
downloaded and edited to remove misidentified and “Grammosolen clade”, to have a closer relationship with
misplotted duplicate records. The map as presented the two known species of Grammosolen than to all the
was prepared using QGIS v. 3.2.0–Bonn, with the other species of Cyphanthera (Garcia & Olmstead 2003;
Clarkson et al. 2004).
map outline and features (coastline, sand-ridge and
lakes layers) from GEODATA TOPO 250K series 3
The evidence from molecular studies prompts a re-
Topographic data (Geoscience Australia 2018, accessed assessment of the diagnostic value of characters found
5 February 2018). in Grammosolen and Duboisia/Cyphanthera. Inclusion
of Cyphanthera odgersii in the Grammosolen clade
on molecular grounds is consistent with the shared
Generic placement of Cyphanthera odgersii occurrence of erect, hippocrepiform anthers dehiscing
by a hippocrepiform slit on the abaxial face (Figs 1,
When it was described, Grammosolen was unique 10, 11). This may prove to be a synapomorphy for the
within the Anthocercideae for the occurrence of expanded genus (together with the otherwise distinctive
multangulate hairs (in particular on the exterior of the genus Anthotroche). In the apparent plesiomorphic
corolla lobes), a di- or tri-dynamous androecium of five condition found throughout Duboisia (four species) and
fertile stamens and a gametic chromosome number of the remainder of Cyphanthera (eight species) the anthers
n=56. At that time attention was drawn to some close are sub-reniform, oblique (twisted towards the vertical
similarities between Cyphanthera odgersii (F.Muell.) and tilted abaxially) and dehisce by a semicircular slit
Haegi and the two known species of Grammosolen. along the distal margin (for example as in C. racemosa
Verticillately-branched trichomes are found in both (F.Muell.) Haegi – Fig. 1). While a dense indumentum
C. odgersii and G. dixonii, while all three species share of dendritic hairs occurs in some species of Cyphanthera,
a spike-like inflorescence and, perhaps most notably, the very dense woolly indumentum found in C. odgersii
hippocrepiform anthers (Haegi 1983). These last is unique in the genus (Haegi 1991). Similar hair-
two features are not otherwise found in Cyphanthera coverings are, however, found in Grammosolen and
(Haegi 1983). However, like all species of Cyphanthera C. odgersii shares elaborate verticillately branched hairs
for which a count had been determined (C. albicans with G. dixonii. Similarly the spike-like inflorescence
(A.Cunn.) Miers, C. microphylla Miers, C. myosotidea of C. odgersii is unique in Cyphanthera but occurs
(F.Muell.) Haegi and C. tasmanica Miers), C. odgersii in all species of Grammosolen, being particularly
has a gametic chromosome number of n=30 (Haegi well developed in G. dixonii. Flowers with only four
1983). On balance, C. odgersii was included within fertile didynamous stamens are characteristic of all
Cyphanthera rather than in Grammosolen for that reason Cyphanthera species. In Grammosolen the androecium
and because it shared with all the other species a corolla consists of five fertile, di- or tri-dynamous stamens; this
glabrous outside and a didynamous androecium of is probably the plesiomorphic condition. If C. odgersii
only four stamens. This classification has been widely is correctly placed in Grammosolen, it is possible that
adopted since that time. the loss of the fifth fertile stamen arose independently

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Swainsona 33 (2020) Grammosolen (Solanaceae - Anthocercideae) revisited

Fig. 1. Semi-schematic rep­

re­sen­tation of anther types
in Grammo­solen and Cyphan­
thera, showing flowers
with part of calyx and
corolla removed to reveal
stamens, together with
B excised stamens depicting
dorsal (abaxial), ventral
(adaxial) and lateral views
of anthers. A Grammosolen
(G. dixonii – Haegi 868G, × 8.5);
B  Cyphanthera (C. racemosa
(F. Muell.) Haegi – Haegi 1957
(NSW, CBG, PERTH), ×  7).
Illustration by R. Roden.

in this species, separately from the event which Grammosolen known at the time produced counts of
resulted in this condition in Cyphanthera and Duboisia. n=56 (Haegi 1983). Given it seems likely that n=56
Alternatively, the presence of a fifth fertile stamen in represents the derived state, the occurrence of the
the other three species of Grammosolen could represent plesiomorphic state in the clade (n=30 in C. odgersii) is
a reversion. The ontogenetic likelihood of such a not problematic.
reversion would require further investigation. Some
insight may be evident in the variable occurrence of On the basis of the compelling evidence from the
flowers with four or five fertile stamens in the related molecular phylogenetic studies and the assessment
species Crenidium spinescens Haegi: the absence of one above, C. odgersii is here transferred to Grammosolen,
of the fertile stamens could be the result of genetic with the generic circumscription emended accordingly.
masking rather than its irreversible loss. This transfer creates no nomenclatural problems at any
level. The species epithet has not been used previously
The multangulate hairs found in all species of in Cyphanthera and the lectotype species of the genus
Grammosolen, but absent throughout Cyphanthera, designated by Hunziker (2001) is C. ovaliflolia Miers
may have been secondarily lost in C. odgersii as a (currently regarded as a taxonomic synonym of
member of the Grammosolen clade or may have evolved C. albicans (Cunn.) Miers subsp. albicans; Purdie et al.
subsequently in the other species. Chromosome 1982; Haegi 1983).
numbers in the tribe were first counted by Barnard
(1949) who recorded n=30 and 2n=60 for Duboisia
leichhardtii and D. myoporoides. Ikanaga et al. (1979) Occurrence of putative intergeneric hybrids in
added a count of n=?28 for D. leichhardtii and confirmed the Anthocercideae
n=30 for D. myoporoides. Further investigations by
Haegi (1981, 1983), resulted in the first chromosome The possible occurrence of several intergeneric hybrids
number determinations for Symonanthus, Anthocercis, was recorded by Haegi (1983) and investigations
Cyphanthera, Grammosolen, Anthotroche and Crenidium. of the alkaloid chemistry of some of these putative
To the extent that counts were obtained (for 21 of the hybrids, together with the putative parents (El Imam
31 species), a strong correlation was demonstrated with et al. 1991) provided supporting evidence. Two of
morphological features characterising genera (Haegi the recorded hybrids involve Grammosolen, with one
1981, 1983). A gametic chromosome number of n=30 species, G. dixonii, being one of the putative parents in
was observed for five of the nine species of Cyphanthera each instance. This is reported on in more detail under
(including C. odgersii) while the only two species of G. dixonii below.

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L. Haegi Swainsona 33 (2020)

Taxonomy opposite or sub-opposite bracts, borne in leafy clusters

of cymes terminating condensed lateral branchlets,
Grammosolen Haegi shortly pedicellate. Calyx ± cyathiform, 5-lobed,
Telopea 2 (1981) 178. — Type species: Grammosolen persistent, lanate-tomentose. Corolla ± narrowly
dixonii (F. Muell. & R.Tate) Haegi tubular with ± regular (4–) 5 (–6)-lobed patent
Newcastelia auct. non F.Muell.: F.Muell. & Tate, Trans. limb, drab white with violet striations in throat [but
& Proc. Roy. Soc. S. Austral. 10 (1888) 81, p.p. as to unknown in G. archeri], glabrous or glandular-hairy
N. dixonii (= Grammosolen dixonii); J.M.Black, Fl. outside, glabrous inside except for the very densely
S. Austral. 3 (1926) 479, p.p. as to N. dixonii, excl. and minutely papillose lobes; aestivation of lobes ±
W.A. material; C.A.Gardner, Enum. Pl. Austral. Occ. 3 induplicate, the margins of each lobe overlapping each
(1931) 111, p.p. as to N. dixonii, excl. W.A. material; other or ± meeting, the lobes free and erect; tube ±
J.M.Black, Fl. S. Austral. edn. 2, 4 (1957) 721, p.p. as narrowly funnel-shaped; lobes broadly ovate to narrowly
to N. dixonii, excl. W.A. material; Beard, Descr. Cat. W. elliptic with truncate base, very densely and minutely
Austral. Pl. (1965) 92, p.p. as to N. dixonii, wrongly papillose on upper (inner) surface, otherwise glabrous.
recorded for W.A. Stamens epipetalous at base of corolla-tube, 4 or 5,
Anthotroche auct. non Endl.: Ising, Trans. & Proc. Roy. usually didynamous, very rarely tridynamous, slightly
Soc. S. Austral. 46 (1922) 605, t. 38, 39, Fig. 1, p.p. as recurved prior to and ± erect at anthesis, included in
to A. truncata (= Grammosolen truncatus); J.M.Black, the corolla-tube; anthers much shorter than filaments,
Fl. S. Austral. 3 (1926) 500, p.p. as to A. truncata; Fl. unilocular, free, erect, hippocrepiform, dehiscing
S. Austral. edn 2, 4 (1957) 754, p.p. as to A. truncata; by an abaxial hippocrepiform slit. Ovary 2-locular,
N.T.Burb., Dict. Austral. Pl. Gen. (1963) 20, p.p.; surrounded at base by a moist, annular, aromatic but
H.Eichler, Suppl. J.M. Black’s Fl. S. Austral. (1965) non-nectariferous disc; ovules few; style included; stigma
226, 273, p.p. as to Newcastelia dixonii and Anthotroche
± capitate, very shortly bilobed. Fruit a smooth capsule
blackii as applied to S.A. specimens including all those
cited; D’Arcy in Hawkes, R.N.Lester & A.D.Skelding, with marginicidal and loculicidal, septifragal dehiscence
Biol. Taxon. Solanac. (1979) 12, 14, 17, 21, p.p.; into 4 valves, somewhat concealed by the dense, long
Haegi, ibid, 121–124, p.p. indumentum of subtending branchlet and calyx; calyx
non-accrescent but as long as or longer than capsule;
Anthocercis auct. non Endl.: F.Muell., Fragm. 10 (1876)
seeds few, subreniform with reticulate testa; embryo
19, p.p. as to A. odgersii (= Grammosolen odgersii, q.v.).
slightly curved, not coiled. Gametic chromosome number
Cyphanthera auct. non Miers: Haegi, Telopea 2(2) (determined for three of the four species) 30 or 56.
(1981) 177; R.W.Purdie, Symon & Haegi, Fl. Austral.
(1982) 27; Hunziker, Gen. Solanac. (2001) 386–388, Distribution. As newly circumscribed, Grammosolen
all p.p. as to C. odgersii (= Grammosolen odgersii). is a genus of four species; these have non-overlapping
Erect ± leafy, taprooted shrubs, suckering from stem ranges, extending from the Avon Wheatbelt through the
base following damage to or removal of aerial parts, Coolgardie, Great Victoria Desert and Mallee regions in
densely tomentose with variously dendritic hairs on southern Western Australia, to the Great Victoria Desert,
all vegetative parts, with corky fissured bark at stem western Gawler Ranges, Eyre Yorke Block and the
base. Leaves alternate, petiolate to subsessile, entire. Murraylands in South Australia. All of the species occur
Flowers bisexual, protogynous, subtended by a pair of as scattered, small populations on local patches of sandy

Key to species
1. Leaves 2.5–7 mm long, orbicular-reniform to obcordate (sometimes broadly ovate at shoot tips),
L:B mostly 0.65–0.96; calyx 2.0–2.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. G. archeri
1: Leaves 7–30 mm long, ovate, ovate-elliptic, ovate-triangular (sometimes broadly so) or subcordate,
L:B mostly 0.8–3.8; calyx 2.5–7 mm long
2. Leaves conspicuously petiolate, with petiole 2–4 mm long; leaf base cuneate; indumentum
very densely and closely tomentose with multangulate-dendritic hairs 0.05–0.15 mm long
above and below . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. G. truncatus
2: Leaves of subsessile appearance, with petiole 0.5–1.5 mm long, rarely 3 mm long in larger leaves; leaf base
rounded-truncate, cordate, cuneate or attenuate; indumentum densely to very densely tomentose to lanate-
tomentose with multangulate-dendritic and verticillately-branched hairs 0.2–1.0 mm long
3. Leaves ovate-triangular to very broadly so, sometimes subcordate; base rounded truncate,
sometimes cordate or broadly cuneate; multangulate-dendritic and verticillately-branched
hairs both present; leaves appressed or distally inclined, more or less overlapping;
corolla lobes mostly 3–6 mm long; stamens 5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. G. dixonii
3: Leaves broadly to narrowly elliptic, base cuneate to attenuate; indumentum consisting entirely of
verticillately branched hairs (other than small, inconspicuous glandular hairs); leaves spreading;
corolla lobes 1.3–2.5 mm long; stamens 4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. G. odgersii

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Swainsona 33 (2020) Grammosolen (Solanaceae - Anthocercideae) revisited

soil or in extensive dune systems, generally on disturbed (–0.7) mm on the underside; base rounded-truncate,
sites within or in association with mallee vegetation. sometimes cordate or broadly cuneate; apex acute or
slightly obtuse, less commonly acuminate, usually
1. Grammosolen dixonii (F.Muell. & R.Tate) Haegi angular or blunt, rarely rounded; margin flat or often
Telopea 2 (1981) 178; R.W.Purdie, Symon & Haegi, undulate, sometimes slightly recurved; midrib obscure
Fl. Austral. 29 (1982) 29; Haegi in Jessop & Toelken, adaxially, slightly prominent abaxially. Flowers borne in
Fl. S. Austral. edn 4, 3 (1986) 1242. — Newcastelia leafy clusters consisting of one to three, 1–3-flowered
dixonii F.Muell. & Tate, Trans. & Proc. Roy. Soc. S. cymes on much-condensed lateral branchlets, sometimes
Austral. 10 (1888) 81, as ‘Newcastlia Dixoni’. — aggregated to form leafy spikes; inflorescence-bracts 1–4
Type citation: “On sand ridges at Ral-Ral on the × 0.5–2 mm, moderately tomentose with multangulate-
River Murray, 30 miles [c. 48 km] from the Victorian dendritic hairs. Pedicel 0.2–0.8 mm, oblique, glabrous.
border; also at Crystal Brook; Mr Samuel Dixon.” Calyx 3.3–5 mm, outside densely to very densely
Lectotype (here designated): Ral Ral, S.A., S. Dixon tomentose with eglandular multangulate-dendritic
s.n. (MEL41019). Isolectotypes: AD97221081A,
and verticillately branched hairs 0.2–0.3 (–0.8) mm
BM629250. Remaining syntype: Crystal Brook, S.A.,
S. Dixon s.n. (AD97221081B). and upper layer 0.4–0.8 (–2.0) mm, inside moderately
pubescent with porrect to antrorse glandular hairs 0.05–
Anthotroche blackii auct. non F.Muell.: H.Eichler, 0.15 mm, and multangulate-dendritic, forked or simple
Suppl. J.M. Black’s Fl. S. Austral. (1965) 273, p.p. as to eglandular hairs 0.15–0.3 mm on the lobes, often only
all S.A. specimens cited.
near the apex; calyx-lobes acute to slightly obtuse or
Erect, spreading or sprawling greyish shrub 0.3–2 m tall rarely acuminate, always angular, (0.4–) 0.5–1.1 (–1.5)
and 0.8–3 (–5) m diam., with branches (at least finally) times as long as tube, 1.0–2.5 (–3) × 1–1.5 mm, sinuses
ascending. Branches very densely lanate-tomentose with acute. Corolla (7.5–) 8–12.5 mm, dull white with
multangulate-dendritic and verticillately branched hairs violet striations, outside moderately pubescent with
mostly 0.6–1.3 mm and fewer hairs c. 1.5 times as long multangulate-dendritic hairs 0.2–0.3 (–0.4) mm on the
as these i.e. 0.8–1.5 mm, and scattered, inconspicuous lobes and upper tube, inside very densely papillose with
glandular hairs 0.1–0.4 mm, the indumentum gradually hairs 0.05–0.1 mm on lobes; tube 4.5–7.5 mm long.,
wearing off with age. Leaves loosely appressed or at diam. 1.0–1.7 mm at base, 2.5–3.5 mm at limb-base;
least distally inclined, more or less overlapping so as to lobes ovate to elliptic with truncate base, (L:B 1.4–2.2
obscure much of the axis, subsessile or with a petiole to (–2.8)), 3–6 (–7.5) × (1.6–) 2–4 mm, slightly narrowed
1.5 (–3) × 1 mm; lamina ovate-triangular to very broadly or broadened at the base, apex acute and usually
so, sometimes subcordate (L:B (0.8–) 0.9–1.5 (–2.5)), blunt or rounded, rarely angular, margin flat. Stamens
(6–) 8–16 (–20) × (4–) 6–15 (–17)  mm, densely to 5, the longer ones (3.5–) 4–5 mm and the shorter
very densely tomentose (including petiole) with stalked ones 2.5–3.5 (–4.2) mm; filament-bases and adjacent
multangulate-dendritic and verticillately branched corolla moderately to densely pubescent with simple
hairs, mainly 0.1–0.3 mm and fewer 0.2–0.5 mm, and or forked eglandular hairs 0.2–0.5 (–0.8) mm, less
scattered inconspicuous glandular hairs, 0.05–0.1 mm, dense abaxially; anthers 0.8–1.0 × 0.8–1.0 mm. Ovary
the branched hairs usually denser and up to 0.5 very broadly ovoid to globose or depressed globose,

A
B

Fig. 2. Grammosolen dixonii.

A flowering branchlet, × 1;
B flower, × 2.5; C calyx of
advanced bud opened out,
with pistil, ×  6; D opened
corolla of same, × 6; E capsule,
× 6; F seed, ×  10. (A–D
D E F Haegi 1331; E–F Haegi 869).
Illustration by M. Perkins.

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L. Haegi Swainsona 33 (2020)

0.6–1.2 × 0.6–1.2 mm; ovules 4–9 in total; style 2.7–3.5 adjacent northern plains west of the southern Flinders
(–4.2) mm, ± equal to the longer stamens and included Ranges and northern Mount Lofty Ranges; and east of
by 0.5–2 mm; stigma c. 0.2 × 0.4 mm; disc undulate, these ranges, in an area on both sides and in the vicinity
0.2–0.5 mm high, usually slightly, rarely much greater of the River Murray (Fig. 3).
in diameter than the ovary (to 1.6 mm), red. Capsule
± inconspicuous, ± embedded in dense long hairs Grammosolen dixonii is found almost exclusively on
of enclosing calyx and axis, ± globose, 3.5–4.5 × calcareous pale red sandy soils in mallee vegetation
3–4.8 mm. Seeds 1–5, 2.5–3.5 × 1.1–1.6 mm; surface- and is frequently associated with Triodia. It is generally
reticulae 0.1–0.35 × 0.1–0.2 mm. Chromosome number: a plant of disturbed sites, which promote seed
n=56 (L. Haegi 676, 1457M). Fig. 2.
germination and may be locally frequent following fire.
Typification. This species was described (under Mechanical damage or burning of the aerial parts may
the basionym, Newcastelia dixonii) by two authors, stimulate the production of suckers from the stem base.
F. Mueller and R. Tate, and two localities are given in the
protologue. Material corresponding with information Phenology. Flowering specimens have been collected in
in the protologue has been located on three sheets, one all months except February, May and June, with most
each in AD, BM and MEL. But for one piece on the from August to January. Fruiting specimens have been
AD sheet, these specimens all match one another and collected from November to January. Galling of fruits
resemble material from the stated area of collection, viz. as a result of insect attack is a common occurrence.
Ral-Ral on the River Murray. Although not labelled as
such, the distinctive piece on the AD sheet, as indicated Conservation status. Grammosolen dixonii is relatively
by the more distinctly cordate leaves and undulate leaf rare, with many populations occurring in depleted
margins, resembles material known from the present remnant vegetation on roadsides in agricultural areas.
study to be restricted to the northern Yorke Peninsula Populations are conserved in several reserves such as the
region and Crystal Brook area. This specimen is almost Lake Gilles, Clements Gap, Pooginook and Cooltong
certainly that referred to in the protologue as having Conservation Parks.
been collected at Crystal Brook. One syntype collection,
therefore, is in fact divided so that part is mounted on Diagnostic features. Grammosolen dixonii is clearly
each of three sheets (AD, BM, MEL) while the other distinct from the other species in the genus, on account
collection consists of a single piece on the AD sheet.
of its ovate-triangular to sub-cordate leaves and the
The specimen MEL41019 is designated Lectotype; it
bears a label in Mueller’s hand and is the most ample combination of stalked multangulate-dendritic and
specimen. It had been labelled (mistakenly in light of verticillately branched hairs on the axes and leaves.
the foregoing discussion) as a holotype. There is no intergradation with other species, though
in the vicinity of Kyancutta on Eyre Peninsula,
Distribution and Ecology. Known only from South populations at the north-western extreme of the
Australia where it occurs in scattered populations at range of G. dixonii closely approach, within 25 km,
similar latitudes, in three main areas: on northern Eyre populations at the furthest south-eastern extent of the
Peninsula; northern Yorke Peninsula together with the occurrence of G. truncatus.

G. odgersii subsp. occidentalis

G. odgersii subsp. odgersii
G. archeri
G. truncatus
G. dixonii

Fig. 3. Southern (central and western) Australia, showing the geographical occurrence of 4 species of Grammosolen, including
two subspecies (from AVH, ALA–accessed 3 July 2019 and corrected) in relation to extent of sand-ridges (tan) and lakes/drainage
systems (grey).

16
Swainsona 33 (2020) Grammosolen (Solanaceae - Anthocercideae) revisited

Variation. Plants with acuminate (rather than 26.xii.1978, L. Haegi 1598 (NSW, AD, BIRM, F, HO); Area
acute) leaf apices, undulate margins and less dense between Oak Dam (Frenchs Dam) and Gypsum Hills, c.
indumentum predominate in the northern Yorke 32 miles (c. 52 km) N of Overland Corner, 9.x.1965, D.E.
Peninsula populations; similar specimens are rare Symon 3627 (AD, CANB, K); 45 km NW of Renmark on
elsewhere. Leaves are mostly in the range 8–16 × road to Canopus Station, 27.ii.1976, L.D. Williams 7768
6–15 mm, but plants with smaller leaves are found (AD); Glossop, between Barmera and Berri, x.1963, Woolmer
mainly on Eyre Peninsula (e.g. Orchard 2935, leaves 7 75 (AD). Yorke Peninsula: 32 km NW of Port Wakefield,
× 6 mm) while the largest-leaved plants occur on Yorke 29.iii.1966, B. Copley 177 (AD); 8 km WNW of Bute,
Peninsula (e.g. Copley 144, leaves to 20 × 17 mm). 15.x.1975, L. Haegi 705 (AD).
Slight zygomorphy of the corolla is apparent in some
plants seen in the fresh state (e.g. Haegi 1331, from Occurrence of putative intergeneric hybrids involving
Northern Lofty Region). Two of the lobes assume an G. dixonii
upper, more vertical position and may be slightly closer Several specimens not referable to known species
together, while the remaining three lobes are oriented were encountered in small areas where populations of
more horizontally, though not otherwise differently G. dixonii overlap with those of species in two other
positioned than in regular flowers. There is insufficient genera of Anthocercideae: Cyphanthera myosotidea
correlation of variation in any of these characters in one case and Duboisia hopwoodii in the other.
either with other characters or geographically for the Closer examination reveals that the specimens are
recognition of any infraspecific taxa. visually intermediate between the two sets of parents,
suggesting a hybrid origin for these plants. To test this
Derivation of name. Named for Samuel Dixon, an notion further, specimens of the intermediates and
Adelaide geologist and naturalist who had a special parents from each of the two localities were subjected
interest in fodder plants and collected the type material to simple morphometric analysis and an assessment of
in the 1880’s (see Sharr 1978). pollen fertility.

Selected specimens (145 examined) Pollen viability of samples from both putative parents
and the morphologically intermediate specimens was
SOUTH AUSTRALIA. Eyre Peninsula: 29.5 km E of tested. A very low rate of staining was recorded in
Kimba, 4.x.1975, R.J. Chinnock 2747 (AD); Cook Range,
all intermediates, contrasting with high rates in the
Middleback Ranges, 21.ix.1981, N.N. Donner 8015 (AD);
60 km SW of Iron Knob on Kimba road, 16.x.1975, L.
putative parents and supporting the assessment of the
Haegi 706 (NSW, AD, SYD); 15 km ENE of Cowell on intermediates as of hybrid origin.
Mitchellville road, 26.viii.1977, L. Haegi 1304 (NSW, AD,
BRI, CANB); Between Cowell and Arno Bay, 27.xi.1961, Phytochemical investigations of the leaves of the
D.N. Kraehenbuehl 528 (AD); Foot of Iron Duke at eastern end putative parents and hybrids (El Imam et al. 1991)
of Middleback Range, 29.xii.1970, A.E. Orchard 2935 (AD, provide further evidence for the hybrid origin of
CANB); c. 6 km W of Plank Point, 2.ii.1977, L.D. Williams the intermediates; these studies revealed that each
9025 (AD, NSW); 10 km SE of Wudinna, 18.xi.1984, D.N. of the parents has a unique combination of tropane
Kraehenbuehl 5051 (AD). Northern Lofty: Sections 141– and pyridine alkaloids and that the alkaloid spectra
150, Hundred of Wiltunga, 13.ii.1966, B. Copley 34 (AD); of each set of morphological intermediates combine
Southern boundary of Section 195, Hundred of Wiltunga, those found in the parents. Together these various
c. 140 km NNW of Adelaide, 20.iii.1966, B. Copley 144 observations provide strong circumstantial evidence
(AD); Mundoora, c. 50 km SSE of Port Pirie, iv.1962, G. for the intermediates being hybrids. The occurrence
Gardiner s.n. (AD96323154); 5.7 km S of Wandearah East of intermediates limited to areas of overlap in the
Post Office on Port Broughton road, 29.viii.1977, L. Haegi distributions of each of the parents is consistent with
1331 (NSW, AD, CORD, K, MEL, MO, PERTH); W of this conclusion. In both cases, flowering periods
Ninnes, which is 120 km NNW of Adelaide, 8.xii.1963, overlap. No manipulated crossing experiments have
D.N. Kraehenbuehl 1049 (AD). Murray: Calperum Station, been performed to demonstrate that hybridisation is
Overflow Track, at a point c. 20 km NNE of Monash, possible.
4.ii.2015, J.R. Guerin 204 (AD); 20 km NE of Blanchetown
on Waikerie road, 5.x.1975, L. Haegi 676 (NSW, AD, BRI, Cyphanthera myosotidea × Grammosolen dixonii
CBG, NT, PERTH); 18 km WSW of Waikerie on Sturt Hwy
A few plants visually intermediate between these
to Blanchetown, 15.xi.1975, L. Haegi 868A–J [population
collection documenting variability] (AD, BRI, CANB, HO,
two species were encountered along the road verge
MEL, NSW, NT, PERTH); 16 km N of New Well on road between 20 and 30 km NE of Blanchetown, South
running almost due N to main Blanchetown–Waikerie road, Australia, the only area where the two species are
15.xi.1975, L. Haegi 869 (NSW, AD); 11 km NNE of Berri, known to be sympatric. Their presence followed major
21.x.1976, L. Haegi 1252 (NSW, AD); 21 km by road NE disturbance involving extensive earthworks associated
of Blanchetown on Waikerie road, 17.xi. 1976, L. Haegi with realignment of the road. Here G. dixonii was
1263 (NSW, AD); - also 13.x.1977, L. Haegi 1457G, H, I, conspicuous and locally common but C. myosotidea
L, M, O, P, Q, R, S, T, U, V, W, X, Y [population collection was rare. A quantitative assessment of these specimens
at site of occurrence of putative hybrid with Cyphanthera confirms their intermediacy when compared with
myosotidea, also documenting variability in G. dixonii] (AD); material of the two species found at the site (Table 1,
2 km NE of Stony Pinch Dam, c. 15 km NW of Renmark, Fig. 4). Similarly, pollen fertility was low in a specimen

17
 L. Haegi Swainsona 33 (2020)

Table 1. Comparison of certain characters in Cyphanthera myosotidea, Grammosolen dixonii and a putative hybrid.

Character Cyphanthera myosotidea Putative hybrid Grammosolen dixonii

Indumentum Inconspicuous, moderately dense Moderately conspicuous, Conspicuous, dense tomentum of

pubescence mainly of simple moderately dense tomentum of eglandular branched hairs
glandular (with a few eglandular) eglandular branched hairs
hairs

Leaf length (mm) 2–10 8–16 8–16

breadth (mm) 1–4 3–5 6–15
shape Ovate to elliptic or narrowly so Ovate Ovate-triangular to very broadly so
length : breadth (1.8–) 2.0–6.0 (– 8.0) 2.4–3.5 (0.8–) 0.9–1.5 (– 2.5)
margins Flat Slightly undulate Undulate

Inflorescence Flowers scattered in distal parts Flowers in open leafy spikes Flowers in dense, leafy spikes

Pedicel Long, conspicuous ± Short, inconspicuous Short, inconspicuous

Corolla-lobe shape Broadly elliptic Ovate-elliptic Ovate to elliptic

length : breadth 0.8–1.3 1.5–1.7 1.4–2.2

Alkaloid spectrum of leaves Hyoscyamine or atropine; Hyoscine; apohyoscine; Hyoscyamine; hyoscine;

(after El Imam et al. 1991). hyoscine (principal alkaloid); 6β-hydroxyhyoscyamine (principal apohyoscine; 3α-acetoxytropane;
norhyoscyamine or noratropine; alkaloid); tropine; two unidentified no particular alkaloid recorded as
6β-hydroxyhyoscyamine; bases. predominant.
3α-tigloyloxytropane;
scopine or oscine; tropine;
3-phenylacetoxynortropan-6,7-diol.

Table 2. Pollen sterility estimates in Cyphanthera myosotidea, Grammosolen dixonii and a putative hybrid.

Cyphanthera myosotidea Putative hybrid Grammosolen dixonii

% sterile pollen 2% 61% 2%

No. of grains counted 183 142 121

Voucher specimen Haegi 1262 Haegi 1265 Haegi 1263

16

14

12
Mean leaf length (mm)

10

8
Fig. 4. Scatter plot portraying
6
intermediacy of putative
hybrids between Cyphanthera
myosotidea and Grammosolen
4 dixonii, as demonstrated by
C. myosotidea leaf characters. Each plotted
2 Hybrid
point represents a specimen;
four leaves measured per
G. dixonii
specimen; all specimens
0 from Murray Region, South
0 2 4 6 8 10 12 14 16
Australia (cited above and
Mean leaf breadth (mm) below).

18
Swainsona 33 (2020) Grammosolen (Solanaceae - Anthocercideae) revisited

of the intermediate material but high in specimens of pollen mother cells at meiosis, while normal meiosis
the parents from the same site (Table 2). Staining grains was observed in the putative parents (see Table 4).
in the hybrid were markedly smaller than those of
the parents. Observation of meiosis in pollen mother El Imam et al. (1991) state that the establishment of
cells of the putative hybrid indicated a high rate of nornicotine and hyoscine as the principal alkaloids,
failure in pairing of chromosomes to form bivalents at together with the co-occurrence of other bases of
Metaphase 1, while normal meiosis was observed in the the parents firmly support the assignment of the
putative parents (see Table 2). intermediate as a hybrid (Table 3).

Noting that 6β-hydroxyhyoscyamine, a minor Specimens of this hybrid were at one time thought to be
component of C. myosotidea, is present as the principal a new species related to Cyphanthera albicans by South
base in the anomalous material and that tropane Australian botanists. Striking features distinguishing
alkaloid yields of this material are intermediate between the hybrid from species of the tribe are the glabrescent
those of the two parents, El Imam et al. (1991) observe branches and leaves combined with the undulate
that this is consistent with hybrid origin. They also margins of the leaves.
conclude that hybridisation of these species appears
to exert an influence on the capacity of the progeny to Specimens seen
convert hyoscyamine into hyoscine, a reaction generally Duboisia hopwoodii × Grammosolen dixonii: SOUTH
involving 6-hydroxyhyoscyamine as an intermediate. AUSTRALIA. Murray Region: 8 km N of Berri, 4.i.1970,
B. Copley 3000 (AD); 8 km N of Berri, 19.xii.1973, B. Copley
Specimens seen (all Murray Region, S.A.) 4181 (AD); 11 km NNE of Berri, 21.x.1976, L.  Haegi
Cyphanthera myosotidea × Grammosolen dixonii: c. 13 km S 1250 (AD); 2 km NE of Stony Pinch Dam, c. 15 km NW
of Waikerie on roadside, 2.ix.1974, N. Gemmell 293 (AD, of Renmark, 26.xii.1978, L. Haegi 1600 & 1601 (AD);
MEL); c. 21 km by road NE of Blanchetown bridge on 2 km NE of Stony Pinch Dam, c. 15 km NW of Renmark,
Waikerie road, 17.xi.1976, L. Haegi 1264 & 1265 (AD); 22.iv.1979, L. Haegi 1773 (AD); sand ridges 3 km W of
- also 13.x.1977, L. Haegi 1457A, B, E, J, K, N [part of Stony Pinch (burnt), 5.x.1980, A.E. Spooner 7265 (AD);
population collection, including both parents] (AD); 20 km Calperum Station, Overflow Track, at a point c. 20 km NNE
NE of Blanchetown bridge on Waikerie road, 3.xi.1978, L. of Monash 4.ii.2015, J.R. Guerin 205 (AD).
Haegi 1593 (AD); 28 km NE of Blanchetown on Waikerie Duboisia hopwoodii: SOUTH AUSTRALIA. Murray
road, 14.ix.1974, D.N. Kraehenbuehl 3392 (AD). Region: c. 128 m N of Renmark, 6.i.1926, J.B. Cleland
Cyphanthera myosotidea: 13 km SSW of Waikerie, 2.ix.1974, s.n. (AD966030806); c. 10 km N of Berri, 25.viii.1962,
N. Gemmell 297 (AD); c. 21 km by road NE of Blanchetown J.B. Cleland s.n. (AD97217266); Calperum Station,
bridge on Waikerie road, 17.xi.1976, L. Haegi 1262 (AD); c.  10 km ENE of Berri, 4.i.1970, B. Copley 2999 (AD); c.
- also 13.x.1977, L. Haegi 1457C, D, F [part of population 15 km NE of Monash, 22.iv.1957, H. Eichler 13823 (AD,
collection, including putative hybrid and both parents] (AD); CANB); 11 km NNE of Berri, 21.x.1976, L. Haegi 1251
Waikerie, 15.x.1968, Miller s.n. (AD96846176); 61 miles (AD); Canopus Station, c. 185 km ENE of Burra and c.
(c. 98 km) SW of Renmark, 18.ix.1967, B.H.A. Tönnies 95 km N of Renmark, 6.viii.1965, A.C. Robinson 34 (AD);
s.n. (AD97424250); 16 km WSW of Waikerie, 18.ix.1967, Calperum Station, Overflow Track, at a point c. 20 km
Wheeler 452 (AD); c. 15 km SW of Waikerie, 26.ix.1971, NNE of Monash 4.ii.2015, J.R. Guerin 203 (AD). NEW
D.J.E. Whibley 3618 (AD). SOUTH WALES. South Far Western Plains: 110 km S of
Broken Hill on Wentworth road, 1.ix.1962, J.B. Cleland s.n.
Grammosolen dixonii: see main treatment above. (AD97217258).
Grammosolen dixonii: see specimens cited under G. dixonii
Duboisia hopwoodii × Grammosolen dixonii
above.
Specimens apparently resulting from hybridisation of
these species were collected during the 1970’s from
2. Grammosolen truncatus (Ising) Haegi
two sites near Berri and Renmark, c. 200  km NE of Telopea 2(2) (1981) 178. — Anthotroche truncata
Adelaide, South Australia. Both sites have been visited Ising, Trans. & Proc. Roy. Soc. S. Austral. 46 (1922)
and only a small number of putative hybrids were seen 605, t. 38, 39, Fig. 1. — Type citation: “Ooldea, East-
at each. A much more recent collection made in 2015 West Railway Line, September 15, 1920, and Barton,
from a third site in the same area (Guerin 205) provides in the same district, September 19, 1920.” Lectotype
evidence of the intermittent continued recurrence of the (here designated): Ooldea, Sept. 1920, Ising 1297
hybridisation event. The parents are markedly different (AD98209191). Isolectotypes: AD, BM, BRI, DNA,
from each other in several morphological features CANB, K, L, MEL, NSW. Remaining syntype:
(Table 3) and the intermediacy of the hybrid is visually Barton, 19.ix.1920, Ising 1374 (AD).
obvious (Figs 2, 5 & 6). The average dimensions of
leaves from several specimens collected in the area of Erect, somewhat stiff, grey-green shrub 0.7–2 m tall and
occurrence have been plotted to demonstrate this diam. Branches very densely and closely tomentose with
quantitatively (Fig.  7). Pollen from a specimen of multangulate and multangulate-dendritic hairs mostly
putative hybrid origin showed markedly higher sterility 0.1–0.2 mm and fewer to 0.25 mm, the indumentum
than found in either parent (Table  4). A high rate of very gradually wearing with age. Leaves mostly
failure in pairing of chromosomes was observed in ± spreading, some in small clusters on much-condensed

19
L. Haegi Swainsona 33 (2020)

Fig. 5. Duboisia hopwoodii

× Grammosolen dixonii.
A flowering branchlet, × 1;
B flower, ×  3; C calyx of
advanced bud opened out,
with pistil, × 5; D opened
corolla of same, × 5. (Haegi
A D 1600). Illustration by M.
Perkins.

D C

Fig. 6. Duboisia hopwoodii.

A flowering branchlet, × 1;
B flower, × 3; C corolla of
advanced bud, opened out,
A × 5; D calyx and pistil of same,
× 5; E berry, × 5; F seed, × 10.
(Haegi 888). Illustration by M.
E F Perkins.

20
 Swainsona 33 (2020) Grammosolen (Solanaceae - Anthocercideae) revisited

Table 3. Comparison of certain characters in Duboisia hopwoodii, Grammosolen dixonii and a putative hybrid.

Character Duboisia hopwoodii Putative hybrid Grammosolen dixonii

Indumentum Glabrous Immature parts conspicuously Conspicuous, dense tomentum of

tomentose; mature parts eglandular branched hairs
glabrescent

Leaf length (mm) 35–105 8–16 8–16

breadth (mm) 2–8 3–5 6–15

length : breadth (6–) 10–35 (–71) 2.4–4.5 (0.8–) 0.9–1.5 (–2.5)

margins Flat Undulate Slightly undulate

Corolla-lobe shape Broadly ovate to orbicular Ovate-elliptic Ovate to elliptic

length : breadth (0.3–) 0.5–1.2 1.5–1.7 1.4–2.2

Stamens 4 4 or 5 5

Alkaloid spectrum of leaves Hyoscine; nicotine; Hyoscine (2nd major Hyoscyamine; hyoscine;
(after El Imam et al. 1991). nornicotine (principal alkaloid); apohyoscine apohyoscine; 3α-acetoxytropane;
alkaloid); N-acetylnornicotine; and 3α-acetoxytropane no particular alkaloid recorded as
N-formylnornicotine. in admixture; nicotine (in predominant.
hyoscine picrate mother liquor);
nornicotine (principal alkaloid);
N-formylnornicotine.

Table 4. Pollen sterility estimates in Duboisia hopwoodii, Grammosolen dixonii and a putative hybrid.

Duboisia hopwoodii Putative hybrid Grammosolen dixonii

% sterile pollen 2.8% 86.0% 1.0%

No. of grains counted 545 372 283

Voucher specimen Haegi 1599 Haegi 1600 Haegi 1598

80

70

60

50
Leaf length (mm)

Fig. 7. Scatter plot portraying

40 intermediacy of putative
hybrids between Duboisia
30
hopwoodii and Grammosolen
dixonii, as demonstrated by
leaf length and leaf length to
20 breadth ratio. Each plotted
G. dixonii
point represents a specimen;
10 four leaves measured per
Hybrid
specimen; all specimens
D. hopwoodii
from Murray Region, South
0
0 2 4 6 8 10 12 14
Australia or adjacent area
in New South Wales (cited
Leaf length to breadth ratio above).

21
L. Haegi Swainsona 33 (2020)

axillary branchlets but generally spaced so as not to Stamens 5, 3–4 mm and 2–3.2 mm; filament-bases
obscure axis, petiolate; petiole 2–4 × 0.5–0.8 mm, densely pubescent with simple, forked and sparingly
tomentose as for lamina; lamina ovate to ovate-elliptic, branched hairs, 0.2–0.25 mm, less dense abaxially;
sometimes broadly or narrowly so, less commonly anthers c. 0.8 × 1.0 mm. Ovary globose to depressed
elliptic (L:B 1.1–1.7 (–2.3)), (5–) 7–13 (–18) × (3.5–) globose, 0.8–1 × 0.8–1.1 mm; ovules 5–10 in total; style
4–10 (–11) mm, very densely and closely tomentose 2.9–4 mm, ± equal to the longer stamens and included
with multangulate and multangulate-dendritic hairs by 0.5–1 mm; disc undulate, 0.3–0.5 mm high, usually
0.05–0.15 mm above and below; base cuneate; apex slightly, rarely much greater in diameter than ovary (to
acute to slightly obtuse, blunt; margin entire, flat, or 1.5 mm), red. Capsule somewhat inconspicuous, partly
often slightly thick or recurved; midrib faintly prominent embedded in hairs of enclosing calyx, axis and leaves,
above, prominent, conspicuous and often with visible globose, 4–4.5 mm diam. Seeds c. 4, c. 2.8 × 1.2 mm;
secondary veins below. Flowers borne in short, leafy surface-reticulae 0.1–0.25 × 0.1–0.25 mm. Chromosome
clusters of one or two, 1–3-flowered cymes on much- number: n=56 (Haegi 1597). Fig. 8.
condensed lateral branchlets, scarcely aggregated to
form leafy spikes; inflorescence-bracts 1.5–3 × 0.7–1 mm, Distribution and Ecology. Grammosolen truncatus
moderately tomentose with multangulate-dendritic occurs in South Australia and Western Australia,
hairs. Pedicel 0.3–0.6 mm, glabrous to moderately extending from north-west Eyre Peninsula and the
pubescent with multangulate-dendritic hairs c. 0.15 mm western Gawler Ranges north-west into the southern
and glandular hairs 0.1–0.15 mm. Calyx 2.5–4 mm, Great Victoria Desert. It is always found on deep sandy
outside very densely tomentose with multangulate- soils, commonly on sand dunes or localised sandy rises. It
dendritic hairs mostly 0.1–0.2 mm, with fewer c. is primarily a plant of burnt or otherwise disturbed sites
0.25 mm, and scattered glandular hairs 0.1–0.15 mm, in mallee vegetation, sometimes associated with Callitris
inside moderately pubescent with antrorse glandular and seems to occur mainly in small scattered populations.
hairs, 0.08–0.15 mm, especially on the lobes; calyx-lobes See G. dixonii above for comments on the co-occurrence
acute to slightly obtuse, angular or blunt, 0.2–0.4 times of these two species on Eyre Peninsula (Fig. 3).
as long as tube, 0.5–1 × 1–1.2 mm, sinuses obtuse.
Corolla (7.3–) 9.5–11 mm, ± dull-white with violet Phenology. Flowering specimens have been collected
striations, outside moderately to densely pubescent from August through February but the main flowering
with multangulate and forked hairs 0.2–0.25 mm on season appears to be in September and October.
lobes and upper tube, inside very densely papillose with Fruiting specimens have been collected in September,
hairs 0.1–0.2 mm on lobes; tube 4.5–6.3 mm, markedly October and January. Data are unavailable for the
widening in upper 0.5–1 mm, diam. 1.3–1.8 mm months March to July.
at base and 2.8–3.5 mm at limb base; lobes elliptic to
ovate with truncate base (L:B 1.7–2.4), (2.5–) 3.5–6 Conservation status. Grammosolen truncatus is an
× (1.5–) 2.0–2.8 mm, sometimes slightly narrowed at uncommon plant known from only about 70 collections.
the base, apex rounded to bluntly acute, margin flat. It is not recorded from any conservation reserves.

C
A

Fig. 8. Grammosolen trun­

catus. A flowering branchlet,
× 1; B flower, × 2.5; C corolla
of advanced bud, opened
out, × 6; D opened calyx and
pistil of same, × 6; E capsule,
× 6; F  seed, × 10. (Haegi 770).
D E F Illustration by M. Perkins.

22
Swainsona 33 (2020) Grammosolen (Solanaceae - Anthocercideae) revisited

Diagnostic features. Grammosolen truncatus is clearly 1990, W.R. Archer 510907 (PERTH). Isotypes: AD,
related to G. dixonii and G. archeri but is perhaps closer BM, CANB, MEL, MO, to be distributed.
to G. archeri. The western extremity of the range of
G. dixonii almost meets but does not overlap with the Spreading grey-green shrub, 0.6–1 m tall and up to 2 m
eastern extent of the occurrence of G. truncatus, while diam. Branches very densely and closely tomentose with
G. archeri has a distinct geographical range separated by multangulate and multangulate-dendritic hairs, mostly
several hundred kilometres from both other species. c. 0.2 mm and fewer to 0.25 mm, the indumentum
gradually wearing off with age. Leaves mostly ±
Variation. Grammosolen truncatus is relatively uniform spreading, some at times in small clusters on much
throughout its range, with both the smallest-leaved condensed axillary branchlets but generally spaced so
specimens (e.g. Symon 8120, Blaylock 2035) and the as not to obscure the axis, petiolate; petiole 0.4–2.2 ×
largest-leaved ones (e.g. Haegi 770) occurring in the 0.4–0.7 mm, tomentose as for lamina; lamina orbicular-
Gawler Ranges populations. reniform to obcordate, sometimes broadly ovate at the
shoot tips (L:B 0.65–0.96 (–1.1)), 2.5–7 × 2.6–10 mm,
Hybrids. None recorded. very densely and closely tomentose with multangulate
and stalked multangulate-dendritic hairs, 0.1–0.15 mm
Derivation of name. From the Latin truncatus, above and below; base usually truncate or shallowly
truncate, in reference to the shortly lobed calyx, as cordate, sometimes broadly cuneate; apex bluntly
indicated in the protologue. obtuse to rounded; margin flat, sometimes thick or
slightly recurved; midrib obscure adaxially, ± prominent
Notes. Some details in the protologue description have abaxially, sometimes with secondary veins. Flowers
not been substantiated in this study. My observations borne in 1–2-flowered cymes in the distal leaf axils of
indicate that the hairs of the indumentum are not short or elongate branchlets, scarcely aggregated to form
“plumose” but multangulate-dendritic; the stamens are leafy spikes; inflorescence-bracts 0.4–0.6 × 0.1–0.2 mm,
included, not “scarcely exsert” and the ovary is glabrous, densely tomentose with multangulate-dendritic hairs.
not “with a few stellate hairs”. Examination of the Pedicel 0.3–0.5  mm, pubescent with multangulate-
greater number of specimens now available indicates dendritic hairs c. 0.15 mm, glandular hairs absent.
that up to four seeds develop: in Ising’s material the Calyx 2.0–2.5 mm, outside very densely tomentose with
capsules were one-seeded. multangulate-dendritic hairs mostly 0.1–0.2 mm and
fewer to 0.25 mm, inside moderately pubescent with
Selected specimens (70 examined) antrorse glandular hairs 0.06–0.1 mm, especially on
WESTERN AUSTRALIA. Great Victoria Desert IBRA the lobes; calyx-lobes acute, blunt, 0.4–0.5 times as long
Region: Anne Beadell Highway, 37 km E by track of Ilkurkla as tube, 1 × 0.8 mm, sinuses acute. Corolla 5.5–6 mm,
roadhouse, 6.x.2010, R. Davis, J. Jackson & D. Ferguson dull white, striations not recorded, outside moderately
11636 (PERTH, CANB n.v.). SOUTH AUSTRALIA. to densely pubescent with multangulate and forked
North Western: c. 135 km N of Cook, just in main dunes, hairs 0.15–0.25 mm on the lobes and upper tube,
18.viii.1980, D.E. Symon 12233 (AD, NSW). Nullarbor: inside very densely papillose with hairs 0.08–0.1 mm on
2 miles [c. 3 km] S of Maralinga on sand ridge, 10.x.1956, lobes; tube 3.5–4 mm long, markedly widened in upper
N. Forde 620 (AD, CANB); 35 km E of Barton on 0.5 mm, diam. 1.2–1.4 mm at base, and 2.0–2.4 mm at
Transcontinental Railway, 24.x.1970, B. Lay 81 (AD, MEL). limb-base; lobes ovate-elliptic with truncate base, (L:B
Gairdner Torrens: 15 km S of Lake Everard Homestead, 1.7–1.8), 2.0–2.5 × 1.2–1.4 mm, sometimes slightly
20.ix.1972, J.Z. Weber 3202 (AD). Eyre Peninsula: 30 km narrowed at base, apex rounded to bluntly obtuse,
NW of Ceduna, 2 km NW of Koonibba Hill, 4.i.1979, M.D. margin flat. Stamens 5, the longer ones 3.0–3.3 mm
Crisp 4756 (CBG, AD, NSW); 60 km NNE of Minnipa on and the shorter ones 2.5–2.7 mm; filament-bases densely
Yardea road, 18.x.1975, L. Haegi 770 (AD, CBG, NSW); pubescent with simple, forked and sparingly-branched
17 km SSW of Hiltaba on Yantanabie road, 19.x.1975, L. eglandular hairs 0.2–0.3 mm, equally dense abaxially;
Haegi 775 (AD, BIRM, BRI); 20 km N of Karcultaby, upper anthers c. 0.5 × 0.6 mm, lower anthers sometimes
20.x.1975, L. Haegi 794 (AD, CORD, MO); c. 29 km by
slightly smaller, 0.4–0.5 × 0.5–0.6 mm. Ovary globose,
road SW of Yardea Homestead on Minnipa road, 29.x.1978,
L. Haegi 1597 (NSW, AD, CANB, F, HO); 12.4 km directly
0.6 mm diam.; ovules 6–8 in total; style 1.9–2.1 mm, ±
SSE of Mt. Stuart, 11.x.1999, D.E. Murfet & R. Taplin equal to the longer stamens and included by 0.5 mm;
19991011 (AD); 2.9 km directly SSW of Corabinnie Hill, stigma c. 0.15 × 0.3 mm; disc undulate, 0.2–0.3 mm
19.x.2001, S.D. Kenny BS 128-276 (AD168685). high, c. 0.8  mm diam., colour and consistency
unknown. Capsule ± inconspicuous, largely enclosed by
3. Grammosolen archeri Haegi, sp. nov. the tomentose calyx, globose, 2.5–3.0 mm diam. Seeds
Grammosolen sp. Mt Ridley (W.R. Archer 1210911) W. 3–4, c. 2.2 × 1.2 mm; surface-reticulae 0.1–0.15 × 0.05–
Austral. Herb., FloraBase—the Western Australian Flora 0.15 mm. Chromosome number not determined. Fig. 9.
(1998–), https://florabase.dpaw.wa.gov.au/ [accessed 5
June 2019]; CHAH, Austral. Pl. Cens. (2006), https:// Distribution and Ecology. Endemic to Western
biodiversity.org.au/nsl/services [accessed 5 June 2019]. Australia where it is to date known only from the
— Holotype: Western Australia, Eastern Mallee IBRA Eastern Mallee IBRA subregion in the South Western
subregion, [Vicinity of ] Lake Halbert [precise locality Province, in an area 70–90 km inland from the coast
details withheld for conservation reasons], 8 Oct. between Esperance and Cape Arid. The three known

23
L. Haegi Swainsona 33 (2020)

E
D

Fig. 9. Grammosolen archeri.

I A flowering branchlet,
× 4.5; B  leaf, spread, upper
G side, × 7.5; C leaf showing
H orientation and underside,
× 7.5; D flower from above,
A F × 6; E flower from side, × 6;
F corolla lobe, late bud stage,
× 12; G corolla opened out,
with androecium, × 7.5;
H  stamens showing adaxial
dehiscence, × 19; I calyx
K opened out, with pistil, × 9;
J capsule, ×  7.5; K seed,
B C J × 11. (W.R. Archer 2909916).
Illustration by F. James.

populations occur below 200 m altitude in the 300 mm usually closely spaced so as to be overlapping, concealing
to 400 mm mean annual rainfall zone, on disturbed the axis (especially in the more distal parts), and with
sites in deep sandy loam soil and in mallee-heath flowers aggregated into leafy spikes. From G. odgersii it
vegetation (Fig. 3). differs in having five (cf. four) fertile stamens and in
lacking verticillately-branched hairs on the vegetative
Conservation status. Rare and poorly known. Listed parts.
under the schedules of the Western Australian Wildlife
Conservation (Rare Flora) Notice as a Priority 1 species, Variation. No appreciable patterns of variation are
reflecting inadequate knowledge of its occurrence and evident in G. archeri, which is recorded from only three
abundance, because of the small number of collections collections across an area about 100 × 20 km.
and relatively poorly collected area it occurs in.
Derivation of name. Named for William R. Archer,
Diagnostic features. Grammosolen archeri differs from botanical collector and horticulturist of Esperance,
all three other species in its distinctive, small, orbicular- Western Australia, who first discovered this species in
reniform to obcordate leaves and in its much smaller the field and has collected the only known material of
flowers and fruits. It seems closest to G. truncatus and, it.
perhaps somewhat less so, to G. dixonii. It is of similar
appearance, with a dense covering of multangulate and Specimens seen (precise locality details withheld for
multangulate-dendritic hairs on vegetative parts and conservation reasons)
with flowers of very similar appearance (dull white WESTERN AUSTRALIA. Eastern Mallee IBRA subregion:
corolla with ovate-elliptic lobes; the occurrence of [Vicinity of ] Lake Halbert, 17 Feb. 1990, W. Archer 1702903
striations in the tube is expected but yet to be observed). (AD; PERTH, to be distributed); 12.x.1991, W.R. Archer
Like G. dixonii it has a spreading habit and subsessile 1210911 (MEL; also PERTH, HO n.v.); [Vicinity of ] Mt
leaves, while it is similar to G. truncatus in being more Ridley, 13.x.1990, W.R. Archer 13109010 (AD; PERTH,
openly branched and having small inflorescences borne BRI, BM, to be distributed); [Vicinity of ] Mt Buraminya,
on short lateral shoots (though these are sometimes 29.ix.1991, W.R. Archer 2909916 (AD; CANB, MEL n.v.;
elongate in G. archeri). In G. dixonii the leaves are K, NSW, PERTH, to be distributed).

24
Swainsona 33 (2020) Grammosolen (Solanaceae - Anthocercideae) revisited

4. Grammosolen odgersii (F.Muell.) Haegi, comb. nov. Distribution and Ecology. Grammosolen odgersii is
Anthocercis odgersii F.Muell., Fragm. 10 (1876) 19; endemic to southern inland Western Australia where
C.A.Gardner, Enum. Pl. Austral. Occ., Part 3 (1931) it occurs in a few widely scattered populations in the
116; Beard, Descr. Cat. W. Austral. Pl. (1965) 95; Avon Wheatbelt and Coolgardie IBRA regions. The
J.W.Green, Cens. Vasc. Pl. W. Austral. (1981) 92. range falls entirely within the 150 mm to 500 mm
— Cyphanthera odgersii (F.Muell.) Haegi, Telopea 2 mean annual rainfall zone (Fig. 3).
(1981) 177; R.W.Purdie, Symon & Haegi, Fl. Austral.
29 (1982) 27. — Type citation: “Ad fontes Victoriae; Conservation Status. See subspecies below.
Young”. Lectotype (here designated): Victoria
Spring [Queen Victoria Spring, Western Australia, Diagnostic features. Grammosolen odgersii is readily
30°26’S 123°24’E], s.dat., Young s.n. (MEL70945). distinguish­ able from all the other three species in
Isolectotype: MEL70234. the genus in having flowers with only four (not
five) fertile stamens, and in the indumentum on the
Grey-green soft shrub 0.5–2.5 m high with ascending or vegetative parts consisting of very long verticillately
radiating, often arched branches. Branches very densely branched hairs, up to 6.5 mm long on the axes and
lanate-tomentose with verticillately branched eglandular up to 1 mm long on the leaves (cf. multangulate and
hairs 0.3–6.5 mm long and scattered simple glandular multangulate-dendritic hairs no more than 0.25 mm
hairs 0.25–0.5 mm. Leaves ± scattered or often crowded long in G. truncatus and G. archeri and a mixture of
towards the ends of branches, spreading, subsessile; multangulate-dendritic and verticillately branched
lamina broadly to narrowly ovate-elliptic (L:B 1.1–3.8), hairs no more than 1.5 mm long in G. dixonii).
(11–) 12–30 (–35) × 7–13 mm, very densely lanate- Grammosolen odgersii is perhaps closest to G. dixonii
tomentose above and below with verticillately branched but differs further from it in leaf-shape: broadly
hairs 0.3–1.0 mm; base cuneate to attenuate; apex acute to narrowly ovate-elliptic (L:B 1.1–3.8), cf. ovate-
and blunt to obtuse or rounded; margin flat, rarely triangular to very broadly so, sometimes subcordate
slightly inrolled or undulate; midrib obscure above and (L:B (0.8–) 0.9–1.5 (–2.5)) in G. dixonii.
below. Flowers in dense axillary and terminal clusters,
often forming leafy spikes; upper inflorescence-bracts to Variation. Although G. odgersii is known from fewer
5 mm, narrowly elliptic, moderately pubescent adaxially than 30 collections, a geographically correlated,
with oblique to antrorse glandular hairs 0.05–0.25 mm, bimodal pattern of variation is evident, based
and densely lanate-tomentose abaxially (and also near on several characters (Table  5). Plants from four
the apex adaxially) with verticillately branched hairs populations at the western end of the distribution
0.5–2 mm. Pedicel 0.5–2 mm, sparsely to moderately differ from all other populations mainly in having
pubescent with simple glandular hairs 0.05–0.75 mm. longer, proportionally narrower leaves, a longer
Calyx 4–7 mm, pubescent outside in lower half as for indumentum and shorter corolla-lobes. These have
pedicel, in upper half very densely lanate-tomentose been recognised as subsp. occidentalis (Haegi 1981).
with porrect to oblique verticillately branched hairs There are no overlaps in the distributions of the two
0.5–2.5 mm, often entirely obscuring calyx, pubescent subspecies and although subsp. odgersii has a wide
inside with porrect to antrorse glandular hairs 0.05– east-west occurrence, there is no perceptible gradation
0.25 mm; calyx-lobes 2–3 × 1.0–1.7 mm, acute to towards subsp. occidentalis.
acuminate, blunt, sinuses acute, angular. Corolla 5.5–
8.5 mm, white with violet striations, sparsely pubescent Derivation of name. Named in honour of William
outside with glandular hairs 0.05–0.2 mm, very densely Henry Odgers (1828–1881), an under-secretary of the
pubescent inside with papillae 0.05–0.1 mm; tube 4.2– Victorian Chief Secretary’s Department, who supported
6.5 mm, diam. 0.8–1.5 mm at base and 2.5–4 mm at Ferdinand von Mueller (the author of the name) in his
limb base; lobes ovate to broadly ovate with truncate cause of carrying out botanical research as an officer of
base (L:B 1.1–2.3), 1.3–2.5 × 1.1–1.6 mm, fused for the Victorian Government (see protologue, cited above;
0.5–1 mm at base, apex rounded, margin flat but whole Sharr 1978).
lobe often somewhat concave. Stamens 4, the upper
ones 2.0–3.0 mm and the lower ones 1.3–2.2 mm; Vernacular name. Woolly anthocercis (Grieve &
filament-bases pubescent with simple and forked or Blackall 1975); woolly cyphanthera (Canackle &
dendritic eglandular hairs 0.1–0.5 mm and sometimes Moore 2014).
glandular hairs 0.05–0.15 mm; anthers c. 0.8 × 1 mm.
Ovary ovoid, 0.6–1.6 × 0.5–1.2 mm; ovules 5–6 in total;
style 2.5–3 mm; stigma 0.2–0.3 × 0.3–0.4 mm; disc
even to slightly undulate, 0.1–0.3 mm high, bright red. Key to subspecies of G. odgersii
Capsule inconspicuous, embedded in dense, long hairs 1. Leaves with L:B of 1.1–2.0 and 20 mm or less in length;
of calyx and axis, ellipsoid to ovoid or broadly so, 3–5 corolla lobes 2.0–2.5 mm . . . . . 4a. subsp. odgersii
× 2–4 mm. Seeds 1–4, 2.8–3.4 × 1.1–1.4 mm; surface 1: Leaves with L:B of 2.3–3.8 and 18–35 mm in length;
reticulae 0.2–0.4 × 0.2–0.3 mm. Gametic chromosome corolla lobes 1.3–1.8 mm . . . 4b. subsp. occidentalis
number: n=30 (Chinnock 3100). Figs 10 & 11.

25
L. Haegi Swainsona 33 (2020)

Table 5. Morphological differences between the two subspecies of G. odgersii.

Character subsp. odgersii subsp. occidentalis

Habit Low shrub 0.5–1m Shrub 1–2.5 m

Length of indumentum 0.2–2.0 mm 0.5–3.0 mm

Length to breadth ratio of leaves (1.1–) 1.4–1.8 (–2.0) (2.3–) 2.5–3.3 (–3.8)

Leaf length 11–20 mm 18–35 mm

Pedicel length 0.5–1.5 mm 0.9–2.0 mm

Corolla-lobe length 2.0–2.5 mm 1.3–1.8 mm

Pubescence at filament bases Simple glandular and eglandular hairs Simple and dendritic eglandular hairs

4a. Grammosolen odgersii subsp. odgersii Specimens seen (all cited)

Anthocercis odgersii F.Muell.: F.Muell., Syst. Cens. WESTERN AUSTRALIA. Coolgardie IBRA Region:
Austral. Pl. (1882) 96; F.Muell., Second Syst. Census. Victoria Desert, Camp 58, Elder Exploring Expedition,
Austral. Pl. (1889) 163 (where distribution given 25 km NE of Streich Mound, 21.ix.1891, R. Helms s.n.
erroneously as Northern Australia (“N.A.”); Ewart, (AD96414102, AD97604848, MEL70235, MEL70946,
Vict. Naturalist 23(9) (1907) 155, p.p. excluding NSW123728, NSW123729); 35  km W of Plumridge
Cowcowing material; Ewart, Vict. Naturalist 24(1) Lakes, 8.5 km WNW of Salt Creek airstrip, 15.ix.1979, J.
(1907) 12–13, p.p. excluding Koch specimen; Ewart, Taylor 498, M. Crisp & R. Jackson (CBG, NSW); 89.9 km
Vict. Naturalist 24(6) (1907) 56, p.p. as to Victoria WSW of Coolgardie, just E of Boorabbin, 18.ix.1976,
Springs material; B.J.Grieve & W.E.Blackall, How R.J. Chinnock 3098, 3099 & 3100 (AD); 82.8 km WSW
to Know W.A. Wildfl. 4 (1975) 610, [110], p.p., as of Coolgardie, near Boondi, 18.viii.1977, R.J. Chinnock
to “Leaves orbicular-ovate” and ref. to Coolgardie 3674 (AD); 20.5 km E of Zanthus along Transcontinental
District. Railway line, 17.ix.1979, M.D. Crisp 5881, J. Taylor & R.
Cyphanthera odgersii (F.Muell.) Haegi subsp. odgersii, Jackson (PERTH; CBG, NSW n.v.); c. 90 km by road SW of
Coolgardie on Great Eastern Highway, 4.vii.1979, L. Haegi
Telopea 2 (1981) 177.
1780 (NSW, BRI, F, PERTH); Near Warangering, Camp
Chloanthes drummondii R.Helms ex Ewart, Vict. 81, Elder Exploring Expedition, 14.xi.1891, R. Helms s.n.
Naturalist 24 (1907) 12, as ‘drummondi’, nom. inval. (AD96414101, AD96430080, MEL70236); Goddard (now
Ponton) Creek E of Zanthus, 3.x.1956, R.D. Royce 5570
Shrub 0.5–1 m high with radiating often arched (PERTH); 20 km E of Zanthus, 2.ix.1968, P.G. Wilson 7629
(PERTH, AD); 20.5 km E of Zanthus at Goddard (now
branches. Branch-indumentum 0.3–2.0 mm. Leaf-
Ponton) Creek, 9.xi.1976, E. Wittwer 1977 (PERTH).
lamina ovate-elliptic to broadly so (L:B 1.3–2.2), 11–20
× 7–11 mm, with indumentum 0.3–0.9 mm. Corolla- 4b. Grammosolen odgersii subsp. occidentalis
lobes ovate to broadly ovate (L:B 1.3–2.3), 2.0–2.5 × (Haegi) Haegi, comb. nov.
1.0–1.5 mm. Filament-bases pubescent with simple Cyphanthera odgersii (F.Muell.) Haegi subsp.
eglandular and glandular hairs. Gametic chromosome occidentalis Haegi, Telopea 2 (1981) 178; R.W.Purdie,
number n=30 (Chinnock 3100). Fig. 10. Symon & Haegi, Fl. Austral. 29 (1982) 28. — Type:
“Western Australia, Avon District” [precise locality
Distribution and ecology. Subspecies odgersii occurs in details withheld for conservation reasons], 23.ix.1976,
Western Australia over a wide area from the far western L. Haegi 1100. Holotype: PERTH. Isotypes: AD,
Great Victoria Desert westwards to near Southern CANB, K, MO.
Cross, in the 150  mm to 500  mm rainfall zone. It is Anthocercis odgersii F.Muell.: Ewart, Vict. Naturalist
a rarely encountered plant which occurs in widely 23(9) (1907) 155, p.p. as to Cowcowing material;
scattered populations on open sites in mallee, on sand Ewart, Vict. Naturalist 24(1) (1907) 12–13, p.p.
dunes and sandplain, often following fire. Flowering as to Koch specimen; Ewart, Vict. Naturalist 24(6)
has been recorded from August to October and fruiting (1907) 56, p.p. as to Koch specimen; B.J.Grieve &
in September and October (Fig. 3). W.E.Blackall, How to know W.A. Wildfl. 4 (1975)
610, [110], p.p. as to “linear-oblong”, illustration and
Conservation status. Uncommon to rare but not reference to Avon District.
threatened.
Erect shrub 1–2.5 m high with ascending branches.
Variation. Little variation is evident among the small Branch indumentum 1.0–6.5 mm. Leaf-lamina
number of specimens of this species collected. ovate-elliptic to narrowly so (L:B 2.3–3.8), 18–35 ×

26
Swainsona 33 (2020) Grammosolen (Solanaceae - Anthocercideae) revisited

C
F
A Fig. 10. Grammosolen od­
ger­sii subsp. odgersii.
A  flowering branchlet, × 1;
B flower, × 3; C corolla of
advanced bud opened out,
× 5; D opened calyx and pistil
of same, × 5; E capsule, × 5;
F seed, × 10. (Chinnock 3100).
D Illustration by M. Perkins.

C
B

D
Fig. 11. Grammosolen od­
ger­sii subsp. occidentalis.
A  flowering branchlet, × 1;
B flower, × 4; C corolla of
advanced bud opened out,
× 5; D opened calyx and pistil
of same, × 5; E capsule, × 5;
A E F  seed, × 10. (Haegi 1100).
Illustration by M. Perkins.

27
L. Haegi Swainsona 33 (2020)

8–13 mm, with indumentum 0.5–1.0 mm. Corolla- Powell (Sydney), Philip Short (Melbourne) and Bob
lobes broadly ovate (L:B 1.1–1.5), 1.3–1.8 × 1.0– Chinnock, Bill Barker, Robyn Barker and Ed Biffin
1.6 mm. Filament-bases pubescent with eglandular hairs (Adelaide) are sincerely thanked for their beneficial
only. Chromosome number not determined. Fig. 11. discussions and in some cases assistance in the field.
Likewise, I acknowledge provision of several helpful
Distribution and ecology. Grammosolen odgersii suggestions by an anonymous referee – these have been
subsp. occidentalis is known from a few small, scattered incorporated in an amended version of the paper. Kathy
populations in the Avon Wheatbelt and adjacent Stephens and Marie Taylor (Sydney) provided excellent
Coolgardie IBRA regions in Western Australia, in technical assistance; Margaret Perkins and Rob Roden
association with salt lakes forming parts of ancient prepared most of the drawings while temporarily
shallow drainage systems. Plants occur in deep sandy to employed at the National Herbarium of New South
sandy loam soils, often on disturbed sites (e.g. following Wales. Fiona James, as a student in botanical illustration
fire) in mallee/woodland (Fig. 3). at the State Herbarium of South Australia, prepared the
excellent plate of Grammosolen archeri. My thanks go
Conservation status. Listed in Western Australia,
(as Cyphanthera odgersii subsp. occidentalis), applying to Bill Barker for preparing the distribution map using
international criteria, as Critically Endangered. For QGIS and useful GIS layers with point data I provided
this reason only generalised locality information is from the ALA download. My deep gratitude goes to my
provided here. Grammosolen odgersii subsp. occidentalis wife Rosalie Haegi for her support, encouragement and
has been the subject of intensive recovery work unstinting help throughout the life of the project, not
protecting existing populations, propagation of plants least in assisting with fieldwork.
for translocation and collection of seed for long-term
germplasm storage (Canackle & Moore 2014). I gratefully acknowledge the loan of specimens from,
and / or assistance received while visiting the following
Variation. Little variation is evident among the small herbaria: BM, BRI, CANB, CBG, K, MEL, NE, NSW,
number of specimens of this species collected. PERTH.

Specimens seen (all cited — precise locality details

withheld for conservation reasons) References
WESTERN AUSTRALIA. Avon Wheatbelt IBRA Region:
Sep. 1904, M. Koch s.n. (no. 1104) (AD, K, MEL70237, Barnard, C. (1949). Microsporogenesis, macrosporogenesis
MEL70238, MEL70944, NSW, PERTH); Oct. 1939, and development of the macrogametophyte and seeds
C. Gardner s.n. (PERTH); 3.xii.1946, C. Gardner 8507 of Duboisia leichhardtii FvM and D. myoporoides R.Br.
(PERTH); 19.x.1990, F.H. & M.P. Mollemans 3470 Australian Journal of Scientific Research B 2: 241–248, plates
(PERTH); 28.x.1997, J.A. Cochrane 2460 (PERTH); 1–3.
12.ix.2007, J.M. Collins 280 (PERTH); 12.ix.2007, L. Duffy Canackle, L. & Moore, N. (2014). All aboard for conservation.
LD140 (PERTH). Coolgardie IBRA Region: June 1990, Landscope 29(4): 38–41.
F.H. & M.P. Mollemans 2981 (AD, PERTH); 13.ix.1990, Clarkson, J.J., Knapp, S., Garcia, V.F., Olmstead, R.G., Leitch,
F.H. & M.P. Mollemans 3114 (AD, PERTH). A.R. & Chase, M.W. (2004). Phylogenetic relationships in
Nicotiana (Solanaceae) inferred from multiple plastid DNA
regions. Molecular Phylogenetics and Evolution 33: 75–90.
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Australian species of Duboisia R.Br. (Solanaceae). Journal of
The study described here commenced while the author the Adelaide Botanic Gardens 16: 27–31.
was employed at the Waite Agricultural Research Darlington, C.D. & La Cour, L.F. (1970). The Handling of
institute, University of Adelaide during 1976 and 1977 Chromosomes, 5th edn, 2nd printing. (George Allen &
and a PhD student at Flinders University of South Unwin: London).
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1982). During this time it was supported by grants from Pharmacognosy 29(4): 263–267.
the Australian Biological Resources Study, for which I Garcia, V.F. & Olmstead, R.G. (2003). Phylogenetics of Tribe
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work recently recommenced in Adelaide and I thank sequence data. Systematic Botany 28(3): 609–615.
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Garcia, V.F. & Collier, S.M. (2008). A molecular phylogeny
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Koeltz Books, Koenigstein, Germany). Flora of Australia Vol. 29, vii, 208 pp. (Australian
Ikenaga, T., Abe, M., Itakura, A. & Ohashi, H. (1979). Alkaloid Government Publishing Service: Canberra).
contents in the leaves of artificial F1 hybrids between Sharr, F.A. (1978). Western Australian Plant names and their
Duboisia myoporoides and D. leichhardtii. Planta Medica 35: meanings – a glossary, xxxvi, 228 pp. (University of Western
51–55. Australia Press: Perth).

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