Swainsona

Government
of South Australia

Board of the
Botanic Gardens and
State Herbarium

Swainsona

formerly Journal of the Adelaide Botanic Gardens

ISSN 2206-1649 (Print) • ISSN 2206-1657 (Online) •flora.sa.gov.au/swainsona

Guidelines to authors

Swainsona, formerly the Journal of the Adelaide Botanic Gardens, is an
open-access, peer-reviewed journal published by the State Herbarium of
South Australia on behalf of the Board of the Botanic Gardens and State
Herbarium (Adelaide, South Australia). The journal is available online at
flora.sa.gov.au/swainsona and through JSTOR.

Editorial Committee: Robyn Barker, Ed Biffin, Tony Kanellos, Jurgen
Kellermann (Editor), Tucy Sutherland & Michelle Waycott.

Scope

Swainsona accepts papers in the following categories:

• plant systematics, taxonomy and nomenclature of vascular plants,
bryophytes, fungi, lichens and algae

• evolution and biogeography

• descriptive plant morphology and anatomy

• plant biology, genetics, and ecology

• obituaries, biography and history

• bibliographic studies, book reviews

• botanical illustrations.

Focus of the j ournal is on taxa from the Australasian region, but manuscripts
from other regions of the world may be considered.

Descriptions of single taxa are accepted, as well as revisions and full length
papers. There is no page limit, but authors should inform the Editor if
they intend to submit larger manuscripts. Larger manuscripts may also
be submitted for the Swainsona Supplement series or be published in a
separate volume of the journal. All submitted papers are refereed by two
referees.

Submission

Papers will be considered only on assurance that they are not being
considered by another publication and will not be withdrawn without
consent of the Editorial Committee.

Manuscripts should be submitted to the Editor electronically as a Word
document (.doc, .docx) or in Rich Text Format (.rtf). Consult the Editor
where uncertain on style. The Editorial Committee may return a submitted
manuscript to bring it to standard.

Address for submissions:

Dr Jurgen Kellermann
Editor, Swainsona

Botanic Gardens and State Herbarium
Hackney Road, Adelaide, S.A. 5000, Australia
E-mail: juergen. kellermann @sa.gov. au

Style

Authors should follow the format in the most recent volume, Yol. 31,
noting title, authorship, abstract, heading, references, in-text publication
referencing, etc. Detailled Guidelines will be available on the journal’s
website.

Italicise in the text the following: scientific names, the short-hand de¬
signation of a herbarium collection (see below), journal and book titles in
text, references and abbreviations, such as et al., auct. non, nom inval. Do
not italicise in text and References standard abbreviations of Latin origin,
such as: c., e.g., etc. Format in synonymy differs (see below).

Keys. Indented keys are preferred to bracketed keys.

Synonymy. Full lists of synonyms and invalid names, with references to
application in significant historical publications, are encouraged. It is
recommended that:

• nomenclatural (homotypic) synonyms be placed chronologically in same
paragraph, synonyms separated by a period and em-dash; taxonomic
(heterotypic) synonyms, then misapplications and invalid names, follow
in separate paragraphs

• type details be dealt with at the end of each paragraph grouping
nomenclatural synonyms, or in a separate section titled Typification.

• authorities be abbreviated according to Brummit & Powell’s (1992)
Authors of Plant Names, and its updates in the International Plant
Names Index ( www.ipni.org )

• references italicised and have consistent abbreviations of book and
journal titles. Books titles are to be abbreviated following Stafleu et
al. (1976—), TL-2 and TL-2 Suppl. Journal titles are to follow Bridson
& Smith (1991), B-P-H/S-, abbreviations are available online through
IPNI; citation of volume, date and pagination have to be consistent,
and either in the form “Benth., FI. Austral. 4: 111 (1868)” or “Benth.,
FI. Austral. 4 (1868) 111”.

Citation of specimens. In general text use the abbreviated method
(collector name and unique collection number, italicised, e.g. J.Smith
1234) or, where lacking a number, collector name and the date and/or
herbarium sheet identifier, italicised (e.g . J.Smith AD23456). Full citation
of specimens should follow the treatment of each taxon, with a selection
of 10-30 specimens, grouped by country, state and region, ordered either
alphabetically by collector and number or north to south, west to east by
locality.

Voucher specimens need to be lodged in recognised herbaria. Gene
sequences must have a GenBank accession number and specimen voucher
number.

Phrase names. A new phrase name will only be agreed if publication of a
formal Latin name is not practical, if no alternative exists, and if there are
strong (e.g. conservation) reasons for establishing one. The format should
follow that set by Australian herbaria: B. Barker, Austral. Syst. Bot. Soc.
Nsltr 122: 12-13 (2005).

References. See most recent volume for reference style. Cite Web-pages
and Databases in text in abbreviated form with date, as with bibliographic
References; in References, cite Web-pages in the form: Author or
Organisation (Date). Title. Web-address [accessed: date]; cite Databases
in form requested by the owner, or by: Abridged name (Date). Title.
Institution. Web-address [accessed: date].

Publication & copyright

Following the reviewing and editing process, new articles are published
online as soon as possible. The online publication date is indicated on
the first page of each article and is considered the valid publication date
according to the International Code of Nomenclature. Hardcopy, collating
all articles published during a calendar year, is printed at the start of the
following year. The printed journal is distributed to exchange partners and
is available for sale (contact the Editor for more information).

Publication in Swainsona is conditional on licence being given to publish
both in a printed journal, electronically on the internet, or in other form
approved by the Crown through the cited copyright holder: Board of the
Botanic Gardens and State Herbarium (Adelaide, South Australia).

Government State Herbarium

of South Australia of South Australia

Swainsona

»!*

ISSN 2206-1649 (Print) • ISSN 2206-1657 (Online) •flora.sa.gov.au/swainsona

Volume 31 (2017/18)

Contents

Leptecophylla in Tasmania: a reassessment of four species

SJ. Jorman & G. Kantvilas .1

A new species of small black disc fungi, Smardaea australis (Pezizales, Pyronemataceae),
is described from Australia

PS. Catcheside, S. Qaraghuli & D.E.A. Catcheside .17

Tephromela baudiniana sp. nov. (lichenised Ascomycetes) from Kangaroo Island

G. Kantvilas &J.A. Elix .27

Two species of Bacidia De Not. with pruinose apothecia from Kangaroo Island

G. Kantvilas . 31

Goodenia asteriscus (Goodeniaceae), a new arid zone species from north-western South
Australia and eastern Western Australia

P.J. Lang & RJ.-P. Davies .37

Pertusaria crassilabra Mull. Arg. - a reinstated name for an Australasian lichen

G Kantvilas .45

The type of Sturt pea found

4.5. George .49

Micarea kartana sp. nov. (lichenised Ascomycetes) from Kangaroo Island, South Australia

G. Kantvilas .55

Notes on the taxonomy of Australian Lindernia subg. Didymadenia (Linderniaceae)

W.R. Barker .59

Antrelloides atroceracea, a new genus and species in the Pezizaceae (Pezizales) from
Australia

P.S. Catcheside & D.E.A. Catcheside .81

The Ptilotus murrayi species group: synonymisation of P. petiolatus under P. murrayi and
description of the new Western Australian species P. unguiculatus (Amaranthaceae)

T.A. Hammer .93

CORRIGENDUM to: A new species of small black disc fungi, Smardaea australis (Pezizales,
Pyronemataceae), is described from Australia

P.S. Catcheside & D.E.A. Catcheside .101

New taxa, names and combinations

Swainsona 31 ( 2017 / 18 )

Antrelloides P.S.Catches. & D.E.A.Catches.

Antrelloides atroceracea P.S.Catches. & D.E.A.Catches.

Bacidia brigitteae Kantvilas.

Goodenia asteriscus P.J.Lang..

Leptecophylla oxycedrus (Labill.) Jarman .

Leptecophylla parvifolia (R.Br.) Jarman.

Lindernia subg. Didymadenia W.R.Barker.

Lindernia sect. Didymadenia (W.R.Barker) W.R.Barker
Lindernia sect. Hemiarrhena (Benth.) W.R.Barker . . ,

Lindernia sect. Heterandrae W.R.Barker.

Lindernia sect. Prolatae W.R.Barker.

Lindernia sect. Scapigerae W.R.Barker.

Lindernia acrandra W.R.Barker.

Lindernia atrata W.R.Barker.

Lindernia brennanii W.R.Barker.

Lindernia calliandra W.R.Barker.

Lindernia cyanoplectra W.R.Barker.

Lindernia cf/eryf/ira W.R.Barker..

Lindernia dunlopii W.R.Barker.

Lindernia enypniastina W.R.Barker..

Lindernia /ei/coc/iroc? W.R.Barker ..

Lindernia lucrusmiana W.R.Barker..

Lindernia mitrasacmoides (O.Schwarz) W.R.Barker . ,

Lindernia murfef/ana W.R.Barker..

Lindernia petrensis W.R.Barker.

Lindernia porphyrodinea W.R.Barker & M.D.Barrett . ,

Lindernia prolata W.R.Barker..

Linderniapronanthera W.R.Barker ..

Lindernia pustulosa W.R.Barker.

Lindernia robyniae W.R.Barker..

Lindernia scopularis W.R.Barker.

Lindernia scutellata W.R.Barker.

Lindernia thyridostoma W.R.Barker.

Lindernia tiwiensis W.R.Barker.

Lindernia venustula W.R.Barker.

Micarea kartana Kantvilas & Coppins..

Ptilotus unguiculatus T.Hammer.

Smardaea australis P.S.Catches. & D.E.A.Catches. . . ,
Tephromela baudiniana Kantvilas & Elix.

28 Sep. 2018.82

28 Sep. 2018.82

15 June 2017.34

24 Nov. 2017.37

15 June 2017.1

15 June 2017.7

. 2 May 2018.61

. 2 May 2018.64

. 2 May 2018.71

. 2 May 2018.70

. 2 May 2018.61

. 2 May 2018.67

. 2 May 2018.77

. 2 May 2018.62

. 2 May 2018.74

. 2 May 2018.78

. 2 May 2018.63

. 2 May 2018.63

. 2 May 2018.74

. 2 May 2018.77

. 2 May 2018.69

. 2 May 2018.76

. 2 May 2018.71

. 2 May 2018.63

. 2 May 2018.65

. 2 May 2018.67

. 2 May 2018.62

. 2 May 2018.77

. 2 May 2018.72

. 2 May 2018.78

. 2 May 2018.65

. 2 May 2018.71

. 2 May 2018.70

. 2 May 2018.68

. 2 May 2018.76

11 Apr. 2018.56

28 Sep. 2018.96

15 June 2017.19

15 June 2017.28

The online publication date is indicated before the page number.

w

Swainsona 31:1-16 (2017)

© 2017 Board of the Botanic Gardens & State Herbarium (Adelaide, South Australia) ©0

Leptecophylla in Tasmania: a reassessment of four species

SJ. Jarman & G. Kantvilas

Tasmanian Herbarium, PO Box 5058, UTAS LPO, Sandy Bay, Tasmania 7005
Email: Gintaras.Kantvilas@tmag.tas.gov.au

Abstract: The occurrence of Leptecophylla juniperina (J.R.Forst. & G.Forst.) C.M.Weiller in Tasmania is
reviewed. Two subspecies of this taxon are re-instated to specific rank: L.oxycedrus (Labill.) Jarman
comb. nov. and L. parvifolia (R.Br.) Jarman comb. nov. Leptecophylla juniperina itself is excluded from the
Tasmanian flora. Tasmanian plants previously identified as L.juniperina are mostly either L.oxycedrus
or the newly described L. pogonocalyx subsp. decipiens Jarman subsp. nov. An identification key is
provided for Tasmanian species of Leptecophylla.

Keywords: Leptecophylla, taxonomy, change of status, new subspecies, Tasmania

Introduction

The genus Leptecophylla was first described by Weiller
(1999) for a group of 12 closely related species formerly
included in Cyathodes Labill. or Styphelia Sm. in the
Epacridaceae, a family which has since been submerged
in the Ericaceae (Kron et al. 2002). The transfer of
these species to the new genus serves to highlight
the close affinities within the group and is widely
accepted by botanists. However, within Leptecophylla ,
relationships among several taxa are more problematic.
This present study focusses on the taxa included
in Weiller s (1999) study as L. juniperina subsp.
juniperina , L. juniperina subsp. oxycedrus , L. juniperina
subsp. parvifolia and L. pogonocalyx. Our treatment
proposes major nomenclatural changes, with the re¬
instatement to specific rank of both of the subspecies
oxycedrus and parvifolia , the exclusion of L. juniperina
from the Tasmanian flora, and the description of a new
subspecies of L. pogonocalyx. A detailed discussion of
the evidence supporting these changes is provided.

Materials

Taxonomic interpretation of all Tasmanian species was
based primarily on fresh material, although the large
collection of Leptecophylla specimens at the Tasmanian
Herbarium (HO; over 800 sheets) was also consulted
extensively. The New Zealand species, L. juniperina ,
was examined only as dried specimens held at HO
or on loan from the National Herbarium of Victoria
(MEL), Auckland Museum Herbarium (AK), Allan
Herbarium, Landcare Research Manaaki Whenua,
Lincoln (CHR), Museum of New Zealand Te Papa
Tongarewa Herbarium, Wellington (WELT), and the
Natural History Museum, London (BM).

Type material of L. juniperina and L. parvifolia was
examined via loans from BM. Labillardiere’s specimens
of L. oxycedrus held at Florence were viewed there by
one of us (GK), as well as via digital images supplied by
the Herbarium Universitatis Florentinae (FI-W). Other
historical specimens collected in the late 18 th and early
19 th centuries were viewed as digital images on JSTOR
Global Plants or on the individual websites of herbaria
in Britain, Europe or America.

Taxonomic discussion

1. Leptecophylla oxycedrus (Labill.) Jarman, comb. nov.

Styphelia oxycedrus Labill., Nov. Holl. Pl. 1: 49,
t. 69 (1805). — Type: in capite Van-Diemen,
J.J. Labillardiere (holo: FI009077!). Cyathodes oxycedrus
(Labill.) R.Br., Prodr. 540 (1810).

For detailed synonomy, see Weiller (1999, p. 205),
under Leptecophylla juniperina subsp. oxycedrus.

The first known European collections of Leptecophylla
oxycedrus (Fig. 1) were made by Jacques Julien de
Labillardiere, naturalist with Bruni D’Entrecasteaux’s
expedition in the Recherche and Esperance which sailed
from France in 1791 in search of the missing navigator
Jean-Francois de La Perouse. The expedition visited
Van Diemens Land (Tasmania) in April-May 1792,
and January-February 1793 (Labillardiere 1800).
Material from the expedition was widely distributed,
with the main plant collections being held in the Webb
Herbarium in Florence.

Labillardiere’s precise collecting sites are not known,
with the location on all his specimens simply indicating
‘In capite Van-Diemen’ (Van Diemens Land). However,
his stay in Tasmania was confined to the south-east and
south of the island, with the main anchorages being

Published online: 15 June 2017 • flora.sa.gov.au/swainsona

ISSN 2206-1649 (Print) • ISSN 2206-1657 (Online)

SJ. Jarman & G. Kantvilas

Swainsona 31 (2017)

Ex Herb. Labillardicre.

i L_ fy//4,u (Zk- Z't TUt&Z- _ j|

Herbarium

Umversitatis Florentinae

FOTOTECA

n.- xtt8

Data 4# 0} CO

Herb. Webbianum.

Fig. 1 . Leptecophylla oxycedrus, collected in Van Diemens Land (Tasmania) by J.J. Labillardiere in 1792/1793. (Image courtesy of
Herbarium Universitatis Florentinae.) Inset: magnified segment of the main image.

2

Swainsona 31 (2017)

in Recherche Bay in the far south. The ships sailed
through the D’Entrecasteaux Channel and around the
northern tip of Bruny Island, with landings made in
several places (Labillardiere 1800). Based on the known
distribution of Leptecophylla oxycedrus , the species
is most likely to have been collected from around
Recherche Bay, where it would have been one of the
first species encountered after landfall in April 1792.
The first description of the plant appeared in 1805 in
Labillardiere’s Novae Hollandiae Plantarum Specimen.

Leptecophylla oxycedrus was originally classified
in the genus Styphelia (Labillardiere 1805) and
later in Cyathodes (Brown 1810). It remained as
a distinct species until 1868, when Bentham, in
Flora Australiensis , synonymised it with Cyathodes
acerosa R.Br. (= Leptecophylla juniperina (J.R.Forst.
& G.Forst.) C.M.Weiller), a New Zealand species.
Bentham’s interpretation was followed by many
botanists (e.g. Rodway 1903 and Curtis 1963 dealing
with the Tasmanian flora; Ewart 1930, Willis 1973 and
Albrecht 1996 with the Victorian flora; Cheeseman
1906, 1925 and Allan 1961 with the New Zealand
flora). A dissenting view was held by Sleumer (1963),
who retained the two separately within the genus
Styphelia. In the most recent taxonomic treatment of
Leptecophylla (Weiller 1999), the two taxa, together
with L. parvifolia , are given subspecific rank within
L. juniperina and all three subspecies are included in
the Tasmanian flora.

Separation of L. oxycedrus from L. juniperina

Evidence from both floral and vegetative characters
indicates that Leptecophylla oxycedrus should retain
its original status as a distinct species, separate from

Leptecophylla in Tasmania: a reassessment of four species

L. juniperina. Weiller (1999) comments that all species
in Leptecophylla are apparently functionally dioecious’,
and refers to the presence of ‘apparently hermaphrodite
and male-sterile flowers on separate plants, with only
male-sterile flowers setting fruit’. In L. oxycedrus ,
dioecy is clearly evident in the dimorphic flowers, with
separate plants producing either flowers with large,
pollen-producing anthers or small empty anthers.
Typically, the corolla tube of pollen-producing flowers
is much longer than in female plants (Figs 2, 3) and is
exserted well beyond the calyx. The lobes are almost
always less than half the length of the tube and often
less than one-third to one-quarter. In shape, the tube
is slightly barrel-shaped (Fig. 3) or narrowly urceolate.
In dried material, the tube often appears cylindrical
to very narrowly campanulate. In female flowers, the
length of the tube is shorter, but still clearly remains
exserted beyond the calyx.

Dioecy occurs in L. juniperina (Fig. 4; Gardner 2011)
but is not as obvious as in L. oxycedrus because of the
smaller size difference between pollen-producing and
female flowers. The corolla tube is shorter than in
L. oxycedrus (cf. Figs 3, 4) and, at the shorter end of
the range, can be equal to or less than the length of the
calyx, a condition not encountered in L. oxycedrus.

Critical vegetative characters that separate L. oxycedrus
from L. juniperina are related to the seasonal production
of leaves and to leaf shape and venation. In the genus as
a whole, leaves are produced in a single flush each year.
They are pre-formed in an annual ‘parcel’ enclosed by
bracts which are brown, scarious and increase in size
from the minute lowermost bracts up to the last large
bracts that reach approximately the length of the first

Fig. 2. Leptecophylla oxycedrus from Recherche Bay, Tasmania, the most likely Type locality of J.J. Labillardiere's species collected
in 1792/93. A Pollen-producing flowers; B female flowers with poorly formed anthers; C general aspect of the plant. Scale = 5 mm.

3

SJ. Jarman & G. Kantvilas

Swainsona 31 (2017)

Fig. 3. Flowers of Leptecophylla oxycedrus from Recherche Bay. A Pollen-producing flowers from three different plants; B corolla
of flowers shown in A. C Female flowers from three different plants; D corolla of flowers shown in C. Scale = 5 mm.

Fig. 4. Flowers of Leptecophylla juniperina. A Reconstituted flower from dried material (AK258766; dissection undertaken with
permission); B corolla from A. C Reconstituted flower from FI0583393; D corolla from C. E Partially opened flower on the lectotype,
photographed from BM000797781. Scale = 5 mm.

leaves (e.g. Fig. 5). There is a zone between the last
totally scarious bract and the first properly formed leaf
where there are a number of ‘transition’ leaves that differ
in shape from both the bracts and the properly formed
leaves. They have abaxial venation that is composed
of finely branched veins, which, although essentially
parallel, do not have the distinctly striate appearance
seen in typical leaves. Much of the ‘transition’ leaf
comprises green photosynthetic tissue.

In the ‘transition’ leaves of Leptecophylla oxycedrus , the
hyaline or scarious part of the margin towards the tip
is quite broad but rolls towards the undersurface as the
young shoot expands. As a result, the transition leaves
more-or-less resemble the typical leaves in outline
(Figs 6A, 6B) and blend inconspicuously among them
(Fig. 7A). They are most easily recognised by checking
the leaf undersurface for the rolled margin in the upper
part of the leaf, and the finely branched venation

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Swainsona 31 (2017)

Leptecophylla in Tasmania: a reassessment of four species

Fig. 5. Seasonal production of leaves in Leptecophylla. Scale
= 5 mm.

(Fig. 6A). Except for a few tiny bracts at the very base
of the branchlets, all of the bracts and many of the
transition leaves fall by the time the branchlet reaches its
ultimate size for that years growth. By the second year
of growth, although some of the uppermost transition
leaves may remain, they are uncommon and essentially
indistinguishable from the fully formed mature leaves
when viewed from their upper surface.

In the transition leaves of L. juniperina, the hyaline or
scarious portion towards the apex is quite narrow and is
not or only scarcely rolled under, giving the transition
leaves an oblanceolate shape (Figs 6C, 7B). The finely
branched venation of these leaves is clearly visible on
the undersurface (Fig. 6C). The ‘transition’ leaves
persist and are held across several years’ growth (Fig. 7B)
and, because of their different shape and venation, and
their frequency and persistence, they give a subtle but
distinctive aspect to the plant. There is also a difference
in the shape of normal leaves, which tend to be more
linear in L. juniperina (Fig. 6D) than in L. oxycedrus
(Fig. 6B). The tendency for the major veins to branch,
particularly towards the leaf apex, also persists into the
true leaves to a greater extent in L. juniperina than in
L. oxycedrus.

The branching venation on the underleaf of
L. juniperina has been commented upon by earlier
botanists, including Robert Brown, who, in 1810,
observed that the Tasmanian Cyathodes {Leptecophylla)
oxycedrus differed from the New Zealand C. acerosa
{Leptecophylla juniperina ) in the branching of the
veins towards the leaf extremities. Hooker (1853), in
the Flora of New Zealand , makes the comment about
C. acerosa\

‘There is a tendency in the leaf (very variable in
amount) to become broader towards the tip, whence
the outer nerves branch to supply the increased
surface, which I do not observe in the following
species [C. oxycedrus\ ; this character Mr Brown
detected, and applied in distinguishing them.’

These vegetative differences in venation, and shape and
level of persistence of transition leaves are accompanied
by important floral differences between the two
species (see above), which have proved to be consistent
wherever the appropriate floral stages have been
available for checking.

Our assessment that L. juniperina and L. oxycedrus
are separate species is not novel, with botanists such
as Brown (1810), Sprengel (1824), de Candolle
(1838), Hooker (1853, 1859) and, more recently,
Sleumer (1963), treating the two separately, albeit
acknowledging their close affinities.

Descriptions of L. oxycedrus are given in Ewart (1930,
as Cyathodes acerosa R.Br.) and Albrecht (1996, as
Cyathodes juniperina (J.R.Forst. & G.Forst.) Druce),
and are not repeated here.

Notes. In recent works (Weiller 1999; Crowden &
Duretto 2013), the presence of sparse, rigid hairs
on the corolla of L. oxycedrus has been used as a key
identification character. However, corolla hairiness does
not appear to be a reliable taxonomic character in this
species and its close relatives.

Labillardiere (1805) described the corolla as scarcely
hairy and the hairs on the corolla limb as rare (‘corollae
limbo ... vix piloso’, and ‘Corollae limbus pilis raris’).
In the illustration accompanying Labillardiere’s work,
it is difficult to see any sign of hairs on the corolla,
although admittedly the floral drawing is rather small.
Labillardiere himself is unlikely to have seen many
flowers, either on a single plant or across the population,
because his visits to Tasmania were well outside the peak
flowering times for L. oxycedrus (August-October). We
have collected flowering material from the area most
likely to be the Type locality of Labillardiere’s specimens
(Recherche Bay), where plants are still plentiful. In
an examination (dissecting microscope) of 60 plants
(three flowers/plant) encompassing two populations at
Recherche Bay, approximately two-thirds of the plants
were found to have a glabrous corolla. The remaining
third had hairs present on the corolla, but these were
sometimes as few as one or two. Hairs were mostly in

5

SJ. Jarman & G. Kantvilas

Swainsona 31 (2017)

Fig. 6. Leaves of Leptecophylla oxycedrus from Recherche Bay (A, B) and L.juniperina from New Zealand (C, D, Auckland area,
HO504556). A, C Transition leaf, showing shape and venation on the upper and lower surface. B, D Typical leaf from mid-season's
growth, showing shape and venation on upper and lower surface. Scale = 5 mm.

Fig. 7. Leptecophylla oxycedrus (A) from Recherche Bay, the probable Type locality, and L. juniperina (B) from New Zealand
(Auckland area, HO504556). Arrows show the position and/or shape of transition leaves (fallen or scarcely discernible among the
typical leaves of L. oxycedrus; readily distinguished in L.juniperina). Scale = 1 cm.

the tube near the base of the anthers, but occasionally
one or a few hairs also occurred on the lobes.

Populations in the southern half of Tasmania, on the
south-eastern, southern and western coasts, typically
have a glabrous corolla or one with so few hairs that, for
identification purposes, it can be considered glabrous.
Further north, on the Bass Strait Islands and in the far
north-west (also in southern Victoria), the populations
seem to be more variable (as indicated by collections
at HO), and a greater proportion of plants have hairs
present on the corolla. In those plants, the hairs,
though sparse, are often more numerous on individual

flowers than seen in southern plants. In an examination
of 25 plants from Slaves Bay (north-western Tasmania),
all plants had one to a few hairs on the tube near the
anthers, and about half (13/25) had at least one or two
hairs on the limb.

The presence of sparse hairs on the corolla is clearly a
variable character within and between populations of
L. oxycedrus. Two other species that normally have a
glabrous corolla (L. juniperina and L. pogonocalyx) have
also been observed occasionally with sparse hairs on the
upper surface of the lobes and at the top of the corolla
tube.

6

Swainsona 31 (2017)

Distribution. The stronghold of Leptecophylla oxycedrus
is in Tasmania but the species also occurs in southern
Victoria (Ewart 1930; Willis 1973; Albrecht 1996).
It has been reported from New Zealand (Hooker
1833; Cheeseman 1906, 1925; Allan 1961), where it
has mostly been treated as a variety of L. juniperina.
However, we very much doubt its occurrence in that
country. Although Leptecophylla juniperina exhibits
a high level of morphological variability (as observed
from loan material), none of the variants we have seen
can be ascribed to L. oxycedrus.

Selected specimens examined

TASMANIA: Walkers Hill, Flinders Island, 20.x. 1976,
M.Allan (H027696); King Island, 24.iv.1802, R.Brown
(BM000797755, BM000797756); Kent Group, Bass Strait,
12-19.xii.1803, R. Brown (BM000797757); Nye Bay,
8.i. 1986, A.M.Buchanan 7702 (H097763); Observatory
site, Bennetts Point, Recherche, ll.ix.2003, A.M.Buchanan
16054 (H0523632); Nettley Bay Road, ll.i.2003,
L.H.Cave (H0583395); Maria Island, Bishop and Clerk,
I4.xii.2011, L.H.Cave 1395 (H0564872); Mt Munro, Cape
Barren Island, 7.x. 1988, P.A.Collier 3550 (HOI 18374);
Strzelecki summit, Flinders Island, 27.x. 1990, R.K.Crowden
& Y.Menadue (H0536147); Marrawah, v.1948, W.M. Curtis
(H053463); Surprise Bay, 6.iii.2003, H.J.Elliott
(H0583396); Old rehabilitated mine tailings near Grassy,
King Island, 26.X.2005, A.M.Gray 1590 (H0535658); Emu
Bay Road, 21 .viii. 1838, RC.Gunn 714 (HO4071); South
Hummock, Three Hummock Island, 5.x. 1995, S.Harris
& J.Balmer (HO445082); Balt Spur, Tasman Peninsula,
26.X.1979, J.Jarman (H031368); Slaves Bay, 13.X.2003,
J.Jarman & G.Kantvilas (H0583399); D’Entrecasteaux’s
Watering Place in Recherche Bay, 174.2004, J.Jarman &
G.Kantvilas (H0583400); Gillams Beach in Recherche
Bay, 4.ix.20l4, J.Jarman & G.Kantvilas (HO583401);
Macquarie Heads, 19.ix.20l4, J.Jarman & G.Kantvilas
(H0583402); Road to Redbill Point, Port Dalrymple,
12.ix.2015, J.Jarman & L.A.Thorne (HO583403); Track
to Cape Raoul, 19.xi.2003, G.Kantvilas (HO583404);
Coxs Bight, 31.xii.1982, D.I.Morris 8285 (H068182);
Maatsuyker Island, viii. 1976, A.Moscal (H032384); Point
Hibbs, 234.1984, A.Moscal 5863 (HO401591); Boat
Harbour, 17.xi.2002, M.R. & M.H.Stanton (H0583397);
Mt Leventhorpe, summit, Flinders Island, 3.iv.2007,
P.Tyson (H0583398); Tasman Island, downhill of quarters,
254x.2007, PA. Tyson 485 (H0545837).

VICTORIA: Cape Woolami, Phillip Island, 8.ix.l981,
A. Opie & S. Van Berkel (HO58969).

2. Leptecophyllaparvifolia (R.Br.) Jarman, comb. nov.

Cyathodesparvifolia R.Br., Prodr. 540 (1810). — Type:
[Tasmania: Mount Wellington (Table Mountain), near
River Derwent], RBrown, Feb-May 1804 (as Styphelia
erythrocarpa) (holo: BM000802314! Bennett No. 2416).
For additional notes and a detailed synonomy, see
Weiller (1999, p. 204) under Leptecophylla juniperina
subsp. parvifolia.

Leptecophylla parvifolia was first collected from Mt
Wellington (Table Mountain) by Robert Brown in

Leptecophylla in Tasmania: a reassessment of four species

1804 during his nine-month sojourn in southern
Tasmania. Brown named the plant Cyathodes parvifolia
(Fig. 8) and described it in his Prodromus florae Novae
Hollandiae, published in 1810. He commented on its
similarity to C. oxycedrus (= Leptecophylla oxycedrus)
which he had seen growing on the Bass Strait Islands
but observed that it was easily distinguished by its small
size. Since Brown’s initial description, L. parvifolia has
generally been accepted at specific rank (e.g. by de
Candolle 1838; Hooker 1859; Bentham 1868; Rodway
1903 and Curtis 1963). However, Sleumer (1963)
considered it a subspecies of Leptecophylla oxycedrus
(as Styphelia oxycedrus Labill. subsp. parvifolia (R.Br.)
Sleum.) and, more recently, Weiller (1999) treated it as
a subspecies of L. juniperina.

Leptecophylla parvifolia and L. juniperina

Several morphological characters readily separate
Leptecophylla parvifolia from L. juniperina. The corolla
shape and size is different (Fig. 9 compared to Fig. 4),
as well as leaf size, shape and venation, and the level
of retention of the transition leaves. Leptecophylla
parvifolia loses most of its transition leaves promptly
and the ones remaining are superficially similar to the
typical leaves. Even in the very oldest transition leaves,
the shape is more ellipical than oblanceolate, in contrast
to the persistent oblanceolate-shaped transition leaves
of L. juniperina.

Leptecophylla parvifolia and L. oxycedrus

In floral morphology, L. parvifolia is similar to
L. oxycedrus, except that the flowers are slightly
smaller (Fig. 9 compared to Fig. 3), and the lobes of
L. parvifolia are somewhat longer and narrower in
relation to the tube compared to those of L. oxycedrus.
Leaves of the two species may be similar in shape,
but those of L. parvifolia are distinctly shorter and
narrower, and they may be elliptical to oblong as well
as lanceolate.

In its typical form, L. parvifolia grows as a compact
rounded shrub, about 50-100 cm tall and often wider
than high, with many branches from ground level. Even
in dense understoreys at its lowest elevation, it rarely
exceeds 2 m in height. On the other hand, L. oxycedrus
typically occurs as a tall shrub greater than 3 m in
height, and has been reported as a small tree reaching
10 m (Curtis 1963, as Cyathodes juniperina). In extreme
coastal environments, it can occur as a dense wind-
pruned, low to medium shrub, but in such situations,
its overall appearance is very different from that of
L. parvifolia. Where living plants are growing in their
natural habitat there is little likelihood of confusion
between these two species.

The two species differ in ecology and distribution.
Leptecophylla parvifolia is widespread in Tasmania, and
abundant in the southern, central and north-eastern
highlands. It occurs in heathy vegetation or in open
forest dominated by various Eucalyptus species, and
can also be associated with high-altitude Nothofagus
cunninghamii forests. It is particularly abundant on

7

SJ. Jarman & G. Kantvilas

Swainsona 31 (2017)

(Sable Mountain
D< invent.

Cf. Steam, Introduction to Robert Breton’s Prodromus
(1960) prefixed to Hist. Nat. Classics facsimile.

R, Brown, Iter Australiense, 1802—5.

fPrescntod by direction or.T. J. Dessert. 1S70.1

x- g

i

u

cJi

Fig. 8. Leptecophylla parvifolia. Holotype collected by Robert Brown from Mt Wellington (Table Mt) in Van Diemens Land
(Tasmania) in 1804.

8

Swainsona 31 (2017)

Leptecophylla in Tasmania: a reassessment of four species

m

1MTT

c

B

♦ 'If* If

D

Fig. 9. Flowers of Leptecophylla parvifolia from Mt Wellington, the Type locality. A Pollen-producing flowers from three different
plants; B corolla of flowers shown in A. C Female flowers from three different plants; D corolla of flowers shown in C. Scale = 5 mm.

mostly grows as a medium to tall shrub in wet eucalypt
forest or coastal scrub, but may also form a small tree
(Fig. 11). Hooker (1859) records Cyathodes oxycedrus (=
L. oxycedrus) as ascending to 3000 ft, but this is possibly
an error resulting from confusion with Leptecophylla
pogonocalyx which, until the work of Weiller (1999),
remained unrecognised in the Tasmanian flora. The

Fig. 10. Leptecophylla parvifolia dominating the low heathy layer in eucalypt forest on Tasmania's Central Plateau.

the Central Plateau, where it occurs in broad expanses
as a dominant or subdominant low shrub in the
understorey (e.g. Fig. 10). It is found most commonly
above 500-600 m, ascending to over 1200 m, but
can occur at lower elevations especially where it has
dispersed from higher ground nearby. In contrast,
L. oxycedrus occurs in lowland coastal areas, where it

9

SJ. Jarman & G. Kantvilas

Swainsona 31 (2017)

Fig. 11 . Leptecophylla oxycedrus forming a small tree at
Recherche Bay. The fence post (scale) is c. 1.2 m tall.

highest elevation for L. oxycedrus known at present is
750 m on Mt Strzelecki on Flinders Island; the highest
on the Tasmanian mainland is 400 m on Mt Raoul on
the Tasman Peninsula. Its furthest recorded distance
from the coast is about 10 km on Mt Leventhorpe, also
on Flinders Island, but this, and other inland sites, are
likely to be subjected to strong maritime influences.
Leptecophylla parvifolia is restricted to the Tasmanian
mainland whereas L. oxycedrus extends to the Bass Strait
Islands and is also recorded from southern Victoria.

Taxonomic status

Morphological differences between L. parvifolia and
the New Zealand L. juniperina are such that there is
no justification in uniting the two. The situation is less
straightforward between L. parvifolia and L. oxycedrus.
However, the widespread distribution of L. parvifolia
in Tasmania, its abundance across very large areas, its
retention of small leaves across a range of ecological
conditions and, typically, its very different aspect
from L. oxycedrus , are strong arguments in favour of
separating the two. This view supports that of Brown
(1810), who described the species, and others who
worked on the Tasmanian flora such as Hooker (1859),
Rodway (1903) and Curtis (1963).

Descriptions of L. parvifolia are given in Bentham
(1868), Rodway (1903) and Curtis (1963), all as
Cyathodesparvifolia R.Br., and are not repeated here.

Selected specimens examined

TASMANIA: Bradys Lake, 17.x. 1975, M. Allan

(H0572582); Western Mountains, Western Tiers, i. 1848,
W.H.Archer (H04131); south slope of Mt Maurice,
9.xii.l979, A.M.Buchanan 76 (H032182); north ridge of
Mt Mangana, South Bruny Island, 9.xi.l984, A.M.Buchanan
4264 (HO407859); Mt Barrow, 27.xii.1959, T.E.Burns 231
(H04116); Lyell Highway, east of turnoff to Rufus Canal,
12.i.2003, L.H.Cave (H0583004); N ridge of Millers
Bluff, 25 km W of Campbell Town, 8.xii.l990, P.Collier
4976 (HO126290); Mt Faulkner, 26.ix. 1954, W.M. Curtis
(H04122); MacKenzies Tier, c. 3 km SSW of dam on Little
Pine Lagoon, 21.ii.20l4, M.F. de Salas 626, M.L.Baker &
G.Kantvilas (H0575139); Snow Hill Marshes, 29.iv.1986,

F. Duncan & M.Brown 83 (HO507457); Herringback,

summit, l.xii.2006, A. Gray 1756 (H0538694);

Poatina Highway, SW of Poatina, 17.xi.2002, J.Jarman
(HO583006); Murchison Highway, near Belmont Road
turnoff, 12.X.2003, J.Jarman & G.Kantvilas (H0583007);
Bruny Island, Lockleys Road, 2.ix.2015, J.Jarman &

G. Kantvilas (H0583008); Cathedral Rock, l.xii.2013,
G.Kantvilas (H0583005); Quoin Mountain, near summit,
20.iv.1986, AMoscal 12887 (HO410253); Cathcart Bluff,
11 .i. 1990, A.Moscal 18371 (HO 144597); Ben Lomond
National Park, at treeline near Rangers Headquarters,
18.xii.1979, M.G.Nohle (H074451); One O’clock Hill,
Murderers Marsh, Mt Dromedary, 20.i. 1981, A.E.Orchard
5277 (H040863); Lake Fenton area, Mt Field National
Park, 10.v. 1986, JM.Powell2153 (H0307760); Lake Sorell,
xi.1908, L.Rodway 496 (HO4079); Breona, Great Lake,
\4.xi.\947, J.Somerville (H053458); Collinsvale, 13.X.1934,
VV.Hickman (H04125).

10

Swainsona 31 (2017)

3. Leptecophylla juniperina (J.R.Forst. & G.Forst.)
C.M.Weiller

Muelleria 12: 200 (1999); Epacris juniperina J.R.Forst.

& G.Forst., Char. Gen. PI. 20, t. 10 (1776). —

Type: [New Zealand], sine loco, Forster, G.Forster’s
Herbarium (lecto, fide Weiller 1999: BM00079771!);

Herb. Pallas (isolecto: BM00079772!).

For a detailed synonomy, see Weiller (1999, p. 203).

Leptecophylla juniperina (Figs 4, 12) was first described
as Epacris juniperina in 1776 from material collected by
Johann and Georg Forster in New Zealand during James
Cook’s second Pacific Voyage in the ships Resolution and
Adventure (Forster & Forster 1776). The same species
was described 15 years later by Joseph Gaertner as Ardisia
acerosa (Gaertner 1791), again based on material from
New Zealand, but this time from specimens collected
by Joseph Banks and Daniel Solander on Cook’s earlier
voyage in the Endeavour in 1769. For many years, the
specific epithet acerosa was the one used most commonly
for the species ( Cyathodes acerosa or Styphelia acerosa),
with the earlier name given by the Forsters generally
being ignored. However, Sleumer (1963) and Curtis
(1963) reverted to the name juniperina and, thereafter,
the epithet has been applied widely.

Leptecophylla juniperina (as Cyathodes acerosa ) was
included in the Tasmanian flora at least as early as
1819, when Roemer & Schultes (1819) indicated it
occurred ‘In insulae Van Diemen’. The source of their
information is not clear and was possibly due to a
misinterpretation of Brown’s (1810) comments in his
Prodromus florae Novae Hollandiae, where he noted
the close similarities between Cyathodes oxycedrus
and C. acerosa. Another possible explanation is that
the information came from an examination of early
collections of plant material from Tasmania. Some of
Robert Brown’s specimens of C. oxycedrus from Bass
Strait collected in 1802 (BM00797756 from King
Island) and 1803 (BM000797757, Kents Group) carry
the name C. acerosa, but this was probably a temporary
field name used by Brown until further information
came to hand. Having the use of Joseph Banks’ library
and herbarium (Vallance etal. 2001), Brown would have
seen Banks and Solander specimens of C. acerosa from
New Zealand before his Australian trip commenced in
1801. He was also unlikely to have had access at that
time to Labillardiere’s Tasmanian specimens of Styphelia
(' Cyathodes ) oxycedrus, a species that was not published
until 1805. By the time Brown published his Prodromus
in 1810, he was apparently satisfied that C. oxycedrus
and C. acerosa were different, and that only one of
them, C. oxycedrus, occurred in Tasmania.

In spite of the distribution given by Roemer & Schultes
(1819), most of the early botanical references after
Robert Brown omitted C. acerosa from the Tasmanian
flora (e.g. de Candolle 1838; Hooker 1859). However,
in 1868, in Flora Australiensis, Bentham synonymised
the Tasmanian C. oxycedrus with C. acerosa under
the latter name. Through this synonymy, C. acerosa
(under various names) has continued to be listed as a

Leptecophylla in Tasmania: a reassessment of four species

Tasmanian plant by many botanists, with the notable
exception of Sleumer (1963).

Exclusion of L juniperina from the Tasmanian flora

The close relationship between Leptecophylla juniperina
and Tasmanian members of the genus is immediately
apparent from the general appearance of the plants,
which have narrow, sharply pointed leaves that are
striate and glaucous below. In spite of these similarities,
L. juniperina can be separated from all Tasmanian
species using vegetative characters and from most using
floral characters.

Important vegetative characters involve leaf shape and
venation, and persistence of transition leaves (see above
under L. oxycedrus ). Leptecophylla ahietina is the only
Tasmanian species that has any tendency to produce
persistent, oblanceolate transition leaves. However,
it is probably the most distinctive of all Tasmanian
Leptecophylla species and is very easily separated from
L. juniperina by a broad array of characters, including
the densely hairy corolla lobes, long corolla tube and
horizontally orientated trichomes between the veins on
the leaf undersurface.

In terms of floral morphology, L. juniperina differs
from all Tasmanian species except L. pogonocalyx in its
relatively short, broad corolla tube. However, the two
can be easily separated on the basis of leaf characters
(venation, subtleties of leaf shape, and differences in
shape and retention of transition leaves).

We have examined specimens of Leptecophylla juniperina
collected by the Forsters (lectotype: BM000797781,
Fig. 12; isolectotype: BM000797782) and Banks
and Solander (BM000802315, BM000802316)

and compared them with numerous specimens of
Leptecophylla from widely dispersed locations in
Tasmania. Comparisons have also been made with
material of L. juniperina on loan from New Zealand.
On this basis, we do not consider L. juniperina to be
a Tasmanian species, and formally exclude it from
the Tasmanian flora. Plants previously identified as
L. juniperina in Tasmania are mostly either L. oxycedrus
or a newly described subspecies of L. pogonocalyx.

Selected specimens examined

NEW ZEALAND, NORTH ISLAND: Cascade Park,
Waitakere Range, Waitemata City, Auckland, ll.ix.1974,
S.J.Astridge & L.RStemmer (CHR259266); Tairua State
Forest, 24.iv.1956, I.L.Barton (AK213083); Jacks Bay, Bay
of Islands, 3.viii.l967, R.Belcher (AK117922); Hauraki
Gulf, Ponui (Chamberlins) Island, 28.viii.1978, EA.Brown
(AK151185); north Waikawau Bay, 26.ix.2002, E.K.Cameron
(AK258766); Waitangi, Manukau County, 3.x. 1900, H.Carse
(CHR332679); Woodhill, West Coast, Waitemata County,
12.vii.1921, H.Carse (CHR332677); summits of hills at
Mangaroa, xi.1826, A. Cunningham [?] (MEL685918); W. of
Mt Messenger, N. Taranaki, iii.1978, A.P.Druce (CHR323871);
Pokai Reserve, S. Mamaku Plateau, x.1978, A.P.Druce
(CHR325967); Mt William, Pokeno, 27.x. 1972, R.O.Gardner

11

SJ. Jarman & G. Kantvilas

Swainsona 31 (2017)

Fig. 12. Lectotype of Leptecophylla juniperina (BM000797781) collected by the Forsters in New Zealand. Inset: magnified
segment of the main image.

12

Swainsona 31 (2017)

(CHR258390); Whangaparapara Bay, Great Barrier Island,
ix.1964, B.L.Gee & M.S.Reid (AK263286); Little Barrier
Island, l.ix.1963, P.Hynes (AK98692); Forestry Department
area, Woodhill, 17.viii.1968, P.Hynes (AK118807); Northcote,
Kauri Glen, 18.ix.1943, D.L.Knowlton (AK132300);
Hapuakohe Range, Waiti Road above Ohinekaua Stream,
6.vii.l991, P.J. de Lange 856 & G.M.Crowcroft (HO504556,
ex CHR473442); Birkenhead, in Le Roys Bush, l4.vii.1982,
D.Lewis (AK271068); Flunua Ranges, Mangatangi Track,
Auckland, 13.ix.2003, T.J.Martin (H0583394); Birkdale,
Auckland, 18.viii.1924, H.B.Mathews ? (AK105578); Little
Barrier Island, 26.vi.1947, H.RMcKenzie (AK211747);
Whangarei, 1957, T.W.Mellor (AK50410); Cascades,
Waitakere Ranges, 30.U983, JM.Powell 2074 (H0584244);
Baie des lies, 1843, M.Raoul (MEL685965); Wairata Forest
Farm, Wairata, Opotiki, x.2014, A.Redpath (H0583393); c.
3 km SW of Waiwera, 26.x. 1980, P.Scofield 29 (AK154355);
Mt Manganui, Tauranga, i. 1960, A.G.Simpson (AK266216);
c. 2 km SW ofWaiwera, 3.X.1981, G.Straka 336 (AK155408);
Mamaku, x.1941, M.R.Woodhead (AK151817); Auckland
(MEL2380635).

SOUTH ISLAND: Opuragi, Totaranui, 5-15.xi. 1769,
J.Banks & D. So lander (BM000802315, BM000802316).

4. Leptecophylla pogonocalyx C.M.Weiller

Muelleria 12: 206 (1999). — Type citation: above
Lake Dove, Cradle Mountain, eastern slopes,
21.xi.1985, C.M.Mihaich 5. Type sheet: From the
eastern slopes above Lake Dove, Cradle Mountain-
Lake St Clair National Park, 21.xi.1985, C.M.Mihaich
5 (holo: H0521873!).

Leptecophylla pogonocalyx was segregated by Weiller
(1999) from what had been known until then as
L. juniperina. Distinguishing characters are a short
corolla tube and pubescent calyx and bracteoles (Weiller
1999). The species is very widespread in the south-west
and west of Tasmania and is now known to cover a
much broader altitude range than when first reported,
occurring from sea level to alpine elevations.

In the present study, the circumscription of
L. pogonocalyx is broadened to include a new subspecies,
L. pogonocalyx subsp. decipiens. In their typical form,
subsp. pogonocalyx (Fig. 13) and subsp. decipiens (Figs 13,
14) can be distinguished unambiguously by the degree
of hairiness of the sepals and floral bracts (glabrous vs
hairy), and this distinction applies across much of the
area they occupy: subsp. pogonocalyx in southern and
western areas; subsp. decipiens in north-western areas.
However, where their distribution patterns converge
(north-western parts of the Central Plateau, parts of the
West Coast), it becomes increasingly difficult to assign
plants to one or other subspecies with certainty because
of the intergrading level of hairiness and the absence of
any other diagnostic character.

Leptecophylla pogonocalyx subsp. decipiens Jarman,
subsp. nov.

A L. pogonocalyce subsp. pogonocalyce C.M.Weiller
sepalis glabris differt.

Leptecophylla in Tasmania: a reassessment of four species

Type: slopes of Mt Leslie, 12.xi.2015, J.Jarman &

G.Kantvilas (holo: H0583888; iso: MEL, CANB,
AK, CHR, WELT).

Medium to tall shrub up to c. 4 m; branchlets
pubescent, with short stiff hairs. Leaves alternate,
spreading or reflexed, narrow-ovate to narrow elliptical-
lanceolate, tapering to a pungent point, 8-17 (-23) mm
long (including pungent point), 1.2-1.6 mm wide;
upper surface convex, green; lower surface glaucous,
striate with 3-5 (-7) unbranched parallel veins; margin
smooth or scabrous above; petiole short, c. 1 mm long,
glabrous or with short hairs on the upper surface; apex
acute, with a stiff sharp point, c. 1 mm long. Flowers
white, solitary, mostly in the upper axils, on straight
or recurved pedicels, 1-3 mm long; bracts glabrous
or with minute, inconspicuous hairs at the tip, ovate-
oblong to broadly triangular, margin ciliolate, apex
obtuse, upper bracts imbricate, decreasing in size to the
lower bracts, lower bracts imbricate or distant; sepals
glabrous or with a small patch of minute, inconspicuous
hairs at the apex, ovate-oblong, apex obtuse, margin
ciliolate; corolla tube 1.5-2.5 (-3) mm long, 2-3 mm
wide, bulbous in the middle, exserted beyond the
calyx or equal to it, corolla lobes spreading or reflexed,
1—1.5 mm long, shorter than the tube, glabrous or
rarely with sparse hairs. Anthers half-exserted. Ovary
5-locular; disk of five scales or lobes. Fruit a drupe,
pink to red, 7-10 mm diam. Figs 13D-F, 14.

Etymology. The epithet decipiens , from the Greek
participle meaning ‘deceiving’, refers to the general
appearance of the subspecies, which is so like that of
several other Tasmanian Leptecophylla species that the
taxon has been overlooked or misidentified in the past.

Distribution and ecology. Leptecophylla pogonocalyx
subsp. decipiens is the typical Leptecophylla occurring
throughout north-western Tasmania, from the lowlands
to over 1000 m elevation. It is known as far south as
the Pieman River and extends onto the northern and
north-western rim of the Central Plateau. It has not
been recorded from southern parts of Tasmania. It
commonly grows as a tall understorey shrub in wet
eucalypt forest. However, like other tall-growing
Tasmanian Leptecophylla species, it matures as a small
plant and can be found flowering at heights below 1 m
in open situations, at high elevations, or along margins
of roadsides and other disturbed areas. Peak flowering
time is October-November, with later flowering at
higher elevations.

Selected specimens examined

TASMANIA: Bowry Creek, Savage River, 9.vi.l993,
A.M.Buchanan 13378 (HO409926); south ridge of St
Valentines Peak, 13.1.1986, P.A.Collier 1165 (H0116892);
Baretop Ridge, 294.2015, M.F. de Salas 1140 & M.L.Baker
(H0578612); Mt Bertha, near summit trig point, 5.ii.2015,
M.F. de Salas 1354 & M.L.Baker (H0578425); Murchison
Highway 7.7 km N of Waratah and Guildford Roads
junction, lO.x.1978, AM.Gray 281 (H028100); near mouth

13

SJ. Jarman & G. Kantvilas

Swainsona 31 (2017)

Fig. 13. Leptecophylla pogonocalyx subsp. pogonocalyx (A-C) and L. pogonocalyx subsp. decipiens (D-F). A, D Pollen-
producing plant; B, E female plant; C flower showing hairy calyx; F flower showing glabrous calyx. Scale A, B, D, E = 5 mm; C, F =
1 mm.

Fig. 14. Leptecophylla pogonocalyx subsp. decipiens. A Pollen-producing flowers from three different plants; B corolla from
flowers shown in A. C Female flowers from three different plants; D corolla from flowers shown in C. Scale = 5 mm.

of Pieman River, 15.i. 1954, W.D.Jackson 136 (HO4096);
Wandle River, 16.xi.20l4, J.Jarman (H0584431); Loyatea
Peak, 19.X.2001, J.Jarman & G.Kantvilas (H0584432);
Black Bluff Track, 19.X.2001, J.Jarman & G.Kantvilas
(H0584433); Savage River Pipeline Road, 27.xi.2003,
J.Jarman & G.Kantvilas (H0584434); Murchison Highway,

near Belmont Road turnoff, 28.xi.2003, J.Jarman &
G.Kantvilas (H0584435); Blackwater Road near spur 5,
10.xi.2015, J.Jarman & G.Kantvilas (HO583890); Norfolk
Road near the junction with Sumac Road, 10.xi.2015,
J.Jarman & G.Kantvilas (H0583891); Henrietta,
12.xi.2015, J.Jarman & G.Kantvilas (H0583892); Bonds

14

Swainsona 31 (2017)

Range, 25.xi.1982, A.Moscal 1044 (H068368); The Clump,
9.xii.l983, A.Moscal 4666 (H079821); Frankland Creek,
18.xii.1983, A.Moscal5003 (HOI 11287); Providence Creek,
Arthur River, 28.iii.1984, A.Moscal 7166 (H0121900);
Trowutta Arch, 7.iv.l984, A.Moscal 7434 (HO92140);
Mother Cummings Peak, 20.ii. 1986, A.Moscal 12368
(H0402240).

Composition of Leptecophylla in Tasmania

On the basis of this study, the genus Leptecophylla
comprises six species in Tasmania, one of which has
two subspecies. These include L. abietina (Labill.)
C.M.Weiller, L. divaricata (R.Br.) C.M.Weiller, and
L. pendulosa (Jarman) C.M.Weiller, in addition to the
taxa documented herein. An identification key to these
species is provided (below).

Acknowledgements

We thank the many people who have assisted during
the course of this study. In particular, Chiara Nepi
of the Herbarium Universitatis Florentinae was very
helpful during a visit there (by GK) and also provided
digital images of Labillardieres specimens collected
in Tasmania. Roy Vickery (formerly Natural History
Museum, London) facilitated the loan of material
from there and provided photocopies of excerpts from
historical documents, and Tony Orchard (Canberra)
provided advice on the choice of Type material.
We thank the staff at MEL, AK, CHR, WELT and
BM for loans from those herbaria, and Kim Hill
(Tasmanian Herbarium) for managing the associated

Leptecophylla in Tasmania: a reassessment of four species

documentation. Special collecting efforts by Lyn Cave
and Penny Tyson (Tasmania) and Anne Redpath,
Rhys Gardner and Ewen Cameron (New Zealand) are
also greatly appreciated. Thanks also to the Auckland
Museum and Ewen Cameron for permission to dissect
and photograph material of Leptecophylla juniperina for
publication.

References

Albrecht, D.E. (1996). Epacridaceae. In: Walsh, N.G. &
Entwisle, T.J. (eds), Flora of Victoria 3: 464-509. (Inkata
Press: Melbourne).

Allan, H.H. (1961). Flora of New Zealand, Vol. 1. (Government
Printer: Wellington, New Zealand).

Bentham, G. [1868] (1869). Flora Australiensis, Vol. 4. (Reeve
& Co.: London).

Brown, R. (1810). Prodromus florae Novae Hollandiae et insulae
Van Diemen. (Johnson: London).

Candolle, A.P. de (1838). Prodromus systematis naturalis regni
vegetabilis, sive enumeratio contractu ordinum, generum,
specierumque plantarumVW: 741. (Treuttel & Wurtz: Paris).
Cheeseman, T.F. (1906). Manual of the New Zealand flora.

(Government Printer: Wellington).

Cheeseman, T.F. (1925). Manual of the New Zealand flora, 2 nd
edn. (Government Printer: Wellington).

Crowden, R.K. & Duretto, M.F. (2013). Flowering plants of
Tasmania: subspecies of Leptecophylla. On: KeyBase. http://
keybase.rbg.vic.gov.au [accessed: December 2013, July
2016].

Curtis, W.M. (1963). The student’s flora of Tasmania, Part 2.

(Government Printer: Tasmania).

Ewart, A.J. (1930). Flora of Victoria. (Government Printer for
University Press: Melbourne).

Key to Tasmanian species of Leptecophylla *

1. Flowers with a short, broad corolla tube: in pollen-producing flowers, the length of the
tube equal to or less than the diameter at the widest point; tube bulbous in shape (e.g.

Fig. 14)

2. Sepals pubescent on the outer surface. L. pogonocalyx subsp. pogonocalyx

2: Sepals glabrous on the outer surface. L. pogonocalyx subsp. decipiens

1: Corolla tube relatively long in relation to its diameter: in pollen-producing flowers, at
least 1.5 times as long as the width; tube barrel-shaped or cylindrical (e.g. Figs 3,9)

3. Style long, at least twice the height of the ovary

4. Corolla lobes with long straggling hairs; bracts distant, bracteoles scarcely

reaching the sepals. L. divaricata

4: Corolla lobes glabrous, rarely with one or two hairs; bracts and bracteoles over¬
lapping . L. pendulosa

3: Style short, less than twice the height of the ovary

5. Leaves longer than 7 mm

6. Inside of corolla lobes densely hairy. L. abietina

6: Inside of corolla glabrous or with scattered sparse hairs. L. oxycedrus

5: Leaves shorter than 7 mm. L. parvifolia

* Leptecophylla juniperina is excluded from the Tasmanian flora.

15

S.J. Jarman & G. Kantvilas

Swainsona 31 (2017)

Forster, J.R. & Forster, G. (1776). Characteres generum
plantarum quas initinere ad insulas mans australis. (B. White,
T. Cadell & P. Elmsly: London).

Gaertner, J. (1791). De fructibus et seminibusplantarum. Vol. 2.
(G.H. Schramm: Tubingen).

Gardner, R. (2011). Sex ratios of some trees native to New
Zealand. Auckland Botanical Society Journal 66(2): 151-155.
Hooker, J.D. (1853). The botany of the Antarctic voyage of
H.M. discovery ships ‘Erebus’ and ‘Terror’, in the years 1839-
1843. II. Flora Novae-Zelandiae, Part 1, Flowering plants.
(Lovell Reeve: London). [Facsimile edn 1963, J. Cramer:
Weinheim].

Hooker, J.D. [1859] (1860). The botany of the Antarctic voyage
of H.M. discovery ships ‘Erebus’ and ‘Terror’, in the years
1839-1843. Ill Flora Tasmaniae, Vol. 1, Dicotyledones.
(Lovell Reeve: London). [Facsimile edn 1963, J. Cramer:
Weinheim].

Kron, K.A., Judd, W.S., Stevens, P.F., Crayn, D.M., Anderberg,
A.A., Gadek, P.A., Quinn, C.J. & Luteyn, J.L. (2002).
Phylogenetic classification of Ericaceae: Molecular and
morphological evidence. The Botanical Review 68(3):
335-423.

Labillardiere, J.J. (1800). Relation du voyage cl la recherche de la
Perouse. (English translation, J. Stockdale: London).
Labillardiere, J.J. (1805). Novae Hollandiaeplantarum specimen.
Vol. 1. (Huzard: Paris).

Rodway, L. (1903). The Tasmanian flora. (Government Printer:
Hobart).

Roemer, J.J. & Schultes, J.A. (1819). Systema vegetabilium:
secundum classes, ordines, genera, species. Cum characteribus,
dijferentiis etsynonymiis. (J.G. Cottae: Stuttgart).

Sleumer, H. (1963). Florae Malesianae precursores XXXVII.
Materials towards the knowledge of the Epacridaceae mainly
in Asia, Malayasia, and the Pacific. Blumea 12(1): 145-171.
Sprengel, K. [1824] (1825). Systema vegetabilium. Editio decima
sexta. Vol. 1. (Librariae Dietrichianae: Gottingen).

Vallance, T.G., Moore, D.T. & Groves, E.W. (2001). Nature’s
investigator: The diary of Robert Brown in Australia, 1801—
1805. (Australian Biological Resources Study: Canberra).
Weiller, C.M. (1999). Leptecophylla, a new genus for species
formerly included in Cyathodes (Epacridaceae). Muelleria
13(2): 195-214.

Willis, J.H. (1973). A handbook to plants in Victoria , Vol. 2.
(Melbourne University Press).

With the exception of images and other material protected by a trademark and subject to review by the Government of
South Australia at all times, the content of this publications is licensed under the Creative Commons Attribution 4.0 Licence
(https://creativecommons.Org/licenses/by/4.0/). All other rights are reserved.

© 2017 Board of the Botanic Gardens and State Herbarium (Adelaide, South Australia)

16

w

Swainsona 31:17-26 (2017)

© 2017 Board of the Botanic Gardens & State Herbarium (Adelaide, South Australia) ©0

A new species of small black disc fungi, Smardaea australis (Pezizales,
Pyronemataceae), is described from Australia

Pamela S. Catcheside a - b , Samra Qaraghuli b & David E.A. Catcheside b

a State Herbarium of South Australia, GPO Box 1047, Adelaide, South Australia 5001
Email: Pam.Catcheside@sa.gov.au

b School of Biological Sciences, Flinders University, PO Box 2100, Adelaide, South Australia 5001
Email: Samra.Qaraghuli@flinders.edu.au, David.Catcheside@flinders.edu.au

Abstract: A new species, Smardaea australis P.S.Catches. & D.E.A.Catches. (Ascomycota, Pezizales,
Pyronemataceae) is described and illustrated. This is the first record of the genus in Australia. The
phylogeny of Smardaea and Marcelleina, genera of violaceous-black discomycetes having similar
morphological traits, is discussed.

Keywords: Fungi, discomycete, Pezizales, Smardaea, Marcelleina, Australia

Introduction

Small black discomycetes are often difficult or impossible
to identify on macro-morphological characters alone.
Microscopic examination of receptacle and hymenial
tissues has, until the relatively recent use of molecular
analysis, been the method of species and genus
determination.

Between 2001 and 2014 five collections of a small
black disc fungus with globose spores were made in
South Australia. Initially the fungus was identified as
Marcelleina atroviolacea Brumm., since it conformed
in all respects with a fungus previously collected in
1964 from Anglesea, Victoria, by Gordon Beaton.
This collection had been sent by Beaton to the Royal
Botanic Gardens, Kew, and described by Rifai (1968)
as M. atroviolacea.

No other species of Marcelleina Brumm., Korf & Rifai
has been recorded in Australia, nor of other possibly
confusing genera such as Smardaea Svrcek and Otidea
(Pers.) Bonord, although species of the latter genus
have been found in New Zealand (Atlas of Living
Australia, ALA). There are no records of any species of
Smardaea in the ALA or Australia’s Virtual Herbarium
(AVH). Those in the ALA and AVH of Marcelleina
atroviolacea are of the collections from South Australia
covered in the present study and of Beaton’s collection
from Victoria. The ALA also records M. atroviolacea
as being in New Zealand, but these collections had
been misidentified and were Pseudoplectania affinis
M.Carbone, Agnello & P.Alvarado (P. Johnston, pers.
comm.; NZFungi2 2016). Pseudoplectania Fuckel also

has dark coloured apothecia and globose ascospores,
but differs morphologically from Smardaea in having
dark hairs on the excipulum.

Marcelleina and Smardaea

Four genera of small black discomycetes with purple
pigmentation, Greletia Donad., Pulp aria P. Karst.,
Marcelleina and Smardaea , had been separated
by characters in part based on distribution of this
purple pigmentation, as well as on other microscopic
characters. Their relationships have been much
discussed (Donadini 1984; Pfister 1985; Moravec
1987; Haffner 1995; Peric 2001; Benkert 2005), but
Korf & Zhuang (1991) followed Donadini (1979) in
accepting two genera: Marcelleina (with Pulparia as a
synonym) and Smardaea (with Greletia as a synonym).

The generic name Marcelleina was proposed by van
Brummelen, Korf and Rifai in 1967 (Brummelen
1967; Rifai 1968; Moravec 1987) in honour of Mme
Le Gal, an eminent French mycologist who specialised
in discomycetes. The genus Smardaea was proposed by
Svrcek in 1969, named after Frantisek Smarda, a Czech
botanist.

Species in the genera Marcelleina and Smardaea have
small, cupulate to saucer-like, sessile or subsessile,
glabrous apothecia with violaceous-brown hymenium.
The receptacle tissue of both is similar: ectal excipulum
of globose or angular cells ( textura globulosa or textura
angularis) and medullary excipulum of interwoven
septate hyphae ( textura intricata). Asci are cylindrical
and their apices do not turn blue in Melzer’s reagent.
Paraphyses are septate and contain purple granules.

Published online: 15 June 2017 • flora.sa.gov.au/swainsona

ISSN 2206-1649 (Print) • ISSN 2206-1657 (Online)

P.S. Catcheside etal.

Swainsona 31 (2017)

The two genera differ in that all Marcelleina species
have globose spores, the paraphyses are usually
unbranched and, though the sterile tissue and
paraphysis contents have purple pigmentation, the asci
and ascospores are not necessarily purple. The spores
of Smardaea may be globose or ellipsoid, paraphyses
are usually branched and there is purple pigment in
asci, ascospores, paraphyses and excipula (Benkert &
Moravec 1986; Moravec 1987; Hansen & Knudson
2000 ).

Nine species of Smardaea are presently recognised
(Index Fungorum), four of which have globose spores.
Of these only S. planchonis (Dun. ex Boudier) Korf
& W.Y.Zhuang has smooth spores; S. reticulosperma
(Donadini, Riousset & G. Riousset) Benkert has
reticulate spores, those of S. verrucispora (Donadini &
Monier) Benkert are warty and S. marchica (Benkert
& J. Moravec) Benkert has subglobose, coarsely warted
spores.

Eleven species of Marcelleina are listed in Index
Fungorum. Seven of these have ornamented spores
with ornamentations ranging from warty-tuberculate,
ridged or partially to completely reticulate. Of the
four smooth-spored species, M. benkertii J.Moravec,
M. chopraiana (L.R.Batra) S.C.Kaushal, M. parvispora
E.Rubio, Tabares & M.A.Martinez and M. atroviolacea,
only the latter has purple colour in all tissues and
structures. The generic position of M. parvispora will be
discussed later in this paper.

Korf & Zhuang (1991) synonymised Marcelleina
atroviolacea and Smardaea planchonis and erected the
new combination Smardaea planchonis.

Distribution and habitat of Smardaea planchonis

Smardaea planchonis , albeit under its various synonyms,
is widespread in the northern hemisphere (Rifai 1968;
Peric 2001). The type locality is France and it has been
found in a number of countries in central and southern
Europe (Boudier 1887; Lagarde 1911; Donadini 1984;
Moravec 1987; Marchetti & Franchi 1993; Haffner
1995; Peric 2001; Benkert 2005; Martin 2005; Cuesta
& Ribes 2006; Lantieri et al. 2009), the U.S.A. (Pfister
1985), Bermuda (Seaver 1928) and has been reported
in Argentina in the southern hemisphere (Gamundi
1960; Rifai 1968).

Smardaea planchonis is reported as growing in
mostly sandy soil, often amongst moss and under
Cupressaceae. For example, in Tuscany it has been
found under Cupressus sempervirens L., C. glabra Sudw.,
Juniperus oxycedrus subsp. macrocarpa (Sibth. & Sm.)
Ball and J. phoenicea L. (Marchetti & Franchi 1993).
Benkert (2005), in his discussion on the distribution
and ecology of S. planchonis , reported its presence
under Juniperus horizontalis Moench., J. chinensis
“plumosa aurea” and Chamaecyparis obtusa (Siebold &
Zucc.) Endl. In the U.S.A. Pfister (1985) recorded it
under Yucca. The Argentinian collections were from a

park in the inland city of Mendoza, on moist soil in
sandy groves.

Materials and methods

Habitat and associated plant communities were noted
in the field. Collection locations were recorded by
GPS, geodetic datum WGS84/GDA94 (Garmin
GPS 12) and in situ photographs taken (Nikon 4500).
Macroscopic characters were described directly from
fresh material. Colours are designated using the Royal
Botanic Gardens Edinburgh Colour Chart (1969) and
given as colour descriptor and number, e.g. “rust 13”,
and in general terms. Fresh material was dried in a food
dehydrator at 35°C for 24 h (Hydraflo 1000FD).

Sections of fresh material and dried specimens were
hand-cut and mounted in various media. For the
amyloid reaction, fresh material was stained with
Melzer’s reagent and dried material was rehydrated in
5% NH 4 OH before staining. Water mounts were used
to determine colour of context, 5% KOH and 5%
NH 4 OH were used to determine reaction to alkali, and
5% H 2 SO 4 to determine the acid reaction.

Measurements were made using an Olympus BH-2
microscope at x400 or xlOOO with a calibrated ocular
micrometer. Spore dimensions are given as: length
range x width range (n = 40) and Q ratio (spore length/
spore width). Dimensions of asci are given as length
range x width range (n = 20). A Nikon 4500 camera
was used to photograph microscopic characters.

Descriptions of Smardaea australis are based on the type
collection, P.S.Catcheside 4079 (AD-C 58765), with
outlying measurements for other collections given in
brackets. Photographs of fruit bodies and microscopic
characters are from the type collection. All South
Australian collections have been accessioned into the
State Herbarium of South Australia (AD). AD numbers
(AD-C nnnnn) are given in the Taxonomy section
together with the Collector’s number (PSC nnnn); in
other sections only the Collector’s number is used.

DNA Extraction, amplification and processing were
as described in Catcheside et al. (2016). To place
species of interest in the Pyronemataceae, sequences
of the ribosomal RNA large subunit gene were aligned
with those of representatives of each of the currently
recognised families and lineages chosen from those used
by Hansen et al. (2013). Sequences were manipulated
with the Geneious 8.1.9 suite of programmes using
Muscle for alignment and, for tree building, either
neighbour joining (Fig. 8) or MrBayes (Fig. 9) using
the HKY85 substitution model, 4 heated chains for
1,100,000 iterations including a burn-in of 100,000.
Correlation between genetic and physical distance
between collections was examined using the Pearson
product-moment test in R.

18

Swainsona 31 (2017)

Taxonomy

Smardaea australis P.S.Catches. & D.E.A.Catches. sp.
nov.

Holotype: South Australia. Sleaford Bay, Coffin Bay
National Park, on low sand dune, 34° 31' 12.8"S, 135°
44' 6.4"E, alt. c. 5 m, amongst moss with Leucopogon
parviflorus (Andrews) Lindl., Melaleuca sp. and Acacia
longifolia subsp. sophorae (Labill.) Court, 21 July 2014,
PS. Catchcside PSC 4079 & D.E.A. Catchcside (AD-C
58765).

Mycobank number: MB819607.

Apothccia scattered to gregarious, sessile; (3-) 5-15 mm
diameter, (1-) 3-8 mm high (Fig. 1). Disc more or
less circular, concave to flat; margin initially slightly
inrolled, later becoming plane, entire to crenate,
often becoming eroded; fuscous-black 36, violaceous-
black 38, blackish-purple; smooth, sometimes slightly
pitted, waxy. Receptacle saucer- to cup-shaped; sessile;
concolourous with disc; smooth to finely scurfy;
attached to substrate by a small central soil pad (Fig. 2:
Section through hymenium and excipulum; Fig. 3:
Hymenium showing asci, ascospores and paraphyses).
Flesh 1—1.5 mm thick; dark brown, purplish. Ectal
excipulum 50-100 pm thick (Fig. 4). The outer
layer composed of thick-walled globose, subglobose,
sometimes polygonal cells (textura globulosa :), cells
15-30 pm across; walls purple-brown; clumps of
cells aggregated to give receptacle a slightly scurfy

A new species of small black disc fungi, Smardaea australis

appearance. Inner layer of thinner-walled, cylindric-
subglobose cells to 40 pm long, arranged with long
axis at right angles to surface of receptacle; colour of
cells ranging from colourless to purple to red-purple in
NFUOH, pale purple to deeper or red-purple in KOH,
more strongly purple/red-purple in water, orange-
red in H 2 SO 4 . Medullary excipulum of interwoven,
septate, branched hyphae 4-10 pm diameter (textura
intricata ), with similar colour reactions as cells of the
ectal excipulum. Suhhymenium of more compacted
hyphae. Hymenium 200-250 pm thick. Asci 8-spored;
cylindrical-clavate, (160-) 170-225 x 8.5-12.5 pm
(Fig. 5); amyloid; attenuating towards base; base forked,
arising from croziers (Fig. 6). Immature asci with purple
colouration in alkali and water, red in acid, no colour
in fully mature asci. Ascospores globose (7.5-) 8.0-
10.5 (-11.5) x (7.0-) 8.0-10.0 (-11.5) pm diameter;
Q = 1-1.08 (-1.09); smooth; uniseriate; in upper
portion of the ascus; hyaline, subhyaline; often with
one large globule and several smaller ones. Immature
ascospores with purple colouration in alkali and water,
red in acid, no colour in fully mature spores. Paraphyses
slender, filiform (Fig. 7); slightly longer than asci;
straight or slightly curved; sparsely septate; tips slightly
swollen 2-7 pm diameter; with granular contents which
are more densely concentrated at tips; granular contents
purple to purple-brown in water and alkali, becoming
browner with age of apothecium; paraphyses branching
especially near base; often nodulose.

Fig. 1. Smardaea australis. Apothecia in situ. Scale = 10 cm.

19

P.S. Catcheside etal.

Swainsona 31 (2017)

Fig. 2. Smardaea australis. Section through hymenium and
excipulum. In water. Scale = 100 pm.

Fig. 3. Smardaea australis. Hymenium showing asci,
ascospores, paraphyses. In water. Scale = 100 pm.

Additional specimens examined.

VICTORIA. Two miles West of Anglesea, terrestrial between
road and dunes, 38° 25' 48.44"S, 144° 8' 40.14"E, 5 July
1964, as Marcelleina atroviolacea Brumm., G.W. Beaton
(MELU F 121706a).

SOUTH AUSTRALIA. Flinders Ranges National Park below
Bunyeroo Gorge car park, on surface of mostly bare clay soil
amongst stones, 31° 25' 00"S, 138° 33' 29"E, alt. c. 300 m,
on the bank of creek bed colonised by moss protonema,
near Callitris glaucophylla Joy Thomps. & L.S.A. Johnson
and Eucalyptus camaldulensis Dehnh., 10 Aug. 2001, P.S.
Catcheside PSC 1020 & D.E.A. Catcheside (AD-C 51219);
Mambray Creek, Mount Remarkable National Park, in soil
amongst moss, 32° 49' 17"S, 138° 03' 37"E, alt. c. 200 m,
in dry woodland near C. glaucophylla, E. camaldulensis,
Leucopogon sp. and Acacia sp., 9 Aug. 2011, P.S. Catcheside
PSC 3637 & D.E.A. Catcheside (AD-C 57291); Wilpena
Pound, Flinders Ranges National Park, on surface of mostly
bare clay soil, 31° 33' 03"S, 138° 35’ 22"E, alt. c. 540 m,
amongst C. glaucophylla, 16 July 2014, P.S. Catcheside PSC
4043 & D.E.A. Catcheside (AD-C 58766); Lincoln National
Park, Eyre Peninsula, in soil and amongst moss, 34° 52’
12"S, 135° 52' 38"E, alt. c. 40 m, on raised bank at side of
road, sandy dune heath, E parviflorus, Melaleuca sp. and
A. longifolia subsp. sophorae, 22 July 2014, P.S. Catcheside
PSC4080 &D.E.A. Catcheside (AD-C 58767).

Etymology. From the Latin australis, meaning southern.
Referring to the geographic distribution of the species
in the southern hemisphere.

Note. The nuclear ribosomal sequences of each of the
five collections, PSC 1020 (GenBank KY067461), PSC
3637 (GenBank KY067462), PSC 4043 (GenBank
KY067463), PSC 4079 (GenBank KY067464) and
PSC 4080 (GenBank KY067465), differ (Fig. 8), with
PSC 1020 and PSC 4043 being heterozygous at single
nucleotide positions in ITS2 and ITS1, respectively.
There are ten variable sites in ITS1 (5.7%), eight in
ITS2 (4.1%) and a further six in the ^960bp of the
ribosomal RNA large subunit gene that was sequenced.
This degree of intraspecific divergence in ITS sequence
is high for ascomycetes (Nilsson et al. 2008). However,
there is no significant correlation of sequence divergence

and distance between collection sites (Pearson product-
moment t = 1.38, df = 19, p = 0.18) and no obvious
grouping with respect to substrate or plant association.
Accordingly we consider each to be strains of Smardaea
australis.

Notwithstanding the morphological similarity of
Smardaea and Marcelleina species, comparison of
ribosomal RNA large subunit sequences shows they are
only distantly related members of the Pyronemataceae
placed in the Otidea lineage and Pezizaceae respectively
(Fig. 9).

Discussion

With the relatively few morphological characters to
distinguish between them, identification of small black
discomycetes is problematic. Spore characters, ascus
amyloidity, pigmentation of tissues have been and
are used to help identification to genus and species
levels. Since the development of molecular sequencing
and phylogenetic analyses, it has become clear that
macro- and micromorphological characters alone
are insufficient to determine taxonomic identity and
phylogenetic relationships.

Taxonomy, morphology and purple pigmentation
of Marcelleina and Smardaea

The similarities of the smooth, globose-spored species
of Marcelleina and Smardaea make separation on
morphological characters of the two genera difficult,
if not impossible. As currently understood, all species
of Marcelleina have inamyloid asci, globose spores and
partial purple pigmentation of tissues. Marcelleina
atroviolacea was alone amongst the marcelleinas in
having purple pigment in all tissues, a character that
suggests it be put in the genus Smardaea. Svrcek’s concept
of the genus Smardaea was of species having inamyloid
asci, oblong-ellipsoid, verrucose spores and all tissues
with violaceous colouration. When Korf & Zhuang
(1991) synonymised M. atroviolacea and S. planchonis, a
synonymy that the present authors accept, they extended

20

Swainsona 31 (2017)

Fig. 4. Smardaea australis. Ectal exipulum. In water. Scale =
10 pm.

the generic concept of Smardaea to include species with
globose, smooth spores. Benkert (2005) commented
that S. planchonis is the only species of Smardaea with
such spores. S. australis becomes the second species in the
genus with that combination of spore characters.

When Beaton identified his Victorian collection
of a discomycete as Marcelleina atroviolacea he was
following Rifai’s concept of M. atroviolacea. Rifai
documented the Australian Pezizales that had been
sent to the Royal Botanic Gardens at Kew, a work
that was published in 1968. The genus Smardaea was
not erected until the following year (Svrcek 1969) so
Rifai’s concept of M. atroviolacea would now place
that taxon in the genus Smardaea. Beaton’s collection
of M. atroviolacea (MELU F 121706a) was examined
and found to be similar in all respects to the South
Australian collections, now named S. australis , based on
comparison of spore and ascus measurements (Table 1),
hymenial tissue, their inamyloid asci, smooth, globose
spores and branched paraphyses with purple pigment.

The purple pigmentation in tissues is obviously an
important character in determining genus and species,
but its variability presents difficulties in interpretation.
The asci and ascospores of all Smardaea species contain
purple pigment while those of Marcelleina species are
not necessarily purple; the paraphysis contents of both
genera contain purple pigment.

In all Australian collections of Smardaea australis the cells
and colouration of excipular tissue were similar: ectal
excipulum of thick-walled, purplish-brown, globose to
subglobose cells, medullary tissue of densely interwoven,
purple hyphae. However, the intensity of the purple
pigmentation in hymenial tissues varied in the different
collections and was seemingly age-dependent. Asci,
ascospores and paraphyses of collection PSC 4043 had
the most intense pigmentation, those in PSC 4080 the
least. The apothecia of PSC 4080 were more mature than
those of the other collections with almost all asci having
mature ascospores. The distribution of pigmentation
also varied in all collections: immature asci had distinct

A new species of small black disc fungi, Smardaea australis

Fig. 5. Smardaea australis. Asci. In water. Scale = 10 pm.

purple colouration as did their ascospores. The purple
granular material in the paraphysis tips was variable,
some paraphyses had strongly pigmented contents while
the contents of others were clear. From our observations
it appears that the pigmentation fades with maturity and
thus is age-dependent. All pigmented tissues became a
slightly darker, duller purple in KOH and NH 4 OH and
red-orange in acid. The purple colour was brightest in
water which was found to be the most useful medium
when determining intensity and distribution of the
pigmentation.

While morphologically the Australian material has
some characteristics of Marcelleina species, the extent
of purple pigmentation and sometimes branched
paraphyses fit better with Smardaea. Morphologically
our new species, Smardaea australis , most closely
resembles the northern hemisphere S. planchonis ,
however, it differs from S. planchonis in having shorter
asci (Table 1): the ascus length of S. australis ranges
from 155-225 pm, while that of S. planchonis is from
190-240 pm, if the measurements of Boudier (1887)
are discounted. The size of the ascospores of both
species is similar. Unfortunately there are currently no
molecular data for S. planchonis , so the two taxa cannot
be compared on a molecular basis.

Phylogenetic relationships of Marcelleina and
Smardaea

Recent phylogenetic studies confirm the placement
of Marcelleina in the Pezizaceae (Hansen et al. 2001;
Hansen et al. 2005; Hansen & Pfister 2006; Perry et al.
2007; Tedersoo et al. 2010; Hansen et al. 2013). The
position of the genus Smardaea in the Pyronemataceae
is rather more complex. Perry et al. (2007) placed
two species, S. reticulosperma (Donadini, Riousset &
G.Riousset) Benkert and S. amethystina (W.Phillips)
Svrcek in the Pyropyxis clade and several species of Otidea
in the adjacent Otidea clade. Tedersoo et al. (2013) show
similar placements. Hansen et al. (2013) separated the
Pyropyxis and Otidea lineages, while Frey et al. in Englers
Syllabus of Plant Families (2016) treat the large family
Pyrenomycetaceae s.l. by dividing it into several groups,

21

P.S. Catcheside etal.

Swainsona 31 (2017)

Fig. 6. Smardaea australis. Croziers. In water. Fig. 7. Smardaea australis. Paraphyses. In water. Scale = 1 00 pm.
Scale = 10 pm.

among them the family Otideaceae Eckblad, in which
Smardaea, Pyropyxis and Otidea are included.

Ribosomal 28S sequences confirm that the Australian
collections are within the genus Smardaea (Fig. 9)
with the closest sister species being S. reticulosperma
and S. amethystina. Smardaea reticulosperma, like
S. australis, has globose spores but these have reticulate
ornamentation. Smardaea amethystina has ellipsoid,
warty spores. In the same clade are taxa lacking
purple pigment, thus separating them from Smardaea'.
Pyropyxis rubra (Peck) Egger and two species of Jafnea,
J. fusicarpa (W.R. Gerard) Korf and J. semitosta (Berk. &
M.A.Curtis) Korf. Pyropyxis rubra has ellipsoid, smooth
spores, but differs from Smardaea species in containing
orange pigment. Jafnea species have ellipsoid spores,
and the apothecia are brown and have hairs, another
character that separates them from Smardaea whose
apothecia are glabrous. Otidea species lie within the
same clade. They are characterised by having apothecia
which are split down one side, are yellow, brown or
ochre, and have smooth, ellipsoid spores.

Thus there seems no particular morphological character
that typifies members of the Otideaceae. Amyloidity of
asci has been used to separate taxa such as the family
Pezizaceae from the Pyronemataceae (Rifai 1968). The
Pyronemataceae, which now include the Otideaceae,
have inamyloid asci; the amyloid reaction of the ascus is
a character associated with the Pezizaceae. Hansen et al.
(2001) consider that amyloidity has been lost in some
lineages, such as Marcelleina.

Marcelleina is in a lineage which is some distance
from the Otideacae (Fig. 9). The phylogenetic data
of Hansen et al. (2013) show Marcelleina and Peziza
gerardii Cooke in the Marcelleina/P. gerardii lineage
in the Pezizaceae as opposed to the Pyrenomycetaceae
s.l. in which the Otideaceae lie. They also pointed
out that Schumacher & Jenssen (1992) had noted
the morphological similarities between P. gerardii
and Marcelleina and Smardaea. Peziza gerardii is a
violaceous-brown discomycete with amyloid asci and
fusoid, ribbed ascospores (Beug et al. 2014), thus
differing from Marcelleina with its inamyloid asci and

Q07 y PSC4080 Lincoln NP
PSC4079

Coffin Bay NP\ Q

PSC1020B

TO.07

Bunyeroo Gorge® PSC1020A

1.0

Mambray Creek

rfp*

r^-#PSC4043B

rtf*

&

0.07

Wilpena Pound

Fig. 8. Smardaea australis is genetically diverse. Neighbour joining tree of the sequences from five collections (-960 bp
comprising 18S partial, ITS1, 5.8S, ITS2, 28S partial ribosomal RNA genes). Figures show the % divergence between sequences,
names are the places of collection and A and B after the collection number refer to genotype differences between dikaryon
components.

22

Swainsona 31 (2017)

A new species of small black disc fungi, Smardaea australis

Taphrina deformans
0.54r

0.55

0.2

- Rhizina unduiata

- Helvetia leucomelaena

- Morchella esculenta

- Gyromitra californica

- Kallistoskypha incarnata JX560170

0.98

0.96

0.95

5.87

Coprotus ochraceus
Cephaliophora tropica
Geopyxis vulcanalis
Pseudombrophiia theioieuca (2)

Pulvinula convexella (1)

Glaziella aurantiaca

Hum aria hemisphaerica (1)

— Octospora melina

Sowerbyella radiculata
Aleuria aurantia
Scute/iinia scute!lata
Cheilymenia viteiiina
Pyronema conf/uens
Otidea unicisa
Otidea onotica
Otidea bufonia
Otidea alutacea
Warcupia terrestris
0.79j— Smardaea amethystina
1 — Smardaea reticuiosperma

{ PSC1020
PSC4079
PSC4080
PSC3637
PSC4043

r KY067461 "l
||KY067464
1KY067465 k
KY067462
KY067463J

lr Pyropyxisrubra (1)

ULr Jafnea fusicarpa DQ220355
Jafnea semitosta J N 012 012

- Sarcosoma giobosum

- Chorioactis geaster

Pseudopithyeiia minuscuia

0.98

Ascobolus carbonarius
Peziza michelii
Peziza vesiculosa
Peziza gerardii

Marcel leina persoonii
Marceiieina pseudoanthracina _J

T

Taphrinaceae
Caloscyphaceae
Rhizinaceae
Helvellaceae
Morchellaceae
Discinaceae
Boubovia lineage
Ascodesmidaceae
Geopyxis lineage
Pseudobromophila lineage
Pulvinula lineage
Glaziellaceae
Humaria lineage
Octospora lineage
Sowerbellia lineage
Aleuria lineage

Scutellinia-Trichophaea lineage
Cheilymenia lineage
Pyronema lineage

Otideaceae

Sarcosomataceae

Chorioactidaceae

Sarcoscyphaceae

Ascobolaceae

Pezizaceae

Fig. 9. Phylogeny of Pyrenomycetes showing the Smardaea and Marceiieina clades are distantly related. Sequences of the
ribosomal RNA large subunit gene were aligned with those of representatives of each of the currently recognised families and
lineages chosen from those used by Hansen etal. (2013).Tree construction used MrBayes. Figures show the posterior probability of
branches (note that the support for some is less secure than those based on the combined RPB1, RPB2, EF-lcx and LSU sequences
used by Hansen). GenBank accession numbers for the sequences used follow each species name.

globose spores. Although Peziza gerardii and species
of Marceiieina have purple pigments they do not have
these in all tissues whereas all species of Smardaea have
purple pigments in all tissues.

The difficulties in species identification and placement
within phylogenetic relationships of some of the small
disc fungi is exemplified by a species described as a
Marceiieina , Marceiieina parvispora E. Rubio, Tabares
& M.A.Martfnez. However much earlier, in 1917, this
taxon had been described as a species of Caloscypha
Boudier: C. incarnata Duverney & Maine. It has a
whitish to pinkish-violaceous hymenium, is found in
association with Eucalyptus species and occurs in various
parts of the world. After morphological examination and
phylogenetic studies, Pfister et al. (2013) transferred the
taxon into a new genus Kallistoskypha Pfister, Agnello,
Lantieri & LoBuglio in the Caloscyphaceae, a family
adjacent to Pezizaceae. Previously, Caloscypha had been
included in the Pyrenomycetaceae (Pfister et al 2013).

Habitat and trophic mode

Collections of Smardaea australis from the lower
Eyre Peninsula, Port Lincoln and Coffin Bay, South
Australia, and near Anglesea, Victoria, were on sandy
dunes in coastal heath. Those specimens from the more
northern localities in South Australia, i.e. Wilpena
Pound and Bunyeroo Gorge, Flinders Ranges, and
Mambray Creek, Mount Remarkable National Park,
were on clay soil with river red gum, E. camaldulensis

(Myrtaceae), and native pine, C. glaucophylla (Cu-
pressaceae). Soil and vegetation types were thus
different for these similar South Australian collections.
The morphologically similar Smardaea planchonis is
reported as growing mostly in sandy soils, as did coastal
collections of S. australis but unlike those to the north
on heavier soils. The five South Australian specimens of
S. australis were collected from sites up to 460 km apart
and, although they do have divergent ITS sequences,
the diversity appears consistent with a single species.
Molecular sequencing of the Victorian collection was
not attempted due to its age.

Healy et al. (2013) investigated the trophic status of
a high diversity of Pezizales. Within the Otideaceae,
which includes the genera Pyropyxis , Jafnea , Otidea and
Smardaea , they consider that Otidea leporina (Batsch)
Fuckel is ectomycorrhizal. Hansen etal. (2013) consider
S. amethystina and S. reticuiosperma to be saprobic
and comment that this is surprising since P. rubra is
parasitic and differs morphologically by containing
orange, not purple, pigments. A species of Jafnea ,
J. semitosta is considered saprobic (Antonin & Moravec
2010; Kuo 2012). There are therefore ectomycorrhizal,
saprobic and parasitic species within the one family
of Otideaceae. The trophic mode of Smardaea species
is still uncertain. Tedersoo et al. (2006) and Tedersoo
et al. (2010) did not include Smardaea in their study
of fungal ectomycorrhizal lifestyles, but determined
Marceiieina to be mycorrhizal.

23

P.S. Catcheside etal.

Swainsona 31 (2017)

Table 1. Measurements of asci and spores and locality in collections of Smardaea australis from Australia and S. planchonis
worldwide, n.d. = no data

Collection/

Reference

Asci Range
(pm)

Mean Asci
(pm)

Spore Range
(pm)

Mean Spore
(pm)

Spore Q
Range

Mean Spore
Q

Locality

Smardaea australis in South Australia

PSC4079

(holotype)

170-225x8.5-12.5

194.75 X 10.32

8.0-10.5 x 8.0-10.0

9.2 X 9.05

1-1.08

1.0175

Eyre Peninsula

PSC 1020

190-225x

10—12.5(—14)

208.4 X 11.6

8.0-11.5x8.0-11.5

10.1 x 9.87

1-1.05(1.09)

1.023

Flinders Ranges

PSC3637

160—190(—210) X
9.5-12

177.25 X 10.8

8.0-10.5 X 7.5-10.5

9.03 X 8.87

1-1.07(1.09)

1.018

Mambray Creek

PSC 4080

160-200x
(8.5—)9—12.5

176.8 x 10.32

7.5-9.5 X 7.0-9.5

8.62 x 8.43

1-1.1

1.02

Eyre Peninsula

PSC4043

155-195x9.5-12

173.45 X 10.37

8.0-1102x8.0-11.0

9.74 X 9.55

1-1.08

1.02

Flinders Ranges

Smardaea australis in Victoria

MELU F 121706a

(160-)185-210x
10-12

192.6 X 11

9.0-10.5 x 9.0-10.5

9.55 x 9.44

1-1.04

1.025

Anglesea

Smardaea planchonis worldwide

Boudier 1887

135-150X10-12

n.d.

10-11(-12) (diam.)

n.d.

n.d.

n.d.

France

Cuesta & Ribes
2006

193.5- 234 x

8.5- 12.4

213.7x10.5

9.7-11.4x9.4-11.2

10.5X 10.3

1-1.1

1.02

Spain

Marchetti &
Franchi 1993

190-230 X

11—12(—15)

n.d.

9-11 (-12) (diam.)

n.d.

n.d.

n.d.

Tuscany

Haffner 1995

195-240x9.7-12

n.d.

7.9-11.3 (diam.)

n.d.

n.d.

n.d.

Austria

Benkert 2005

200-235 X

11—13(—16)

n.d.

10-12 (diam.)

n.d.

n.d.

n.d.

Germany

Peric 2001

198-228(-230) X

12—13(—15)

n.d.

8.5-11.5 (diam.)

n.d.

n.d.

n.d.

Montenegro

Pfister 1985

215-225 X 11-12

n.d.

8-10 (diam.)

n.d.

n.d.

n.d.

U.S.A.

Gamundf 1960

191-222 X

10.7-14.5

n.d.

None given

n.d.

n.d.

n.d.

Argentina

Lantieri et al. (2009) suggested that S. planchonis is
mycorrhizal. Collections were mostly recorded under
Cupressaceae or Pinaceae, perhaps suggesting that
this species is mycorrhizal with trees of those families.
Smardaea australis cannot be mycorrhizal with conifers
since there were no Callitris species in the Eyre
Peninsula localities. The trophic status of Smardaea
remains unresolved.

In summary

The similarities in morphology and pigmentation of
the Smardaea and Marcelleina species are striking given
the ancient phylogenetic divergence implied by the
28S rDNA sequences (Fig. 9) and other analyses that
infer two distinct lineages (Hansen et al. 2001; Hansen
et al. 2005; Hansen & Pflster 2006; Perry et al. 2007;
Tedersoo et al. 2010; Hansen et al. 2013). These data
suggest cases of convergent evolution, perhaps driven by
exposure to high levels of insolation, where the intense
purple pigmentation would have provided protection
from ionising radiation.

In the absence of molecular data on northern
hemisphere collections of Smardaea planchonis , it is

not possible to determine for certain, whether the
Australian collections are synonymous with those
from the northern hemisphere or Argentina. However,
the shorter asci, variability and relative paleness of the
violaceous pigmentation, absence of probably associated
plant species and the distance from northern hemisphere
and South American collections are consistent with the
Australian collections being a new species.

Acknowledgements

We should like to thank two anonymous referees for
their very thoughtful and helpful comments and
suggestions that improved this manuscript. We are
grateful to the Directors of the State Herbarium of
South Australia (AD), the Queensland Herbarium
(BRI) and the Melbourne University Herbarium
(MELU) for making specimens available for the study.
We also wish to thank Frank Kutsche for assistance
in obtaining relevant scientific collecting permits for
South Australia and Rangers of Conservation and
National Parks for their assistance and interest. We are
very grateful to the State Herbarium of South Australia
and Flinders University for their continual support.

24

Swainsona 31 (2017)

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South Australia at all times, the content of this publications is licensed under the Creative Commons Attribution 4.0 Licence
(https://creativecommons.Org/licenses/by/4.0/). All other rights are reserved.

© 2017 Board of the Botanic Gardens and State Herbarium (Adelaide, South Australia)

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26

w

Swainsona 31:27-30 (2017)

© 2017 Board of the Botanic Gardens & State Herbarium (Adelaide, South Australia) ©0

Tephromela baudiniana sp. nov. (lichenised Ascomycetes) from Kangaroo
Island

Gintaras Kantvilas a & John A. Elix 6

a Tasmanian Herbarium, PO Box 5058, UTAS LPO, Sandy Bay, Tasmania 7005
Email: Gintaras.Kantvilas@tmag.tas.gov.au

b Research School of Chemistry, Australian National University, Canberra, Australian Capital Territory 2601
Email: John.Elix@anu.edu.au

Abstract: A new lichen species, Tephromela baudiniana Kantvilas & Elix, is described from coastal rocks
on Kangaroo Island, South Australia. It is characterised chiefly by its unique chemistry, comprising the
rare metabolites, 9-O-methylalternariol and alternariol. The species is compared with other saxicolous
species of the genus occurring in Australia.

Keywords: alternariol, biodiversity, lichens, new species, South Australia, Tephromelataceae

Introduction

This paper is a further contribution to the
documentation of the lichens of Kangaroo Island, South
Australia (Elix & Kantvilas 2013a, b; Kantvilas 2016;
Kantvilas & Elix 2014; Kantvilas & Kondratyuk 2013;
Kantvilas & van den Boom 2013, 2015; Kantvilas &
Wedin 2015; McCarthy & Kantvilas 2013a, b, 2016
a, b), and describes a new species of the cosmopolitan
genus Tephromela M.Choisy (Tephromelataceae).
Tephromela is characterised by a crustose thallus with
a unicellular, trebouxioid photobiont, apothecial
ascomata, eight-spored, Biatora- to Lecidella- type
asci, and simple, hyaline, non-halonate ascospores;
the hymenium is invariably pigmented a diagnostic
crimson, referred to as atra- red by Meyer & Printzen
(2000). Detailed descriptions and illustrations of the
genus and a discussion of its affinities are provided by
Fryday (2011), Spribille et al. (2011), Kantvilas (2015)
and many others, and are not repeated here. Species
of Tephromela may be lichenicolous or occur free-
living on a wide range of substrata, including rocks,
bark and wood. They tend to be defined by thalline
characters (e.g. presence or absence of soredia, etc.),
apothecial anatomy and morphology, ascospore size
and thallus secondary chemistry. The apothecia can be
highly variable, ranging from immarginate to biatorine
to “lecanorine”. The seemingly lecanorine apothecial
margin, a feature of the many species ascribed to the
T. atra group, has been interpreted by Spribille et al.
(2011) and Kantvilas (2015) as a thalline cushion that
subtends the apothecium and may envelop it at the
margins.

Variation in the widespread T. atra group was
investigated at a global scale by Muggia et al. (2013),
who found that DNA sequence data do not support

some of the phenotypically distinct entities that have
been described as species. They considered that some of
the variability with respect to apothecial and ascospore
size, substratum preference and thallus chemistry may
be better interpreted as locally developed features that
are part of the infra-specific diversity of one highly
variable taxon. Subsequently, however, Cestaro et al.
(2016) demonstrated that at least in one segregate of
T. atra , the North American species 77 pacifica Bjork
& Muggia, differences in chemical composition were
underpinned by phylogenetic data. In Australia (and
elsewhere), variation, particularly chemical variation,
continues to be treated at species rank (see Kalb
2004, 2008; Elix 2012, 2013a, b; Elix & Kalb 2006,
2008) and its taxonomic importance requires further
detailed investigation. The new species described below
represents a further example of this variation.

Material and methods

The study is based on collections by the first author
housed in the Tasmanian Herbarium (HO). Anatomical
and morphological observations were undertaken using
light microscopy, with thin hand-cut sections mounted
in water, 10% KOH, 50% HNO 3 , lactophenol cotton
blue, ammoniacal erythrosin and Lugol’s iodine, with
and without pretreatment with KOH. Ascospore
measurements are based on 30 observations and are
presented in the format: least value— average—
value. Chemical composition was investigated by thin-
layer chromatography using standard methods (Orange
et al. 2001; Elix 2014). Nomenclature of ascus types
essentially follows Hafellner (1984) and Kantvilas
(2009); that of pigments follows Meyer & Printzen
( 2000 ).

Published online: 15 June 2017 • flora.sa.gov.au/swainsona

ISSN 2206-1649 (Print) • ISSN 2206-1657 (Online)

G. Kantvilas & J.A. Elix

Swainsona 31 (2017)

Tephromela baudiniana Kantvilas & Elix, sp. nov.

Tephromelae atrae (Hudson) Hafellner similis sed
9- O-methylalternariolum et alternariolum, substantias
chemicalias adhuc in Tephromela ignotas, continenti
differt.

Typus: South Australia, Kangaroo Island: Rocky
River Track, c. 250 m from mouth of river, 35°58'S
136°39'E, 10 m alt., on coastal rocks on the bank of
a fast-flowing fresh-water stream, 25 Sep. 2015, G.
Kantvilas 511/15 & B. de Villiers (holo: HO 581706;
iso: AD).

Mycobank number: MB820508.

Thallus free-living, markedly rimose-areolate, whitish
grey, esorediate, forming extensive, irregular, rather
discontinuous patches to 10 cm or more in extent,
essentially comprised of numerous irregularly circular
thalli c. 10 mm wide that coalesce; individual areoles
irregularly rhomboid, 0.4—1.5 mm wide, 0.7—1 mm
thick, plane, verruculose or bullate; prothallus not
developed, but actively expanding thallus margins
often discoloured dark bluish grey at the very edge.
Apothecia scattered, roundish, basally constricted,
“lecanorine”, 0.5-1 mm wide; disc black, glossy, plane
to undulate to convex, epruinose; thalline “margin”
typically prominent but occasionally not developed,
entire, often slightly inrolled, persistent, concolorous
with the thallus, in section to 100-200 pm thick.
Proper excipulum to 20-60 (-100) pm thick, thickest
at the base, becoming excluded at the edges, pale to
intense yellow-brown, K± intensifying yellow, N+
intense brownish orange. Hypothecium to 20-45 pm
thick, hyaline to pale yellowish, thickest in the centre
of the apothecium, tapering towards the margins and
sometimes ± excluded. Hymenium 90-130 (-170) pm
thick, not inspersed or with occasional oil droplets,
crimson red, K+ intensifying red, N+ orange (< atra -
red), persistently coherent in water, K and N, with

an intensely atra- red pigmented epihymenial layer.
Paraphyses 2-3 pm thick, stout, simple; apices not or
gradually expanded to 6 pm wide, with a thick coat of
gel and heavily caked and conglutinated with pigment.
Asci clavate, 43—60 x 12—18 pm, of the Biatora- type:
having a well-developed amyloid tholus, a masse axiale
with ± parallel flanks and a rounded apex, and a darker,
more intensely amyloid edge adjacent to the masse
axiale. Ascospores hyaline, ellipsoid, 8-10.7-13 x 4.5-
6.4— 9 pm. Pycnidia not found. Fig. 1.

Chemistry. Thallus K+ yellow, C-, PD-, UV+ blue;
containing atranorin (minor), 9-O-methylalternariol
(major) and alternariol (minor).

Etymology. The specific epithet commemorates the
French navigator Nicolas Baudin who made the
second European landing on Kangaroo Island in 1802
(including just to the north of type locality) and who
charted and named many of the island’s features.

Remarks. Although Muggia et al. (2013) questioned
the rigorous application of chemical characters in
the delimiation of Tephromela species, this criterion
underpins the taxonomy of the genus in Australia
where 27 species are recorded (McCarthy 2016);
see Elix (2013a) for an identification key. It is the
unusual secondary chemistry of the new species
that distinguishes it unequivocally from other,
morphologically highly similar, saxicolous species
(Table 1). Tephromela baudiniana is chemically unique
within the genus. Alternariol and 9-O-methylalternariol
occur very rarely as major metabolites in lichens and are
known as such only in Pertusaria praecipua A.W.Archer
& Elix from Papua New Guinea (Archer & Elix 1998).
This chemical character is highly significant. Alternariol
is derived biosynthetically from a single, non-reduced

Fig. 1 . Tephromela baudiniana (holotype). A Habit, scale = 2 mm; B Detail, scale = 1 mm.

28

Swainsona 31 (2017)

Tephromela baudiniana sp. nov. from Kangaroo Island

Table 1. Salient features of saxicolous, free-living species of Tephromela in Australia.

secondary chemistry (key
compounds)

asexual

propagules

apothecial

morphology

ascospore size

distribution

T. arafurensis Rambold

atranorin, perlatolic and
glomelliferic acids

esorediate

± immarginate to
"lecanorine"

9-14x5-7 pm 1

northern Australia
(NT, WA)

T. atra (Huds.) Hafellner

atranorin, a-collatolic, alectoronic
and bourgeanic (±) acids

esorediate

"lecanorine"

11-17x5.5-9 pm 2

widespread,

cosmopolitan

T. baudiniana Kantvilas & Elix

atranorin, alternariol,
9-O-methylalternariol

esorediate

"lecanorine"

8-13x4.5-9 pm

Kangaroo Is (SA)

T. buelliana (Mull.Arg.) Kalb

atranorin, a-collatolic acid

± immarginate to
"lecanorine"

10-14x5.5-9 pm 3

SA, Vic, Brazil

T. granularis Kantvilas

atranorin, a-collatolic, alectoronic
and bourgeanic (±) acids

sorediate

"lecanorine"

10-15x5-10 pm 2

Tas

T. korundensis (Rasanen) Kalb

atranorin

esorediate

± immarginate

6.5-12x6-9 pm 1

NEQId

T. lillipillensis Elix

atranorin, alectoronic acid

esorediate

"lecanorine"

7.5-14x6-7.5 pm 1

NSW, Qld

T. promontorii (Zahlbr.) Kalb

atranorin, a-collatolic and
alectoronic acids

esorediate

± immarginate

7-13x5.5-7 pm 4

SA, NT, Vic, South
Africa

*T. skottsbergii (Darb.) Fryday

atranorin, a-collatolic and
alectoronic acids

esorediate

"lecanorine"

10-17x5-8 pm 1

Qld, NSW, Chile

T. stenosporonica Elix & Kalb

atranorin, stenosporonic acid

esorediate

"lecanorine"

11-13x7-9 pm 1

northern Australia
(NT, WA)

T. territoriensis Elix & Kalb

atranorin, physodic acid

esorediate

"lecanorine"

7.5-8 x 5-6.5 pm 1

NT

Data sources. 1 Elix (2009); 2

Kantvilas (2015); 3 Vainio (1890); 4

1 Elix (2013a).

*T. skottsbergii is characaterised further by having an hymenium inspersed with oil droplets.

polyketide chain, whereas a-collatolic and alectoronic
acids, found in the widespread, morphologically
similar T. atra, are derived from two partially reduced
polyketide chains (Armaleo et al. 2011; Saha et al.
2012 ).

The new species has proved to be technically
challenging to study. Although seemingly abundantly
fertile, most apothecia sectioned proved to be devoid of
mature asci or almost so, and literally dozens of sections
were required to compile the observations necessary for
a comprehensive description. Kantvilas (2015) noted
that the ascospores of Tephromela may vary significantly
in size, depending on their stage of development. In
the present work, most measurements were of relatively
young spores still within the ascus, and so their
dimensions may have been underestimated. The thallus
of the new species included numerous crystals detached
from the rock substrate, which protrude through the
upper surface or are enclosed within the medulla. We
have not observed this in previous studies of the genus,
even in collections from highly crystalline rocks such as
granite, and it is not clear whether this is a product of
the species or the substrate. No pycnidia were located,
and the abundant speck-like features on the surface of
the thallus proved to be mainly apothecial initials or
crystalline inclusions.

Morphologically, T. baudiniana falls within the very
broad range of variation exhibited by Tephromela atra ,
which is characterised by containing atranorin plus
a-collatolic and alectoronic acids, usually together with
bourgeanic acid. However, the very “chinky” rimose
thallus, the absence of a prothallus and the relatively

small apothecia place it at the extreme of variation for
that species.

Distribution and ecology. The new species is known
only from the type locality where it was locally very
abundant. It grew on dry, exposed siliceous rocks lining
the course of a fast-flowing fresh water stream, but
close to its mouth and therefore also subject to marine
influences. Thus associated lichen species included truly
littoral species such as Caloplaca gallowayi S.Y.Kondr.,
Karnefelt & Filson, Catillaria austrolittoralis Kantvilas
& van den Boom and Opegrapha spodopolia Nyl., as well
as species more typical of inland, non marine habitats,
for example, Xanthoparmelia australasica D.J.Galloway
and Amandinea spp.

Acknowledgements

We sincerely thank Dr Ernest Lacey, Microbial
Screening Technologies, Smithfield, NSW, and
Dr Andrew Piggott, Department of Chemistry &
Biomolecular Sciences Macquarie University, NSW, for
assistance with HPLC/MS analysis of an extract of this
species. Dr Jean Jarman provided the photographs that
illustrate this paper.

References

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(lichenised Ascomycotina) in Papua New Guinea: three new
species and two new reports. Mycotaxon 69: 311-318.
Armaleo, D., Sun, X. & Culberson, C.F. (2011). Insights from
the first putative biosynthetic gene cluster for a lichen
depside and depsidone. Mycologia 103: 741-754.

29

G. Kantvilas & J.A. Elix

Swainsona 31 (2017)

Cestaro, L., Tonsberg, T. & Muggia, L. (2016). Phylogenetic
data and chemical traits characterize a new species in the
lichen genus Tephromela. Herzogia 29: 383-402.

Elix, J.A. (2009). Tephromela. Flora of Australia 57: 74-83.

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(lichenized Ascomycota) from Australia. Australasian
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Ascomycota) from Australia. Australasian Lichenology 73:
45-54.

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chromatographic data and biosynthetic relationships for lichen
substances, 3rd edn. (Published by the author: Canberra).

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(Lecanoraceae, lichenized Ascomycota) from Australia.
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Elix, J.A. & Kalb, K. (2008). Additional new lichen taxa
(lichenized Ascomycota) from Australia. Australasian
Lichenology 63: 30-36.

Elix, J.A. & Kantvilas, G. (2013a). New taxa and new records
of Amandinea (Physciaceae, Ascomycota) in Australia.
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Elix, J.A. & Kantvilas, G. (2013b). New taxa and new records
of Buellia sensu lato (Physciaceae, Ascomycota) in Australia.
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Calvitimela and Tephromela from the southern subpolar
region. Lichenologist 43: 225-239.

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Gliederung der Sammelfamilien Lecanoraceae und
Lecideaceae. Beihefte zur Nova Hedwigia 79: 241-371.

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Kalb, K. (2008). New or otherwise interesting lichens. IV.
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Kantvilas, G. (2009). The genus Mycoblastus in the cool
temperate Southern Hemisphere, with special reference to
Tasmania. Lichenologist 41: 151-178.

Kantvilas, G. (2015). Observations on the genus Tephromela
(lichenised Ascomycetes) in Tasmania, with the description
of a new species. Herzogia 28: 430-444.

Kantvilas, G. (2016). A synopsis and key for the lichen genus
Caloplaca on Kangaroo Island, with the description of two
new species. Journal of the Adelaide Botanic Gardens 29:
53-69.

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Kantvilas, G. & Kondratyuk, S.Y. (2013). New species of
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some calcicolous species of Lecania A.Massal. (lichenised
Ascomycetes: Ramalinaceae) in southern Australia. Journal
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the Ascomycete genus Arthonia Ach. from Kangaroo Island.
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its island territories. (Australian Biological Resources Study:
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2017].

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turina sp. nov. (lichenized Ascomycota: Verrucariaceae) from
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sp. nov. (lichenized Ascomycota: Verrucariaceae) from
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© 2017 Board of the Botanic Gardens and State Herbarium (Adelaide, South Australia)

30

w

Swainsona 31:31-36 (2017)

© 2017 Board of the Botanic Gardens & State Herbarium (Adelaide, South Australia) ©0

Two species of Bacidia De Not. with pruinose apothecia from Kangaroo
Island

Gintaras Kantvilas

Tasmanian Herbarium, PO Box 5058, UTAS LPO, Sandy Bay, Tasmania 7005
Email: Gintaras.Kantvilas@tmag.tas.gov.au

Abstract: Two species of the lichen genus Bacidia (Ramalinaceae) with pruinose apothecia from
Kangaroo Island, South Australia, are described and illustrated: B. septosior (Nyl.) Zahlbr., a long-
overlooked species that is widespread in south-eastern Australia, and B. brigitteae Kantvilas, described
as new and known only from Kangaroo Island where it occurs in coastal /We/a/euca-dominated
woodland.

Keywords: taxonomy, lichens, South Australia, Ramalinaceae

Introduction

Bacidia De Not. (Ramalinaceae) is a large, cosmopolitan
genus of lichens that occurs in virtually all vegetation
and climatic zones of the world, ranges from sea-level
to alpine elevations, and colonises a wide variety of
substrata, including rock, wood, bark, soil and living
leaves. The genus is well represented in Australia where
33 names in Bacidia have been recorded (McCarthy
2016), many based on Australian types, and all the
major herbaria hold large numbers of specimens that
are mostly unidentified or misidentified.

For decades, Bacidia in the traditional sense served
as a useful “form” genus for crustose lichens with a
green photobiont, biatorine or lecideine apothecia,
eight-spored asci and hyaline, transversely multi-
septate ascospores. However, as has occurred with most
other form genera, such as Buellia De Not., Catillaria
A.Massal., Lecanora Ach. and Lecidea Ach., new insights
and the application of characters such as excipulum
structure and, in particular, ascus type (Hafellner 1984)
have led to the gradual segregation out of Bacidia of
smaller, more natural units, sometimes classified in
different families such as the Pilocarpaceae. Some of
these smaller genera that occur in the Australian flora
and that accommodate taxa formerly included under
Bacidia include Arthrorhaphis Th.Fr. (Obermayer
2001), Bacidina Vezda (Vezda 1990), Bapalmuia Serus.
(Kalb et al. 2000), Bryobilimbia Fryday, Printzen &
S.Ekman (Fryday et al. 2014), Fellhanera Vezda (Vezda
1990), Fellhaneropsis Serus. & Coppins (Kantvilas &
Lucking 2009), Jarmania Kantvilas (Kantvilas 2008),
Sarrameana Vezda & P.James (Kantvilas & Vezda
1996) and Scoliciosporum A.Massal. (Kantvilas 2008).

In recent decades, there have been considerable
advances in the study and delimitation of Bacidia ,
mainly in the Northern Hemisphere, and a large

body of literature pertaining to individual species or
local regions now exists. Significant contributions of a
broader application, and ones that assist understanding
the species found in Australia, include the account for
North American corticolous species (Ekman 1996), and
flora treatments or keys for the Iberian Peninsula (Llop
& Hladun 2002), Great Britain (Coppins & Aptroot
2009), the Sonoran Desert (Ekman 2004) and North
America (B. McCune, unpublished mss). However, in
Australia, the genus remains extremely poorly known
and is, in some respects, the last of the “big genera”
demanding detailed study and revision. It is noteworthy
that since the resurgence of lichenology from the mid-
20 th Century onwards, the only contributions on
Bacidia in Australasia remain the Flora account for New
Zealand by Galloway (2007), and descriptions of new
species by Elix (2009) and Coppins & Fryday (2007).
In this authors considered view, the many taxa listed by
McCarthy (2016) represent only the very tip of a very
large and complex iceberg.

For some years, the author has been gradually
examining the types and reliably identified specimens
of many Bacidia species in the major herbaria of the
world, with the intention of bringing some order to
those taxa that occur in southern Australia, especially
Tasmania and Kangaroo Island, where most of his
taxonomic studies have been focused. This work has
proved to be extremely challenging due to the large
numbers of species and specimens involved, and the
subtle variation in critical species-level characters such
as apothecial pigmentation, excipulum structure and
ascospore dimensions. A comprehensive account for
the region is clearly some way off, even though a few
particular species have now been resolved. In this paper,
two species with pruinose apothecia, a feature that is
rather restricted in the genus, are treated; the work
represents the first hesitant steps towards a broader
study.

Published online: 15 June 2017 • flora.sa.gov.au/swainsona

ISSN 2206-1649 (Print) • ISSN 2206-1657 (Online)

G. Kantvilas

Swainsona 31 (2017)

Material and methods

The study is based mainly on collections of the author
housed in the Tasmanian Herbarium (HO), with
additional material sourced from other herbaria as
indicated in text. Anatomical and morphological
observations were undertaken using light microscopy,
with thin hand-cut sections mounted in water, 10%
KOH, 50% HNO 3 , Lactophenol Cotton Blue,
ammoniacal erythrosin and Lugols Iodine, with
and without pretreatment with KOH. Ascospore
measurements are based on at least 50 observations
for each taxon and are presented in the format: 5 th
percentile—/awrag?—95 th percentile, with outlying values
given in brackets. Chemical composition was investigated
by thin-layer chromatography using standard methods
(Orange et al. 2001). Nomenclature of apothecial
pigments follows Ekman (1996).

The species

Bacidia septosior (Nyl.) Zahlbr.

Catal. Lich. Univ. 4: 240 (1926); Lecidea septosior
Nyl. in Crombie, J. Linn. Soc., Bot. 17: 400 (1880).
Typ e: [Australia, New South Wales] on bark of trees,
near Port Jackson, R. Brown 562 (holo: BM!).

Thallus crustose, whitish grey to grey green, rather
scurfy, smooth to somewhat verruculose, or, more
commonly, patchy and evanescent, to 40-150 pm thick,
forming irregular, undelimited patches to 15 cm wide
or more; prothallus absent; photobiont chlorococcoid,
with cells globose, 7-13 pm diam. Apothecia biatorine,
0.6-1.6 mm diam., scattered, urceolate and strongly
basally constricted to almost substipitate when young,
soon becoming disciform; disc plane at first, often

persistently so, or sometimes becoming convex, black
to brown-black, matt, occasionally bluish grey pruinose
when very young but soon becoming epruinose; proper
excipulum persistent, sometimes becoming reduced
in the oldest, most convex apothecia, with outer parts
glossy red-brown and the upper edge intensely bluish
grey pruinose when young, later becoming brown-black
to black and the pruina restricted to the upper inner edge
or absent, in section cupulate, 90-160 pm thick, at the
upper edge red-brown to brown, K± purple-brown, N+
orange, internally ± colourless, composed of radiating,
branched and anastomosing hyphae 1-1.5 pm thick in
a gel matrix, lacking enlarged terminal cells, inspersed
with bands of angular crystals to c. 5 pm wide that
are insoluble in K. Hypothecium 140-200 pm thick,
colourless throughout, or pale yellowish, intensifying
yellowish in K and N in the upper part. Hymenium
120-160 pm thick, sparsely inspersed with oil droplets;
epithecium grey-green, brownish green or brownish,
K± intensifying greenish, N+ purple, sometimes rather
fleetingly so and then fading to pale orange, also often
slowly forming a precipitate of minute, dark blue
crystals; paraphyses 1.5-2 pm thick, simple, not or
only weakly capitate, ± separating readily in K, but with
apices remaining conglutinated in epithecial pigment
and sometimes expanded to 3 (-4) pm wide; asci of
the Bacidia- type, narrowly cylindrical, with an amyloid
tholus not entirely pierced by a ± non-amyloid, conical
masse axiale, 90-110 x 14-20 pm. Ascospores arranged
side-by-side or very loosely coiled in the ascus, filiform
to narrowly cylindrical with rounded apices, easily
fractured, (55-) 64.5-77.6-96 (-104) x 4-4.5- 5.5
(-6) pm, 20-35-septate. Pycnidia very rare, immersed;
conidia filiform, curved, aseptate, 15-20 x 0.5 (-1) pm.
Chemistry: no substances detected by t.l.c. Figs 1, 2B.

Fig. 1. Bacidia septosior

habit (Kantvilas 261/13).
Note the youngest
(smallest) apothecia with a
characteristically pruinose
inner margin. Scale = 5 mm.

32

Swainsona 31 (2017)

Two species of Bacidio from Kangaroo Island

Note on the type. The sheet in BM bearing Brown's
type specimen also has a second collection in very poor
condition, annotated in the same hand with “on the
bark of tree, S. Coast of Tasmania, R. Brown”.

Discussion. This highly distinctive species is
characterised by the presence of crystals in the
excipulum in combination with the pruinose apothecial
margin and the particular suite of apothecial pigments.
The pruina are best seen in young apothecia, and older
apothecia may be entirely epruinose or with the pruina
reduced to just the inner upper edge of the excipulum.
Whilst some specimens may be dominated by black
epruinose mature apothecia, careful examination
will usually reveal vestiges of pruina on at least some
apothecia. The crystalline inclusions in the excipulum
are invariably present although rarely they may be few in
number. The excipular pigment is laurocerasi- brown of
Ekman (1996). The epithecial pigment is more variable
but, in most collections, the dominant colour in water
is greenish or greenish grey or greenish brown. With
the addition of KOH, the pigment remains unchanged
or the greenish hints intensify, whereas with HNO 3 , it
turns a vivid purple, then slowly fades to a pale orange;
if this pigment is sufficiently concentrated, a precipitate
of minute dark blue crystals may develop. The pigment
appears to match Bacidia- green II of Ekman (1996).

Whilst this species, as defined above, is extremely clear-
cut, there are herbarium collections, particularly from
Tasmania, that have essentially identical apothecial
pigments and ascospores, but lack both the apothecial
pruina and the crystalline excipular inclusions; such

specimens are excluded from this species and require
further study. The most similar species to B. septosior
(Nyl.) Zahlbr., in that it has pruinose apothecia with
crystals in the excipulum, is B. brigitteae , which is
described and discussed below. Also extremely similar
is Bacidia ceratina (Stirt.) Hellb., described from
New Zealand (lectotype in GLAM examined). This
name was synonymised incorrectly with B. laurocerasi
(Delise ex Duby) Vain, by Galloway (1985). Although
B. laurocerasi and B. ceratina have some superficial
features in common, the latter has crystalline excipular
inclusions and occasionally pruinose apothecial margins
identical to what is seen in B. septosior. Interestingly,
neither of these features is noted in the original
description of Stirton (1875). I have considered the
possibility that B. ceratina may be an earlier name for
B. septosior , but have declined to synonymise these
taxa. Brown’s Australian type of B. septosior matches
subsequent collections of this species exactly. In
contrast, the New Zealand type specimen of B. ceratina
has a very thick, papillate-verruculose thallus that
has never been observed in Australian specimens.
Furthermore, no well-developed ascospores could be
found in the B. ceratina specimen, but the hymenium
is consistently thinner than that of B. septosior , and
therefore presumably the ascospores may well be
shorter.

Distribution and ecology. Bacidia septosior appears to
be widespread in lowland, especially coastal areas of
south eastern Australia (New South Wales, Victoria,
Tasmania, the Bass Strait Islands and Kangaroo Island),
where it occurs mostly on soft, spongy bark, such as on

Fig. 2. Anatomy of Bacidia, showing typical, Bacidia -type asci and filiform, multi-septate ascospores. A B. brigitteae;
B B. septosior. Scale = 20 pm.

33

G. Kantvilas

Swainsona 31 (2017)

Fig. 3. Bacidia brigitteae (isotype). A Habit, showing persistently pruinose apothecia. Scale = 5 mm. B Detail, showing the
characteristic, granular to goniocyst-like thallus. Scale = 1 mm.

the bases of eucalypts, Callitris , species of Melaleuca , or
shrubs of the Asteraceae such as Olearia or Bedfordia. It
appears to prefer marginally moister conditions, such as
swampy Melaleuca-dommdMtd woodland or older trees
in locally damper microhabitats within dry eucalypt
woodland. It is usually part of a rich association of
epiphytic lichens, the more common of which include
Bacidia brigitteae Kantvilas, Megalaria grossa (Pers. ex
Nyl.) Hafellner, Menegazzia caesiopruinosa P.James,
M. subpertusa P.James & D.J.Galloway, Pannaria
elixii P.M.Jorg. & D.J.Galloway, P. obscura Mull.
Arg., Parmotrema perlatum (Huds.) M.Choisy, Physcia
poncinsii Hue and Pseudocyphellaria aurata (Ach.) Vain.

Specimens examined

SOUTH AUSTRALIA, Kangaroo Island: Cape du Couedic
Road, 1 km S of Rocky River Settlement, 35°57'S 136°44'E,
70 m alt., 1994, H. Streimann 54997 (AD, CANB); Willson
River, 35°52'S 137°56'E, 1997, R.J. Bates 48365 (AD);
Moffatt Road, 35°49'S 138°00'E, 70 m alt., 2011, G.
Kantvilas 251/11 & B. de Villiers (HO); West Bay, 35°53'S
136°33'E, 10 m alt., 2011, G. Kantvilas 299/11 (AD, HO);
Creek Bay Farm, headwaters of Lubra Creek, 35°49'S
138°06'E, 40 m alt., 2011, G. Kantvilas 379/11 (AD, BM,
HO); Brown Beach, 35°48'S 137°50'E, 10 m alt., 2012,
G. Kantvilas 435/12 & B. de Villiers (AD, CANB, HO);
North Cape Road, 35°36'S 137°35'E, 5 m alt., 2013, G.
Kantvilas 261/13 & B. de Villiers (HO); the Old Cannery,
American River, c. 1 km SW of Ballast Head, 35°46'S
137°48'E, 3 m alt., 2013, G. Kantvilas347/13 &B. de Villiers
(AD, HO); Rocky River, 35°56'S 136°37'E, 10 m alt., 2015,
G. Kantvilas 518/15 (HO); South-West River, 36°01'S
136°52'E, 10 m alt., 2015, G. Kantvilas 423/15 (HO).
VICTORIA: Furnells Landing, 20 km S of Cann River
township, 37°43'20"S 149°08T4"E, 1 m alt., 2008 G.
Kantvilas 155/08 &J.A. Elix (HO).

TASMANIA: southern coast, [1804], R. Brown s.n. (BM);
Marrawah to Montague Track, 40°48'S 144°50'E, 30 m
alt., 1986, G. Kantvilas 113/86 (HO); Swan Basin, 42°12'S
145°16'E, sea-level, 2000, G. Kantvilas 32/00 (HO);
Buckland Military Training Area, S of Bluestone Tier,
42°30'S 147°48'E, 300 m alt., 2003, G. Kantvilas 325/03

(HO); eastern side of Stanley Hwy, 40°47'S 145°16'E, 5 m
alt., 2011, G. Kantvilas 452/11 (HO); end of Bolduans Road,
40°47'S 145°02'E, 1 m alt., 2011, G. Kantvilas 474/11 (HO);
Stony Point, 40°45'S 144°59'E, 2 m alt., 2016, G. Kantvilas
275/16 (HO). Bass Strait, Flinders Island: Patriarchs Inlet,
39°57’S 148°11’E, 1983,/.S. Whinray s.n. (HO, MEL).

Bacidia brigitteae Kantvilas, sp. nov.

Bacidiae septosiori similis et item excipulo crystallis
angularibus insperso sed thallo granuloso, apotheciis
persistente griseo-pruinosis, hymenio tenuiore, 80—
90 pm crasso, ascis parvioribus, 60-75 pm longis, 12-
15 pm latis, et ascosporis brevioribus angustioribusque,
43-80 pm longis, 2.5-4 pm latis, modo 8-18-septatis
praecipue dijfert.

Typus: South Australia, Kangaroo Island: Grassdale
Lagoon, 36°00'S 136°53'E, 20 m alt., on Melaleuca in
mallee woodland, 4 Oct. 2015, G. Kantvilas 326/15 &
B. de Villiers (holo: HO 580598; iso: AD).

Mycobank number: MB821079.

Thallus granular or ± goniocyst-like, bright to dull
green, less commonly becoming evanescent and
endophloeodal, forming irregular, often discontinuous,
undelimited patches to 10 cm wide or more;
individual granules crowded together or dispersed,
30-60 (-80) pm wide, sometimes somewhat isidioid
in appearance, comprising tightly clustered photobiont
cells in a sheath of rather short-celled hyphae 2.5-5 pm
wide; prothallus absent; photobiont a unicellular
green alga with globose cells 5-12 pm diam. Apothecia
biatorine, 0.5-1.5 mm diam., scattered, superficial,
basally constricted, disciform; disc mostly plane or
occasionally becoming undulate or convex, pale brown
but persistently and thickly pale grey pruinose from the
earliest stages; proper excipulum persistent, somewhat
inrolled when young and usually remaining ± taller
than the disc, only rarely excluded when the disc is
highly convex, at first pale grey to pale brown, soon
becoming brown to dark brown to black, persistently
grey pruinose in the upper part and concolorous
with the disc, on the underside epruinose, in section

34

Swainsona 31 (2017)

Two species of Bacidia from Kangaroo Island

cupulate, 50—110 pm thick laterally, 90—150 pm
basally, colourless within but sometimes pale brownish,
K-, N± pale orange at the upper edge, composed of
radiating, anastomosing hyphae 2.5-3.5 pm thick in
a gel matrix, lacking enlarged terminal cells, inspersed
with clusters of angular, rectangular or rhomboidal
crystals up to 10 pm across that do not dissolve in K
but dissolve in N. Hypothecium 100-180 pm thick,
colourless throughout or pale yellowish grey in the
upper part, weakly intensifying yellowish in K and N.
Hymenium 80-90 pm thick, colourless, occasionally
sparsely inspersed with oil droplets, overlain by a
greyish brown epithecium, K+ purplish brown, N+
crimson and fading; asci of the Bacidia- type, elongate-
cylindrical, with an amyloid tholus not entirely pierced
by a ± non-amyloid, conical masse axiale, 60-75 x
12-15 pm; paraphyses simple or sparsely branched,
1.5 (-2) pm thick, with apices not capitate. Ascospores
filiform with blunt or acute apices, side-by-side or,
less commonly, coiled in the ascus, (43-) 50 —60.1—77
(-80) x (2.5-) 3-3.3-4 pm, 8-18-septate. Pycnidia not
found. Chemistry: no substances detected by t.l.c. Figs
2A, 3.

Etymology. The new species is named in honour
of Brigitte de Villiers, who has accompanied me
throughout my excursions on Kangaroo Island,
with great patience strongly supported my lichen
investigations there, and co-collected the type as well as
many other specimens.

Remarks. When first collected, this taxon was
shoehorned, with considerable reservations, into
B. septosior on the basis of having apothecia with an
excipulum inspersed with crystals, and generally filiform,
multiseptate ascospores. Morphological differences
such as the persistently pruinose, disciform rather
than somewhat urceolate apothecia and the granular
thallus were initially considered unusual variations of
B. septosior. The opportunity to observe this species in
the field and make subsequent, rich collections soon
confirmed that this taxon was indeed distinct, and that
the distinctive morphological thalline and apothecial
characters mentioned above were consistently evident.
There are further anatomical differences between the
two species. In B. brigitteae, the hymenium is always
thinner, with concomitantly shorter asci. Furthermore,
the ascospores are consistently shorter and narrower,
with fewer septations. Apothecial pigmentation
appears superficially rather similar and at least some
of the described differences may be due to differences
in pigment concentration, especially with respect to
the excipulum. However, the epithecial pigment of
B. brigitteae is different but difficult to characterise:
there is no hint of greenish coloration in either water
or KOH, yet it reacts crimson in HNO 3 , but with no
subsequent development of a dark blue precipitate.

According to Ekman (1996), a goniocyst-like thallus
is unusual in Bacidia , and is more characteristic of the
genus Bacidina. Yet the new species lacks the expanded,
pseudoparenchymatous hyphae at the outer edge of the

excipulum that characterise Bacidina , and appears to
belong comfortably in Bacidia.

Distribution and ecology. Bacidia brigitteae is known
only from Kangaroo Island, where it grows on the soft,
furrowed bark of Melaleuca in coastal old growth mallee
woodland or Melaleuca-6omm2Ltz<l woodland, in close
proximity to rivers and lagoons where, presumably,
humidity levels are higher, at least intermittently.
Associated species include Bacidia septosior , Caloplaca
kaernefeltii S.Y.Kondr., Elix & A.Thell, Leptogium
crispatellum Nyl., Pannaria obscura Miill.Arg., Physcia
rolandii Elix, Psoroglaena halmaturina P.M.McCarthy
& Kantvilas, Rinodina australiensis Miill.Arg. and
Strangospora pinicola (A.Massal.) Korb.

Specimens examined

SOUTH AUSTRALIA, Kangaroo Island: Ravine des
Casoars, along riverbank, c. 0.5 km inland from coast,
35°48'S 136°35'E, 15 m alt., 2012, G. Kantvilas 482/12&B.
de Villiers (AD, HO); South-West River, 36°01'S 136°52'E,
10 m alt., 2015, G. Kantvilas 426/15 & B. de Villiers (AD,
HO).

Acknowledgements

I thank Jean Jarman for the photographs and preparing
the illustrations for publication. The opportunity
to study the holdings of Bacidia at Londons Natural
History Museum is gratefully acknowledged, as are
loans from there, GLAM and G.

References

Coppins, B.J. & Aptroot, A. (2009). Bacidia De Not. (1846).
In: Smith, C.W., Aptroot, A., Coppins, B.J., Fletcher, A.,
Gilbert, O.L., James, P.W. & Wolseley P.A. (eds) The lichens
of Great Britain and Ireland, pp. 189-207. (British Lichen
Society: London).

Coppins, B.J. & Fryday, A.M. (2007). Three new species of
Bacidia s.lat. (Ramalinaceae) from Campbell Island, New
Zealand. Bibliotheca Lichenologica 95: 155-164.

Ekman, S. (1996). The corticolous and lignicolous species of
Bacidia and Bacidina in North America. Opera Botanica
127: 1-148.

Ekman, S. (2004). Bacidia. In: Nash III, T.H., Ryan, B.D.,
Diederich, P., Gries, C. & Bungartz, F. (eds), Lichen flora
of the Greater Sonoran Desert region. 2: 18-28. (Lichens
Unlimited: Tempe).

Elix, J.A. (2009). New crustose lichens (lichenized Ascomycota)
from Australia. Australasian Lichenology 64: 30-37.

Fryday, A.M., Printzen, C. & Ekman, S. (2014). Bryobilimbia, a
new generic name for Lecidea hypnorum and closely related
species. Lichenologist 46: 25-37.

Galloway, D.J. (1985). Flora of New Zealand Lichens. (P.D.

Hasselberg, Government Printer: Wellington).

Galloway, D.J. (2007). Flora of New Zealand Lichens. Revised
Second Edition. (Manaaki Whenua Press: Lincoln).

Hafellner, J. (1984). Studien in Richtung einer naturlicheren
Gliederung der Sammelfamilien Lecanoraceae und
Lecideaceae. Beiheft zur Nova Hedwigia 79: 241-371.

35

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Kalb, K., Lucking, R. & Serusiaux, E. (2000). Studies in Bacidia
sensu lato (lichenized Ascomycetes: Lecanorales). I. The
genus Bapalmuia. Mycotaxon 75: 281-309.

Kantvilas, G, (2008). Observations on the genus Scoliciosporum
in Australia, with the description of a second species of
Jarmania. Lichenologist A 0: 213-219.

Kantvilas, G. & Lucking, R. (2009). Fellhaneropsis pallidonigrans,
a south-eastern Australian lichen. Muelleria 27: 171-173.
Kantvilas, G. & Vezda, A. (1996). The lichen genus Sarrameana.

Nordic Journal of Botany 16: 325-333.

Llop, E. & Hladun, N. (2002). A key to the Iberian species of
the genus Bacidia with notes on some species. Bibliotheca
Lichenologica 82: 91-98.

McCarthy, RM. (2016). Checklist of the lichens of Australia and
its island territories. (Australian Biological Resources Study:
Canberra). Version 22 January 2016. http://www.anbg.gov.
au/abrs/lichenlist/introduction.html [accessed: 20 March

2017].

Obermayer, W. (2001). Arthrorhaphidaceae. Flora of Australia
58A: 33-36.

Orange, A., James, P.W. & White, F.J. (2001). Microchemical
Methods for the Identification of Lichens. (British Lichen
Society: London).

Stirton, J. (1875). Additions to the lichen flora of New Zealand.
Journal of the Linnean Society, Botany 14: 458-474.

Vezda, A. (1986). Neue Gattungen der Familie Lecideaceae
s.lat. (Lichenes). Folia Geobotanica et Phytotaxonomica 21:
199-219.

Vezda, A. (1990). Bacidina genus novum familiae Lecideaceae
s.lat. (Ascomycetes lichenisati). Folia Geobotanica et Phyto¬
taxonomica 25: 431-432.

With the exception of images and other material protected by a trademark and subject to review by the Government of
South Australia at all times, the content of this publications is licensed under the Creative Commons Attribution 4.0 Licence
(https://creativecommons.Org/licenses/by/4.0/). All other rights are reserved.

© 2017 Board of the Botanic Gardens and State Herbarium (Adelaide, South Australia)

36

w

Swainsona 31:37-43 (2017)

© 2017 Board of the Botanic Gardens & State Herbarium (Adelaide, South Australia) ©0

Goodenia asteriscus (Goodeniaceae), a new arid zone species from north¬
western South Australia and eastern Western Australia

PJ. Lang 0 & RJ.-P. Davies b

a State Herbarium of South Australia, GPO Box 1047, Adelaide, South Australia 5001
Email: peter.lang@sa.gov.au

b Natural Resources Adelaide and Mount Lofty Ranges, Department of Environment, Water and Natural
Resources, GPO Box 1047, Adelaide, South Australia 5001
Email: richard.davies@sa.gov.au

Abstract: Goodenia asteriscus PJ.Lang, a perennial, facultatively stoloniferous, rosette-forming herb is
described, illustrated and mapped, with distinguishing features and affinities also discussed. While this
species belongs to a group characterised by unequal wings on the adaxial corolla lobes, it does not
closely resemble any other species. An amendment to the Flora of Australia key is provided.

Keywords: biodiversity, new species, taxonomy, Australia, Goodenia subsect. Goodenia

Introduction

In May 2011, the second author collected a potentially
new Goodenia species while employed by Outback
Ecology Services to undertake an environmental
assessment survey of the proposed Southern Borefield
for the Metal-X Wingellina Nickel Project in far
eastern Western Australia (WA), c. 100 km SW of
Irrunytju (Wingellina) in the Ngaanyatjarra Aboriginal
Lands. Davies (2011) noted its affinity to Goodenia
heteromera F.Muell., a species not recorded from WA,
but recognised it as different. He sent the collection
{R.J. -P. Davies & L. Ransom LR868 ) to PERTH where
it was examined by M. Hislop, who could not place
it satisfactorily with any currently recognised species
of Goodenia in WA. Hislop (pers. comm., 2011)
confirmed the apparent affinity to G. heteromera , noting
a similarly-shaped indusium, but pointed out that there
were no signs of stolons on the specimen and that it
had significantly larger seeds with much wider wings
outside the range given for that species by Carolin
(1992, p. 235). He suggested that it be referred to as
Goodenia sp. aff. quasilihera (Davies & Alford 2012).

In November 2011 a further six collections were made
by RJPD along a proposed access track for the borefield
(Davies & Chapman 2011). These were referred to PJL
whose investigations of this and other material resulted
in the description and account presented here.

Matching specimens assembled by M. Hislop and
loaned from PERTH showed that the taxon had been
collected twice elsewhere in WA. The first known
collection was made in 1963 (A.S. George 4809) and the
second in 1973 ( D.E.Symon 8437). Neither had been
identified to species, but an AD duplicate of the latter

had been identified as Goodenia fascicularis F.Muell. &
Tate by R.C. Carolin in 1976.

A review of AD collections was made to see if it was
also present in South Australia (SA), focussing on
unidentified material and species of similar appearance
from the adjoining North-western Region (NW).
This uncovered four collections, all of which had been
misidentified as G. fascicularis.

In 2016, a short visit to the Anangu Pitjantjatjara
Yankunytjatjara Lands in the NW of SA by PJL enabled
additional field observations and collections for this
paper, including the type.

Goodenia asteriscus PJ.Lang, sp. nov.

Type: PJ.Lang 2928 & C.J.Brodie, 11 Oct. 2016,
12.1 km NW Kunytjanu homeland, 48.3 km SSE
from Pipalyatjara on road to Watarru, 26°35'11"S
129° 16'55"E, North-western Region, South Australia
(holo.: AD 278175; iso.: CANB, DNA, K, PERTH).

Goodenia sp. affin. quasilihera (L. Ransom 868)
R.J.Davies & J.Alford, Metals XLtd Wingellina Nickel
Proj. Level 1 FI. & Veg. Assessment Wingellina Borefield
Apr. 2012 (2012).

Goodenia fascicularis auct. non F.Muell. & Tate: Symon
in W.R. Barker et al., J. Adelaide Bot. Gard. Suppl. 1:
133 (2005), partly.

Perennial, facultatively stoloniferous, rosette-forming
herb 8-22 cm high, developing a woody taproot
and thickened basal stem retaining old pedicel bases;
vegetative parts strigose to sericeous, predominantly
clothed in simple hairs that are antrorse, lightly to
strongly appressed, laterally basifixed, straight to
gently and irregularly curved, moderately thick and
stiff, whitish (to pale gold) and translucent, sometimes

Published online: 24 Nov. 2017 • flora.sa.gov.au/swainsona

ISSN 2206-1649 (Print) • ISSN 2206-1657 (Online)

PJ. Lang & R.J.-P. Davies

Swainsona 31 (2017)

also with some obscure sparse minute short glandular
hairs, and spreading, long, thin, irregular, white,
septate, cottony hairs in and approaching axils and
occasionally extending sparsely to lower parts of leaves.
Leaves in dense basal rosettes, sometimes also terminal
on flowering racemes transitioning to form secondary
rosettes, procumbent to inclined adaxially, spathulate or
(narrowly-) obovate to oblanceolate-elliptic, (15—) 25-
65 x 5-18 mm (including petiole), all entire or earlier
leaves entire and later ones dentate to pinnatifid and
sometimes pinnatisect basally, with mid-dense antrorse
simple hairs and obscure sparse small glandular hairs
or the latter absent; lamina gradually tapering to an
indistinct petiole, 10-35 mm long; margins with 0-8
lobes, each 1-9 x 1-3.5 mm; apex acute; stolons 40-75
(-100) mm long or absent. Inflorescences radiating from
base of rosette, with 0-19 single-flowered ebracteolate
scapes (20-) 30-90 (-110) mm long, stiff when mature,
(usually) straight and persisting long after flowers or
fruit have fallen, and 0-21 erect to decumbent 2-5
(-8) -flowered, ebracteolate racemes (30-) 60-140 mm
long; bracts leaf-like, (narrowly) oblanceolate, 10-
25 x 1.5-5 mm, entire, with acute apex, all (excepting
sometimes the lowest) subtending pedicels; racemes

often contracted distally and grading to stolons bearing
terminal rosettes; scapes, raceme axes, and pedicels with
antrorse ± appressed simple hairs only, denser and more
spreading at articulation with ovary. Sepals (narrowly)
lanceolate to very narrowly elliptic 3.5—6 x 1—1.4 mm,
their basal V 3 adnate to lower V 2 — % of the ovary, outside
with mid-dense antrorse, appressed, strigose, simple
hairs and sparse short glandular hairs, inside glabrous.
Corolla predominantly yellow, deeper towards the throat
and paler on wings, grading to grey-violet with darker
anastomosing venation on lower adaxial lobes and
base of abaxial lobes to dark violet inside throat, 12.5-
18 mm long, all of outside with dense indumentum of
short glandular hairs, strigose hairs sparse, confined to
lobe apices, or absent; tube 1.6-3 mm long to dorsal
sinus of adaxial lobes, with pouch moderately developed
and extending 1-1.5 mm below sepal lobe attachment
but 0.5-0.8 mm short of ovary base; without enations,
erect short simple hairs on much of inside of tube and
throat and extending forwards along lateral margins of
a raised, thickened yellow zone below the median lobe;
auricles exceeding indusium, overlapping each other
and obscuring indusium in flower; corolla lobes often
decurved. Abaxial corolla lobes (3.5-) 4.5-8 (-9.3) x 1.5-
2.3 mm, fused for 5-8 mm beyond junction with adaxial
lobes, apex acute; wings along terminal 2.5-6.5 mm of
lobes, 3.5-7.5 (-9) mm x 1.3-3.2 mm, projecting 0.8-
2.5 mm beyond lobe apex, with entire margins. Adaxial

Fig. 1. Goodenia asteriscus, 92 km SW of Wingellina, WA, on 21 Nov. 2011 [RJ.Davies 464). A Old plant with a woody base,
persistent radiating scapes and entire leaves; B closer view of flowers.

Fig. 2. Goodenia asteriscus, c. 12 km NW Kunytjanu homeland, SA, on 11 Oct. 2016. A Decumbent plant with lobed leaves {PJ.Lang
2928&CJ.Brodie, Holotype); B young plant with prostrate inflorescences in early stage of transition to stolons {PJ.Lang 2926&CJ.Brodie).

38

Swainsona 31 (2017)

Goodenia asteriscus, a new arid zone species

corolla lobes 7-11.5 x 1-1.9 mm, fused to throat for a
further 1-3.5 mm to their divergence at dorsal sinus;
auricle yellowish-white, cuneate-cymbiform, 3.2-5.5
x 1.5—2.1 mm, with a fringe of hairs 0.2—5 mm long
along its junction with lobe; wings unequal, above
the auricle vestigial and 0.15-0.4 (-0.8) mm wide, on
opposite side along terminal 2.3-4.2 mm of lobe, 3-7
x 1-2.5 mm, with entire margins. Stamen filaments
linear, dorsiventrally flattened, 2.5-4 mm long, anthers
(narrowly) oblong 1.6-2.4 x 0.4-1 mm, base auriculate.
Ovary 3-4.8 x 1.9-2.8 mm, narrowly obovate, tapering
basally into a short stalk for 0.3-1.5 mm above
articulation with pedicel; septum incomplete, % length
oflocules, glabrous; ovules 38-48. Style straight, 3.3-4.5
x 0.8-1.0 mm, expanding to 1.8 mm at junction with
indusium, with spreading fine soft simple hairs 0.2-
0.3 mm long, denser on underside and forming a beard
at indusium junction. Indusium light brown, convex
above, dorsi-ventrally compressed, obverse-trapeziform,
slightly wider than long, 1.5—2.2 x 1.6—2.6 mm, glabrous
on back, apical margin straight on dorsal side, retuse on
ventral side, orifice narrow, bristles dense, white, flexed
upwards on dorsal lip, reflexed on ventral lip, grading
in length from 0.2 mm at margins to 0.7 mm at centre.
Fruit (widely-) ellipsoid, 7-10 x 5-7 mm. Seeds (3.5-)
4-6.2 x (3-) 3.5-5.2 mm, not mucilaginous; seed body
brown, flat to lightly convex, broadly lacriform-obovate,
seed coat shallowly reticulate; wing (0.7—) 1—1.9 mm
wide, pale-brown to stramineous, continuous with seed
body. Figs 1-6.

Affinities and distinctive features. Goodenia
asteriscus belongs in Goodenia sect. Goodenia subsect.
Ebracteolatae K.Krause, based on its yellow corolla,
prominent seed wing and the absence of bracteoles.
Within this subsection it fits Group 8 in the Flora of
Australia key of Carolin (1992) defined by “Flowers not
bracteolate; corolla wings unequal on [adaxial] 1 lobes”.
It is a distinctive species that does not closely resemble
any other.

Goodenia asteriscus is most like G. heteromera and
G. pascua Carolin in the very strong differentiation of
the adaxial (or posterior) corolla lobes, and with these
two species also shares a convex brownish indusium
with reflexed white bristles, articulate pedicels, and
reticulate seeds with prominent wings. Goodenia pascua
is readily distinguished by its well-developed long
racemes, the absence of pedicels arising directly from
the rosette, and its smaller flowers (6-10 mm long) with
the outside of the corolla covered in strigose hairs. The
corolla of G. pascua has a distinctive brown throat, and
the corolla of G. asteriscus most closely resembles that
of G. heteromera where the adaxial lobes differ from the
abaxial ones, not only by their markedly unequal wings
(very narrow or vestigial on the side above the auricle),
but also by having purple-violet coloration and dark
veining extend further up from the base of the lobes

1 in error as "abaxial" as noted by Sage (2000, p. 375)

2 This difference is sometimes obscured on pressed specimens as the
darker coloration becomes stronger and more extensive on drying.

towards the apex 2 (Figs 6 & 7). However, compared
to G. heteromera , G. asteriscus has larger flowers 12.5—
18 mm long (vs. 6-11 mm), with a well-defined corolla
pouch (vs. weakly or not developed); larger auricles
much exceeding the indusium (vs. similar in length
to indusium), but with much shorter fringing hairs;
and larger seeds 3.5-5.2 mm diam. (vs. 1.5-2.5 mm)
with a proportionately wider and more confluent wing
(vs. edge of seed body bevelled near wing junction).
Also, the indusium differs in detail, being broader
and dorsiventrally compressed with a thin orifice (cf.
more elongate and barrel-shaped with a gaping orifice
in G. heteromera). A distinctive feature is the marked
gradation in bristle length across the indusium orifice
(cf. gradation weak or not apparent in G. heteromera)
(Figs 6 & 7).

Fig. 3. Goodenia asteriscus, flowers. A PJ.Lang 2928 &
C.J.Brodie (Holotype); B PJ.Lang 2926 & CJ.Brodie.

Fig. 4. Goodenia asteriscus ( D.E.Symon 8437). A Mature fruit;
B dissepiment.

Fig. 5. Goodenia asteriscus, seeds [RJ.Davies 461).

39

PJ. Lang & R.J.-P. Davies

Swainsona 31 (2017)

Fig. 6. Goodenia asteriscus

{PJiang BS23-25475). A Pressed
flower, showing auricles much
exceeding the indusium, and
markedly unequal adaxial lobe
wings; B closer view of different
flower, showing indusium with
gradation in length of bristles
at orifice.

Fig. 7. Goodenia heteromera.

A Pressed flower, showing
adaxial corolla lobes with
unequal wings ( PJ.Lang
2275); B closer view, showing
indusium with orifice bristles of
± even length and the relatively
small auricles {DJ.Duval2298).

Fig. 8. Goodenia elongata.

A Pressed flower, showing
adaxial corolla lobes with equal
wings and elongate auricle
exceeding the indusium, which
has its left side partly folded
under ( M.Fagg 354); B closer
view, showing transversely
oblong indusium with orifice
bristles of ± even length and
dorsal hairs; glandular hairs on
outside of corolla visible on
lower left adaxial lobe ( R.Bates
6675).

40

Swainsona 31 (2017)

Goodenia asteriscus, a new arid zone species

Recent DNA sequencing (Shepherd et al., in prep.)
places G. asteriscus (represented by R.J.-P. Davies &
L.Ransom LR868) as sister to G. elongata Labill., before
linking to clades with G. heteromera and G. pascua.
In G. elongata the adaxial corolla lobes are relatively
undifferentiated with their wings either equal (in most
specimens observed) (Fig. 8A) or only a little narrower
above the auricles. This species also shares the broadly
similar indusium, articulate pedicels, and reticulate,
prominently winged seeds, but in addition matches
G. asteriscus by its glandular hairs on the outside of the
corolla (although varying from mid-dense to sparse)
(Fig. 8B), large elongate auricles, and large flowers.

Goodenia elongata is a species of damp habitats
confined to south-eastern Australia. In addition to the
equal to sub-equal adaxial corolla wings, it is readily
distinguished from G. asteriscus by its rhizomatous
and erect habit with well-developed cauline leaves and
weak rosette formation (vs. facultatively stoloniferous
rosettes), variable indumentum of stems and leaves
ranging from glabrous to mid-dense and sub-appressed
to strongly spreading (vs. mostly mid-dense and lightly
to strongly appressed), and a transversely oblong
indusium more than twice as wide as long (vs. slightly
wider than long) bearing simple hairs on its back (vs.
glabrous) and shorter orifice bristles of uniform length
(Fig. 8B) (vs. grading to longer in the centre).

Stolon production in G. asteriscus is much less strongly
developed than in G. heteromera , a trait which enables
that species to rapidly colonise wet clay soils in its flood
plain habitats subject to temporary inundation. Fully
differentiated stolons were seen in only a few collections
of G. asteriscus (e.g. D.E.Symon 8437 , P.J.Lang 2959
& C.J.Brodie), while some others had intermediate
states in transition from racemes. Furthermore,
individual plants varied widely in the proportion of
racemes to single-flowered scapes arising from the
main rosette. Counts of racemes vs. scapes ranged
from 10:1 and 17:3 at one extreme through to 1:5
and 0:25 at the other. Variation in the ratios of these
inflorescence types was independent of rosette size,
and together with occasional development of stolons
from racemes is indicative of different growth phases,
possibly determined by conditions in the immediate
environment.

The particularly large auricles (Fig. 6), also present in
G. elongata, are a notable feature of G. asteriscus. In all
fresh flowers of G. asteriscus observed, they overlapped
each other and completely obscured the indusium (Figs
IB & 3), similar to the effect noted and illustrated by
Holland (2015, p. 64, fig. 2) for G. ejfusa A.E.Holland.
Holland (2012, p. 690) found enclosing of the
indusium by the auricles in G. minutiflora F.Muell.
to be variable and suggested that this might be due to
flowering stage or an artefact of pressing. However, field
observations of G. asteriscus showed it to be consistent
during flowering. In G. heteromera the auricles are only
large enough to partly cover the indusium (Fig. 7B).
Yeo (1993, p. 150) suggested that sheltering of the

indusium by the auricles was important in controlling
access to pollen and favoured pollination by bees.

Although G. quasilibera Carolin is similar to G. asteriscus
in having glandular hairs on the outside of the corolla,
it differs significantly in other characters and analyses
of both cpDNA markers and the nrDNA Internal
Transcribed Spacer indicate that it is not closely related
(Shepherd et al., in prep). Other major differences
include non-reduced adaxial lobe wings, a prominent
corolla pouch or spur (often extending beyond the base
of the ovary), the indusium with a concave margin at
its orifice and glandular and simple hairs on its dorsal
surface, and distinctive black seeds with a narrow
overlapping wing.

Goodenia fascicularis, with which earlier collections were
sometimes confused, is readily distinguished by the
outside of its corolla lacking glandular hairs but covered
in strigose hairs, a broader and less convex indusium
with bristles uniform in length, and wings ± equal in
width on the adaxial corolla lobes.

The Goodenia key in Flora of Australia (Carolin 1992,
p. 164) should be amended for Group 8 by replacing
couplet 6 with the following:

6. Plant stoloniferous

6a. Corolla > 12 mm long, outside mainly covered by

short glandular hairs. G. asteriscus

6a: Corolla < 11 mm long, outside mainly covered by

simple strigose hairs. G. heteromera

6: Plant not stoloniferous

6b. Rosette-forming herb. G. asteriscus

6b: Herb or subshrub with erect leafy stems. 7

Distribution. The known distribution of G. asteriscus
spans 220 km from west of Cavenagh Range in WA
to Mt Lindsay (Watarru) in the NW Region of SA,
in areas that are not dominated by dense dunefields
(Fig. 9). It is also reported for the Northern Territory,
just across the border from SA, where P. Latz observed
sterile plants matching those of P.Latz24770.

Ecology. The species grows in hard clay and loam soils
and is able to colonise compacted gravelly surfaces of
road margins (. P.J.Lang 2926 & C.J.Brodie) and an
airway scrape’ {D.E.Symon 847). It has been mostly
found on limestone plains with outcropping calcrete
or nodular limestone and associated plant communities
dominated by Triodia scariosa hummock grassland,
often with emergent Eucalyptus socialis subsp. eucentrica
open mallee or Acacia kempeana open shrubland. The
associated suite of plants recorded in the Wingellina
Nickel project area (Davies 2011, Davies & Alford
2012 ), and those observed near Kunytjanu, closely
fit floristic Group 22 ‘Spinifex Hummock Grassland
on Limestone Plains’ of Lang et al. (2003, p. 171),
an extensive community in the Anangu Pitjantjatjara
Yankunytjatjara Lands of South Australia.

41

PJ. Lang & R.J.-P. Davies

Swainsona 31 (2017)

A collection from NE slopes of a rocky hill known as
‘Scarface’ in the Tomkinson Ranges {J.Z.Weber 5397),
is an exception to this habitat. It was confirmed by the
discovery of a single plant (. P.J.Lang 2959) on a rocky
slope of the same range system composed of dark red-
brown ultra-mafic rock.

Flowering. Mainly Sep.-Jan.

Conservation status. Goodenia asteriscus has been
infrequently collected and the populations observed
in the Wingellina borefield area (RJPD) and NW
of Kunytjanu (PJL) amount to only a few hundred
individuals in total. However, the widespread
occurrence of its major habitat type, in terrain with
limited road and track access, means that many more
populations may exist. Until targeted surveys are
undertaken, and in the absence of any discernable
threatening processes, it is best assigned to the ‘Data
Deficient’ category (IUCN 2017).

Etymology. The epithet, a Latin noun in apposition, is
derived from the Greek asteriskos meaning ‘little star’. It
alludes generally to the radiating pattern of peduncles
and secondary rosettes, but more particularly to the
disposition of persistent flower scapes reminiscent of an
asterisk symbol. A number of Goodenia species retain
their scapes long after flowers or fruits have abscissed,
but this is a pronounced feature in G. asteriscus. Similar
long, stiff, straight persistent radiating scapes are also
seen in G. pedicellata L.W.Sage & K.W.Dixon (Sage
& Dixon 2005, p. 515, fig. 1) and G. gibbosa Carolin
(e.g. E.A. Shaw 450, 451) although these are not closely
related.

Goodenia asteriscus other specimens examined

WESTERN AUSTRALIA: RJ.Davies 460 {-462,
-464, -465, -466), 93 (-91, -92) km direct SW of
Wingellina, c. 9 (-10.5) km direct W of Tjuntjuntjarra
Track, on proposed access track route, (20-) 21
Nov. 2011 (AD); R.J.-P.Davies & L.Ransom LR868,
Great Victoria Desert, c. 100 km SW of Wingellina
township, towards southern end of proposed Southern
Borefield for Metal-X Wingellina Nickel Project, 6 May
2011 (PERTH); A.S. George 4809, N end of Cavanagh
[Cavenagh] Range, 8 Jul. 1963 (PERTH); M.Henson
& M.Hannart 32638, 28 km SSE of Warburton
[coordinates plot as: 113 km W of Warburton & 28 km
S of Jameson Community], 29 Nov. 2011 (PERTH);
D.E.Symon 8437, Blackstone Range, Wingelina
[Wingellina], 8 Jan. 1973 (AD, PERTH; NSW n.v.).

SOUTH AUSTRALIA: North-Western: N.N.Donner
6458, up to c. 1 km SW of Krewinkel Hill, 2 Sep. 1978
(AD; G n.v.)-, P.J.Lang BS23-25475, 12.2 km NW of
Kunytjanu Homeland on road to Pipalyatjara, 3.0 km
direct WNW of Krewinkel Hill, 5 Sep. 1995 (AD,
PERTH); P.J.Lang 2926, 2927, 2930 & C.J.Brodie,
(12.1—) 12.2 km NW Kunytjanu homeland, 48.2
(-48.3) km direct SSE from Pipalyatjara, 12 Oct. 2016
(AD); P.J.Lang 2959 & C.J.Brodie, 0.85 km SW Kalka

community, on mid-slopes of Dulgania Hill range,
12 Oct. 2016 (AD); P.Latz 24770, 5 km NNW of
Aparatjara, on track to Lake Wilson, 53 km E of Kalka,
3 Sep. 2009 (NT n.v., image seen, AD n.v., PERTH
n.v.)-, D.E.Murfet BS23-27534, 14.5 km from Mt
Lindsay NE towards Kintore Ave turn-off, 10.8 km
direct NNE Wartaru [Watarru], 5 Sep. 1995 (AD);
J.Z.Weber 5397, Tomkinson Ranges, Dulgunia Hills,
NE slopes of Scarface, c. 9 km direct NW of Mt Davies
summit, 2 Sep. 1978 (AD; COLO n.v.).

Goodenia quasilibera type specimens examined

WESTERN AUSTRALIA: A. S. George 2171, Thomas
River valley, N of Homestead, 10 Dec. 1960 (Holotype,
PERTH).

Acknowledgments

Support from Sara Weir and Anangu trainee rangers
at Pipalyatjara in facilitating a revisit by PJL of the
population NW of Kunytjanu homeland, and from
Chris Brodie for field assistance on that trip, was much
appreciated. We are grateful for the initial investigation
of WA collections by Mike Hislop, the loan of
specimens from PERTH herbarium, and refinements
to this paper suggested by Kelly Shepherd and Dave
Albrecht.

References

Carolin, R.C. (1992). Goodenia. Flora of Australia 35: 147-281
(235).

Davies, R.J.-P. (2011). Metals X Limited Wingellina Nickel
Project: Level 1 Flora and Vegetation Assessment of the
Wingellina Borefield July 2011 — Draft report. (Outback
Ecology Services: Jolimont, WA). https://consultation.epa.
wa.gov.au/seven-day-comment-on-referrals/wingellina-
nickel-project/supporting_documents/Flora%20and%20
Vegetation%20Borefield%20Jul%20201 l.pdf [accessed: 15
Aug. 2017].

Davies, R.J.-P. & Alford, J. (2012). Metals XLimited Wingellina
Nickel Project: Level 1 Flora and Vegetation Assessment of
the Wingellina Borefield April 2012. (Outback Ecology
Services: Jolimont, WA) https://consultation.epa.wa.gov.au/
seven-day-comment-on-referrals/wingellina-nickel-project/
supporting_documents/Flora%20and%20Vegetation%20
Borefield%20Apr%202012.pdf [accessed: 15 Aug. 2017].

Davies, R.J.-P. & Chapman, A. (2011). Metals X Limited
Wingellina Nickel Project: Proposed Borefield Drill Line
Targeted Flora Assessment November 2011. (Outback Ecology
Services: Jolimont, WA). https://consultation.epa.wa.gov.au/
seven-day-comment-on-referrals/wingellina-nickel-project/
supporting_documents/Borefield%20Drill%20Line%20
Targeted%20Flora.pdf [accessed: 15 Aug. 2017].

Holland, A.E. (2012). Nomenclature and synonymy of several
Goodenia R.Br. (Goodeniaceae) species from northern
Australia. Austro bailey a 8(4): 688-695.

Holland, A.E. (2015). Goodenia ejfusa (Goodeniaceae), a new
species from north Queensland, Australia. Telopea 18:
61—66.

42

Swainsona 31 (2017)

Goodenia asteriscus, a new arid zone species

Blackstone Range

O

, Cavenagh Range

% J

Vin

# Goodenia asteriscus

.state borders

- road

sand dunes

W.A.

25

50

Wingellina

(Irrunytju)

100
■ km

N.T.

m

S.A.

Kunytjanu

Q-

A

n

o

, Mt Lindsay
(Watarru)

Fig. 9. Goodenia asteriscus, known distribution.

IUCN Standards and Petitions Subcommittee [IUCN] (2017).
Guidelines for using the IUCN Red List Categories and
Criteria. Version 13. (International Union for Conservation
of Nature and Natural Resources: Cambridge). http://www.
iucnredlist.org/documents/RedListGuidelines.pdf [accessed:
15 Aug. 2017].

Lang, P.J., Canty, P.D., Nesbitt, B.J., Baker, L.M. & Robinson,
A.C. (2003). Vegetation. In: Robinson, A.C., Copley, P.B.,
Canty, P.D., Baker, L.M. & Nesbit, B.J. (eds), A Biological
Survey of the Anangu Pitjantjatjara Lands South Australia
1991-2001. (Dept, for Environment & Heritage, South
Australia: Adelaide).

With the exception of images and other material protected by a trademark and subject to review by the Government of
South Australia at all times, the content of this publications is licensed under the Creative Commons Attribution 4.0 Licence
(https://creativecommons.Org/licenses/by/4.0/). All other rights are reserved.

© 2017 Board of the Botanic Gardens and State Herbarium (Adelaide, South Australia)

Sage, L.W. (2000). New taxa in Goodenia (Goodeniaceae) from
the south-west of Western Australia, with an update to the
Goodenia key in the Flora of Australia. Nuytsia 123(2):
367-377.

Sage, L.W. & Dixon, K.W. (2005). Goodenia pedicellata
(Goodeniaceae), a new species from the Pilbara bioregion of
Western Australia. Nuytsia 15(3): 513-516.

Yeo, P.F. (1993). Secondary Pollen Presentation: Form, Function
and Evolution. Plant Systematics and Evolution Supplements-.
I-VII, 1-268.

43

w

Swainsona 31:45-48 (2018)

© 2018 Board of the Botanic Gardens & State Herbarium (Adelaide, South Australia) ©0

Pertusaria crassilabra Mull. Arg. - a reinstated name for an Australasian
lichen

Gintaras Kontvilas

Tasmanian Herbarium,Tasmanian Museum and Art Gallery, PO Box 5058, UTAS LPO, Sandy Bay, Tasmania 7005
Email: Gintaras.Kantvilas@tmag.tas.gov.au

Abstract: Pertusaria crassilabra Mull. Arg., based on a Victorian type specimen and hitherto considered
a synonym of P. melanospora Nyl., is reinstated and recorded for south-eastern Australia (including
Tasmania) and New Zealand. The species is described and illustrated. Pertusaria amaurospora Hellbom
and P. melanospora var. sorediata Elix & A.W.Archer are synonyms of P. crassilabra. The occurrence in
Australasia of P. melanospora sens, str., based on a type from South America, is considered doubtful.

Keywords: Australia, biodiversity, lichenised fungi, New Zealand, Pertusariaceae

Introduction

Pertusaria DC. is one of the largest genera of lichenised
fungi and, with 191 formally recorded taxa, certainly
one of the largest in Australia (McCarthy 2017). The
genus has been extensively studied in the region, in
particular by the Australian lichenologist, Alan Archer,
who, alone or in collaboration, compiled several,
continent-wide accounts (Archer 1997, 2004; Archer
& Elix 2016). Regional studies for Tasmania (Kantvilas
1990, Kantvilas & Elix 2008) and New Zealand
(Galloway 1985, 2007) have also been published.

Although most species of Pertusaria have colourless
ascospores, a small number are known to develop brown
or grey-brown spores at maturity or at senescence. One
such species is the aptly named P. melanospora Nyl. This
saxicolous species is based on a type from Chile (Nylander
1855) but has been more widely recorded, including
for Australia (Archer 1997) and New Zealand (Archer
& Elix 1993) and Southern Africa (Stizenberger 1890).
Along the way, several Australasian Pertusaria names
have been consigned to synonymy with P. melanospora ;
viz. P. crassilabra Mull .Arg., P. amaurospora Hellbom and
P. limescens Zahlbr. More recently, Elix & Archer (2013)
described a sorediate variety of P. melanospora , based on
material from Kangaroo Island, South Australia, but
also recorded from elsewhere in south-eastern Australia
and New Zealand.

The present study is based on a review of Australasian
herbarium material filed as P. melanospora , field work
in Tasmania, Kangaroo Island and New Zealand’s
North Island, study of specimens of P. melanospora
sens. str. from South America, and an examination of
relevant type material. As a result, a long-disused name,
P. crassilabra is reinstated for the Australasian taxon,
and the occurrence of P. melanospora in Australasia is
considered doubtful.

Material and methods

Anatomical and morphological observations were
undertaken using light microscopy, with thin hand-cut
sections mounted in water, 10% KOH, lactophenol
cotton blue and ammoniacal erythrosin. Ascospore
measurements are presented in the format: 5 th
percentile-^z'mzgz , -95 th percentile, with outlying values
given in brackets; these are based on 45 observations
for each taxon. Routine chemical analyses by thin-
layer chromatography follow standard methods in
lichenology (Orange et al. 2001).

Taxonomy

Pertusaria crassilabra Mull.Arg.

Hedwigia 32: 126 (1893). - Type: [Australia] Victoria:
Loutit Bay [near Lome], 1891, Luehmann s.n. (holo:
G!).

Pertusaria amaurospora Hellbom, Bih. Kongl. Svenska
Vetensk.-Akad. Handl. 21(3) 13: 73 (1896). - Type:
New Zealand, South Island, Maunganui, near
Taurunga, Dec. 1874, S. Berggren [lecto.: S! (L5124),
fide Galloway 1985; isolecto.: S! (L986, L5125,
L5126)].

Pertusaria melanospora var. sorediata Elix &
A.W.Archer, Australas. Lichenol. 73: 8 (2013). - Type:
Australia, South Australia, Kangaroo Island, Cape St
Albans, 35°48’S, 138°07’E, 20 m alt., on laterite in
pasture, 24 Sep. 2010, G. Kantvilas 176/10 (holo.:
HO!; iso.: AD!).

Pertusaria melanospora auct. non Nyl.: A.W.Archer,
Biblioth. Lichenologica 69: 104 (1997), FI. Austral.
56A: 146 (2004); A.W.Archer & Elix, New Zealand
J. Bot. 31: 111 (1993), Austral. Pertusaria 85 (2016);
D.J.Galloway, FI. New Zealand Lich. (ed. 2) 1150
(2007).

Published online: 9 Feb. 2018 • flora.sa.gov.au/swainsona

ISSN 2206-1649 (Print) • ISSN 2206-1657 (Online)

G. Kantvilas

Swainsona 31 (2018)

Thallus crustose, very tightly adnate, rimose-areolate,
dull yellowish or brownish yellow, up to 10 cm wide
or more, sorediate; individual areoles 0.1-0.5 mm
wide, to 200 (-500) pm thick, plane or slightly convex,
contiguous or dispersed over an effuse, patchy, pale
greyish prothallus, with an upper cortex 20-30 pm
thick, composed of short-celled hyphae 4-5 pm wide;
soredia whitish or greenish white, coarse, sometimes
becoming elongate, corticate and “pseudoisidiate” at
the most mature stage, arising in irregular soralia 0.3-
0.5 mm wide, formed by a disintegration of the thallus
upper cortex, typically sparse and inconspicuous,
but sometimes spreading across the entire thallus;
photobiont a unicellular green alga with individual
cells globose, 6-14 pm diam. Apothecia verruciform,
smooth or rather wrinkled, apically flattened, when
well developed 0.4-1.3 mm wide, constricted at the
base and appearing lecanorine; ostioles mostly 1-4
per verruca, at length gaping, to c. 0.4 mm wide and
appearing disc-like. Epithecium brownish grey, weakly
K+ violet, C+ violet. Ascospores 4-8 per ascus, at first
hyaline, later bluish grey to brownish, K+ violet, C+
violet, ellipsoid to ovate, 48 —58.4—7A (—76) x 2 A—31.8—
40 (-42) pm; wall mostly 10-15 pm thick. Pycnidia
immersed, flask-like, 60-100 pm wide, visible as black
specks in the surface of the thallus; conidia filiform,
mostly ± straight, 16-22 x 0.8-1 pm. Figs 1-2.

Chemistry. Arthothelin; thallus K-, KC+ orange,
C+ orange, P-, UV+ dull orange. Elix & Archer
(2013) also record 4,5-dichloronorlichexanthone,
2 ,5-dichloronorlichexanthone, 2,4-dichloronorliche-
xanthone (±), thiophanic acid (±) and atranorin (±) in
trace concentrations.

Remarks. Pertusaria crassilabra is very distinctive and
has no known confusing species within its distributional
range. Perhaps the most similar species, on account

of its verruciform apothecia and sometimes yellowish
thallus, is P. lophocarpa Korb., but this common taxon
differs by having an esorediate thallus containing
2 ’-0-methylperlatolic acid as the major secondary
substance, has larger apothecia (to 1-3 mm wide)
and colourless ascospores (unpubl. observations and
Archer 2004). Fertile material of P. crassilabra is rather
uncommon (for example, none has been collected in
Tasmania) but the yellowish, areolate, C+ orange thallus
with scattered soralia is unmistakable. The degree of
development of soredia is highly variable. In some
specimens, including the type specimens, they are sparse
and associated mainly with abraded parts of the thallus.
Such a specimen is illustrated by Archer & Elix (2016,
Fig. 75) as P. melanospora var. melanospora. In others, the
soredia spread across the entire upper surface and may
become elongate, corticate and essentially pseudoisidiate
(for example, see Fig. 2). There does not appear to be
any obvious correlation between development of soredia
and habitat factors and all specimens seen display at least
some evidence of soredia. The K+, C+ violet pigment
has not attracted comment from previous Australasian
workers, although it is noted by Schmitt et al. (2006)
in their study of some related North American taxa. It
is consistently present and displays the same reactions
as sedifolia-gcey of Meyer & Printzen (2000), a pigment
that is scattered amongst lichens, occurring, for example,
in species of Micarea Fr. and Rimularia Nyl., and in
Pertusaria erubescens (Taylor) Nyl.

Although clearly related to P. melanospora sens. str. and
sharing ± identical ascospores and thallus chemistry,
P. crassilabra consistently has a distinctly thinner, chinky-
areolate, sorediate thallus, morphological characters
that justify recognising these taxa at species rank. In
the course of this study, I have examined a wide range
of Australasian specimens labelled as P. melanospora in
various herbaria (as listed below) and found all to be

Fig. 1. Pertusaria crassilabra

habit (Kantvilas 176/10),
showing the rimose thallus,
scattered soralia and basally
constricted, verruciform apo¬
thecia. Scale = 2 mm.

46

Swainsona 31 (2018)

Pertusaria crassilabra - a reinstated name for an Australasian lichen

Fig. 2. Pertusaria crassilabra detail (Kantvilas 176/10). Note
the"/.ecanora-like"apothecia with multiple, black ostioles, and
the coarse, pseudoisidiate soredia (top left). Scale = 1 mm.

sorediate, and therefore matching P crassilabra. The
presence of P. melanospora in Australasia is thus highly
doubtful, given that all specimens cited by Galloway
(1985) and Archer & Elix (1993) are sorediate.

The type specimen of P. amaurospora is clearly sorediate
and conspecific with P crassilabra. Galloway (1985)
regarded Pertusaria limescens Zahlbr., based on a type
collection from New Zealand’s South Island, as a
synonym of P. amaurospora , albeit with some doubt as
he did not examine the type collection. Subsequently,
both of these taxa were listed as synonyms of
P. melanospora by Archer & Elix (1993). I am reluctant
to accept P. limescens as a synonym until authentic
material can be located and studied. The original
description of Zahlbruckner (1941) makes no mention
of certain critical characters, such as, for example, the
distinctly yellowish thallus or the pigmented ascospores;
instead the thallus is described as K+ reddish, C-, and
the spores as uncoloured.

Distribution and ecology. Pertusaria crassilabra is
known from Kangaroo Island and mainland South
Australia, from Victoria, Tasmania, New South Wales,
the Australian Capital Territory, and the North and
South Islands of New Zealand. It has a very wide
distribution on exposed rocks, but chiefly in drier areas
and especially at the coast. Elix & Archer (2013) list
a suite of lichens with which P. crassilabra (under its
synonym, P. melanospora var. sorediata) may be associated
with and which indicate its broad ecological amplitude.

Selected specimens examined

TASMANIA. Furneaux Islands, East Kangaroo Island,
40°10’S 147°54’E, 3 m alt., 20.iv.1969, J.S. Whinray s.n.
(MEL); Cockle Bay Lagoon, 42°42’S 147°56’E, 1 m alt.,
24.vii.2011, G. Kantvilas 218/11 (HO); Spiky Bridge,
42°11’S 148°04’E, 60 m alt., ll.viii.2011, G. Kantvilas
225/11 (CANB, HO); Gunners Quoin, 42°46’S 147°19’E,
440 m alt., l.iv.2013, G. Kantvilas23/13 (HO); Lucas Point,

Fishermans Haul, 43°02’S 147°20’E, 1 m alt., 8.xii.2013,
G. Kantvilas 407/13 (HO); Cape Surville, 42°57’S 148°00’E,
25.ix.20l4, G. Kantvilas 476/14 (HO); Dorman Point,
42°54’S 147°44’E, 5 m alt., 13.xii.20l4, G. Kantvilas497/14
(HO); mouth of Interview River, 4l°35’S 144°53’E, 3 m
alt., 314.2015, G. Kantvilas 140/15 (HO); The Nut, 40°46’S
145°18’E, 2 m alt., 25.X.2016, G. Kantvilas 399/16 (HO).
SOUTH AUSTRALIA. Kangaroo Island: Windmill Bay,
35°51’S 138°07’E, 20 m alt., 17.ix.2012, G. Kantvilas 469/12
(AD, HO); c. 3.5 km NE of Stokes Bay, 35°37’S 137°13’E,
50 m alt., 19.ix.2012, G. Kantvilas 536/12 (AD, HO);
Western River Cove, W end of beach, 35°40’S 136°58’E, 2 m
alt., 27.ix.2015, G. Kantvilas 413/15 &B. de Villiers (HO).
VICTORIA. Point Wilson, 15 km NE of Geelong, 38°04’S
144°30’E, iii.1980, A. Geddes s.n. (MEL); Lake Condah,
38°04’05”S 141°49’53”E, 28.iii.2011, V Stajsic 5724
(MEL, NSW); Phillip Island, Kitty Miller Bay, 38°30’34”S
145°10’20”E, 2.iv.2013, V Stajsic 7379 &J.G. Eichler (HO,
MEL).

NEW ZEALAND, NORTH ISLAND. Waitakere Ranges,
Mercer Bay Track, summit of Farley Point, 36°59’S 174°28’E,
10.xi.2016, G. Kantvilas s.n. (AK, HO); Union Bay, N end
of Karekare Beach, 36°59’12”S 174°28’24”E, 1.5 m alt.,
10.xi.2016, G. Kantvilas s.n. (AK, HO).

Further specimens from South Australia, New South Wales,
the Australian Capital Territory and South Island, New
Zealand, held in CANB, are cited by Elix & Archer (2013)
and are not repeated here.

Pertusaria melanospora Nyl.

Ann. Sci. Nat., Bot., ser. 4, 3: 159 (1855). - Type:
Chile, Quilmenco, C. Gay s.n. (lecto.: H-NYL 23603!,
fide Archer & Elix 1993; isolecto: H-NYL 23604!).

A description of this species is provided by Messuti
(2005). It is characterised by a relatively thick,
continuous, esorediate, yellowish thallus containing
arthothelin as the major substance, verruciform
apothecia, (4-) 6-8-spored asci and bluish grey to
brownish ascospores. Messuti (2005) gives the ascospore
dimensions as 55-81 x 30-39 pm; in the present study,
these were measured as (36-) 44-57-74 x (24-) 25-
31-36 pm (n = 43). Both the ascospores and the greyish
epithecium react a faint K+ violet, C+ violet.

Messuti & Archer (2003) and Schmitt et al. (2006) list
several synonyms of P. melanospora {viz. P. calcarea (Nyl.)
Nyl., Melanaria atacamae C.W. Dodge and P. ochracea
Kremp.), based on South American types, but these were
not examined during the present study; the published
descriptions of the taxa make no mention of soredia.
The status of the African report of this South American
taxon (Stizenberger 1890) remains unresolved.

Specimens examined

CHILE, JUAN FERNANDEZ ISLANDS. Isla Robinson
Crusoe (Mas a Tierra), Gipfel, SW of Tres Puntas,
“Vogelgipfel”, alt. 380 m. s. m., 54.1917, C. & I. Skottsberg
(S 55391); Santa Clara, northeast coast opposite El Morro
del Spartan, 15.xii. 1965, H.A. Imshaug38182 (S).

47

G. Kantvilas

Swainsona 31 (2018)

CHILE. Prov. Coquimbo, Kiistenkordille, Serra de Frai
Jorge, 17.viii.1917, C. & I. Skottsberg (S 55390); Valparaiso,
Alto del Puerto, I4.viii. 1940, R. Santesson 2997 (S); Prov.
Aconcagua, Punto Los Molles, 50 m alt., 13.xi.1976, W.A.
Weber & B. Johnston ( Lich. Exsicc. Colo. 545) (MEL, S).

Acknowledgements

The greater part of this work was undertaken at the
Swedish Museum of Natural History (S) and I thank
the Director and staff, especially Mats Wedin, for their
hospitality during my visit. The photographs were
prepared by Jean Jarman. I also thank the curators of
the Herbaria CANB, G and MEL for their hospitality
and/or loans of specimens. Some material studied
was collected during field surveys co-funded by the
Australian Biological Resources Study (ABRS) and
BHP Billiton under the Bush Blitz Program. Laboratory
work was supported by an ABRS Tactical Taxonomy
Grant.

References

Archer, A.W. (1997). The lichen genus Pertusaria in Australia.

Bibliotheca Lichenologica 69: 1-249.

Archer, A.W. (2004). Pertusariaceae. Flora of Australia 56A:
116-172.

Archer, A.W. & Elix, J.A. (1993). Saxicolous species of Pertusaria
(Lichenes) common to New Zealand and Australia. New
Zealand Journal of Botany 31: 111-116.

Archer, A.W. & Elix, J.A. (2016). Australian Pertusaria.

(Privately published by the authors).

Elix, J.A. & Archer, A.W. (2013). A new sorediate variety
of Pertusaria melanospora (lichenized Ascomycota,
Pertusariaceae). Australasian Lichenology 73: 8-9.

Galloway, D.J. (1985). Flora of New Zealand Lichens. (P.D.
Hasselberg, Government Printer: Wellington).

Galloway, D.J. (2007). Flora of New Zealand Lichens. Revised
second edition. (Manaaki Whenua Press: Lincoln).

Kantvilas, G. (1990). The genus Pertusaria in Tasmanian
rainforests. Lichenologist 22: 289-300.

Kantvilas, G. & Elix, J.A. (2008). Additions to the lichen genus
Pertusaria in Tasmania. Sauteria 15: 249-263.

McCarthy, P.M. (2017). Checklist of the lichens of Australia
and its island territories. (Australian Biological Resources
Study: Canberra), http://www.anbg.gov.au/abrs/lichenlist/
introduction.html [Version 12 April 2017].

Messuti, M.I. (2005). The genus Pertusaria (Pertusariales:
Pertusariaceae) in the Juan Fernandez Archipelago (Chile).
Lichenologist 37: 111-122.

Messuti, M.I. & Archer, A.W. (2003). Two new synonyms of
Pertusaria melanospora. Lichenologist 35: 409-410.

Meyer, B. & Printzen, C. (2000). Proposal for a standardized
nomenclature and characterization of insoluble lichen
pigments. Lichenologist 32: 571-583.

Nylander, W. (1855). Additamentum in floram cryptogamicam
chilensem. Annales des sciences naturelles, Botanique, Series 4,
3: 145-187.

Orange, A., James, P.W. & White, F.J. (2001). Microchemical
Methods for the Identification of Lichens. (British Lichen
Society: London).

Schmitt, I., Lumbsch. H.T. & Bratt, C. (2006). Two new
brown-spored species of Pertusaria from south-western
North America. Lichenologist 38: 411-416.

Stizenberger, E. (1890). Lichenaea africana. Berichte iiber
die Thdtigkeit der St. Gallischen Naturwissenschaftlichen
Gesellschaft 1888-1889: 105-149.

Zahlbruckner, A. (1941). Lichenes Novae Zelandiae a cl.
H.H. Allan eiusque collaboratoribus lecti. Denkschriften
der Akademie der Wissenschoften in Wien, Mathematisch-
Naturwissenschafliche Klasse 104: 249-380.

With the exception of images and other material protected by a trademark and subject to review by the Government of
South Australia at all times, the content of this publications is licensed under the Creative Commons Attribution 4.0 Licence
(https://creativecommons.org/licenses/by/4-0/). All other rights are reserved.

© 2018 Board of the Botanic Gardens and State Herbarium (Adelaide, South Australia)

48

w

Swainsona 31:49-53 (2018)

© 2018 Board of the Botanic Gardens & State Herbarium (Adelaide, South Australia) ©0

The type of Sturt pea found

Alex S. George

School of Veterinary and Life Sciences, Murdoch University, 90 South Street, Murdoch, Western Australia 6150
Email: a.george@murdoch.edu.au

Abstract: The type collection of Sturt pea was thought to have been lost soon after the publication of
its original name, Donia formosa G.Don. For a long time, it was assumed (wrongly) to be at the Natural
History Museum (BM), but it has now been located in the herbarium in Geneva (G). Annotations on the
sheet show that the collection details have also been mistaken: instead of the'Curlew River', collected
by Phillip Parker King or Frederick Bedwell, it was collected by Allan Cunningham on the Malus Islands
in the Dampier Archipelago.

Keywords: Sturt pea, Donia formosa, Clianthus formosus, Swainsona formosa, Willdampia formosa,
Fabaceae, Lambert Herbarium, typification

Introduction

Sturt pea is a spectacular flower that has evoked
admiration since its discovery by William Dampier in
1699. Explorers and collectors who came across it in
the 19 th century tended to exclaim over it (Symon &
Jusaitis 2007; George 1999b). It was described in pre-
Linnaean literature, and first named in the Linnaean
system in 1832. Since then it has been treated in many
works such as floras, horticultural works and field
guides, and it has attracted the interest of many artists.
On 23 November 1961 it was proclaimed as the floral
emblem of South Australia ( Government Gazette of
South Australia no. 51: 1731).

One matter in Sturt peas nomenclatural history has
received little attention and can only now be resolved—
the type collection, namely the locality, the collector,
and the present whereabouts of the type specimen itself.
George Don (1832) published two names for what we
regard as Sturt pea. His protologues are:

D. SPECIOSA ; herbaceous, very villous ; leaflets
opposite, lanceolate, acute ; flowers umbellate ; calyx
5-cleft; legume silky. G. Native of New Holland, at
Regent’s Lake. Flowers large, crimson.

D. FORMOSA ; plant herbaceous, very villous ; leaflets
opposite, obovate ; flowers racemose ; calyx 5-cleft ;
legume silky. A. G. Native of the North-west coast of
New Holland, at the Curlew river. Capt. King.

The symbol ^ refers to the herbaceous habit and
G to the requirement to grow it in a glasshouse. The
differences in leaflet shape and inflorescence were
considered insignificant by Brown (1849), who
recognised just one species for which he accepted the
name Clianthus dampieri A.Cunn. This specific epithet

was accepted by Bentham (1864) and all other writers
until Ford and Vickery (1950).

The person cited as collector of the type of Donia
formosa , ‘Capt. King.’, was Phillip Parker King (1791—
1856), captain of HMC Mermaid (and later HM Survey
Ship Bathurst) on several surveying voyages around the
Australian coast from 1817 to 1822. He sometimes
assisted the plant collector Allan Cunningham, as well
as collecting on his own account. King gave the name
‘Curlew River’ to a tidal creek just east of the mouth
of a river later called the Ashburton, Western Australia
(c. 21 °4l’S, 114°58’E), but it never became an official
name. There is a Curlew Bank just offshore, named by
J.W. Combe around 1899/1900, apparently alluding
to King’s name (Murray & Hercock 2008). Later, the
collector of the type was thought to be the master’s mate,
Frederick Bedwell, who is recorded as bringing aboard
a specimen of ‘Dolichos’ from an excursion ashore at
the Curlew River on 19-21 February 1818 (Curry et
al. 2002). It has been thought that this specimen was
Sturt pea, e.g. Symon & Jusaitis (2007, p. 18), but note
that the quotation given there refers to Cunningham’s
landing on the Malus Islands on 1 March (see below),
not the Curlew River.

Cunningham’s journal (A.E.Orchard, pers. comm.) tells
a different story. His entry for 20 February 1818 reads:

Our first officer had landed on the Main and had
visited the Saltmarsh at the Back of the Beach, &
reports the quantities of crystallized Salt he saw on
these flats. He brought me a specimen of a Dolichos
with acute stalks, which he had gather’d on the
Sands (D. foliolis rhombeo-hastatis, reticulato-ramis,
pedunculis petiolo duplo longioribus, racemis parvis,
leguminibus glabris, solitariis, stipulis ovatis, acutis,
a small annual plant).

Published online: 11 April 2018 • flora.sa.gov.au/swainsona

ISSN 2206-1649 (Print) • ISSN 2206-1657 (Online)

A.S. George

Swainsona 31 (2018)

The officer was Bedwell, but features of the plant rule
out this being Sturt pea, especially its glabrous pods.

Further, Cunningham’s entry for 1 March, when he
landed on the Malus Islands (21°31’S, 116°41’E), reads:

I was not a little surprised to find Kennedia speciosa,
a plant discover’d in July 1817 in sterile bleak open
flats near the Regent Lake on the Lachlan River in
Lat. 33°13’S and Long. 146°40’E. It is not common,

I could only see 3 plants, of whom one was in flower.

... This island is the Isle Malus of the French.

Robert Brown (1849, p. 72) repeated this account
almost verbatim but, apart from Bentham (1864), no
later worker has noted the correct locality.

‘Kennedia speciosa was Cunningham’s manuscript
name for Sturt pea. His collection from Regent’s Lake
(now Lake Cargelligo) New South Wales in July 1817,
while on an expedition with John Oxley, became the
type of Donia speciosa G.Don.

The type of Donia formosa was originally in the hands
of Aylmer Bourke Lambert (1761-1842), a man of
private means who accumulated a large herbarium
(Miller 1970). Lambert welcomed botanists to study in
his herbarium and library, and these included George
Don, whose brother David worked for Lambert as his
assistant from 1820 to 1836. After Lambert’s death, his
herbarium was divided into lots and sold in 1843.

In discussing the species, Robert Brown (1849)
appears to have thought that there was more than
one specimen of Cunningham’s since he wrote that
the specimen on which Cunningham based his
name Clianthus Dampieri “cannot now be found
in his Herbarium, as Mr. Heward, to whom he
bequeathed his collections, informs me : nor can I
trace Mr. Lambert’s plant” (i.e. from King’s voyage
in the Mermaid). Given that Cunningham found a
single plant in flower on Malus Island, it is possible
that he gathered just the one specimen that was in
Lambert’s herbarium. Heward gave Bentham a set
of Cunningham’s legumes “in so far as his material
allowed” (Orchard 2014) but there is no specimen of
this collection of Sturt pea at Kew, implying that there
was no duplicate in Heward’s set.

In the Flora Australiensis 2: 214 (Bentham 1864),
this collection was cited as “Dampier’s Archipelago,
A. Cunningham ”. In his Preface in volume 1 of the
Flora , Bentham discussed the collections that he had
used in its preparation. Regarding Allan Cunningham’s
plants he wrote “I have, I believe, been able to
examine the whole of them.” This included a set in
the Hookerian herbarium at Kew as well as a set in
R. Heward’s herbarium that contained Cunningham’s
private herbarium (these specimens are now also at Kew,
donated by Heward in 1862). While there are specimens
of the type of Donia speciosa at BM and K, there is, as
stated above, none of D. formosa. Bentham wrote:

With the few Australian species described from the
herbarium of the late A.B. Lambert, I have had much
difficulty. [...] I have, therefore, in most instances
been obliged to rely chiefly on circumstantial
evidence for the identification of such of these plants
as are only known by the brief diagnoses of G. Don
and others.

While he had further material of Sturt pea to study, this
indicates the difficulty he had in seeing Don’s material.
It seems likely that his citation of Dampier Archipelago
was based upon a reading of Brown’s account of 1849.

In many later accounts treating the species, the presence
of the type collection has received little attention.
When they made the combination Clianthus speciosus ,
Ascherson and Graebner (1909) did not mention
the type collection, nor did they mention the name
Donia formosa. Ford and Vickery (1950) accepted the
collection details of the protologue of Donia formosa
but did not cite a herbarium for the type, while
Thompson (1990, 1993) repeated these details and
cited the holotype as being at the Natural History
Museum (BM). This was repeated by later authors,
e.g. George (1999a), Orchard (2017), but searching
has failed to find it there. Until recently, following
Brown’s comment and being unable to locate the type, I
believed that it had been lost and that a neotype should
be designated. My attention was then drawn to a paper
that discussed the fate of Lambert’s herbarium (Miller
1970) and listed herbaria that held material from
it. From my own searching I knew that the type was
not at Kew (K), the Natural History Museum (BM),
Cambridge (CGE) or Oxford (OXF), so I emailed the
other herbaria listed. I received one positive reply, from
the Phanerogamic Herbarium, Conservatoire et jardin
botaniques, Geneva (G), with a scanned image that
proved to be of the elusive type (Figs 1-2).

The sheet is G 00418563. The annotations on it are:

“New: Holland Curlew River NW Cape. Capt King”,
with ‘Curlew River’ struck out and “ “Malus Island”,
and others of Dampiers Archip 0 - 1818 A.C.” added.

At the foot of the sheet is written: “Clianthus
magnificus G.Don”, with ‘magnificus’ struck out and
an annotation in pencil “formosus” written above.
Another note in pencil reads “vide Dampier voy 3
vol 2”, this refers to Fig. 2 of Tab. 4 in Dampier’s A
Voyage to New Holland etc. in the Year 1699, volume 3
(Dampier 1703).

I do not know the hand of the original, upper
annotation with ‘Curlew River’. The amendments to
it are in Cunningham’s hand. It appears that, when he
published the species in 1832, George Don was not
aware of these amendments.

When compared with a sample of his writing given by
Miller (1970, p. 507), the annotations at the foot of
the sheet appear to be all in the hand of George Don.

50

Swainsona 31 (2018)

The type of Sturt pea found

Fig. 1. The holotype of Donia formosa G.Don, sheet G 00418563 in the Phanerogamic Herbarium, Conservatoire et jardin
botaniques, Geneva, Switzerland (reproduced by permission).

51

A.S. George

Swainsona 31 (2018)

1

S /?

j?**

' S'

/ i . — <-

Fig. 2. Enlargement of the annotations at the foot of the holotype sheet.

The monogram appears to be ‘G.Don’. The pencil
annotation (changing the epithet and adding the
reference to Dampier) are in the same hand—for
example, compare T, rn and W in the two epithets,
and ‘D’ of Dampier with ‘D’ of Don. In changing
the epithet one cannot but wonder if Don took into
account Dampier’s description of the flowers as “very
beautiful”.

The specimen fits Dons description in having obovate
leaves (although this is a common shape for the species)
but particularly in having a racemose inflorescence
(typically it is umbellate).

I conclude that the specimen can be regarded as the
holotype.

The nomenclature for the species is:

Donia formosa G.Don

Gen. Hist. 2: 468 (1832). — Clianthus formosus
(G.Don) Ford & Vickery, Contr. New South Wales
Natl. Herb. 1: 303 (1950). — Swainsona formosa
(G.Don) Joy Thomps., Telopea 4: 4 (1990). —
Willdampia formosa (G.Don) A.S.George, W. Austral.
Naturalist 22: 191 (1999). — Type citation: "’North¬
west coast of New Holland, at the Curlew river, Capt.
King”. — Holotype: W.A., Malus Islands, Dampier
Archipelago [c. 20°31’S, 116°4TE], [1 March] 1818,
A. Cunningham (G 00418563).

Note: The Malus Islands are named not for the
apple, but for French physicist Etienne-Louis Malus
(1775-1812), noted for his studies of light and
optics. Coincidentally they lie less than 20 kilometres
from East Lewis Island, where Dampier collected
his specimens in 1699 (not at Shark Bay as given by
Symon & Jusaitis 2007).

Clianthus dampieri A.Cunn. ex Lindl., Trans. Hort.
Soc. London series 2, 1: 522 (1835), nom. illeg ., nom.
superfl., published as a new name for Donia formosa
G.Don., based on same Cunningham collection and
Dampier’s specimen. — Clianthus dampierii F.Muell.,
Syst. Census Austral. PI. Suppl. 1: 4 (1884), orth. var.

This leaves the question of the ‘Dolichos’ that Frederick
Bedwell brought aboard the Mermaid from his
excursion at the Curlew River on 20 February 1818. At
Kew there is a specimen collected by Cunningham on
the north-west coast in February 1818, determined by

J. Grimes as Cullen stipulaceum (Decne) J.W.Grimes,
but this does not fit Cunningham’s notes as it is a
robust perennial having flowers with a glandular-silky
ovary. A strong possibility is Swainsona pterostylis (DC.)
Bakh.f. which occurs at the mouth of the Ashburton
River, though there is no specimen of the species among
Cunningham’s collections. Another is Canavalia rosea
(Sw.) DC. which occurs commonly along the north¬
west coast but is a robust perennial. Yet another is
Crotalaria cunninghamii R.Br. which is common at the
locality but is a shrub and has pubescent pods. From his
journal it is clear that Cunningham frequently culled
his dried specimens, discarding those that had not
dried satisfactorily or were inadequate for other reasons.
Bedwell’s plant may have met this fate.

Acknowledgements

I am grateful to the editor and the reviewer of an earlier
draft of this paper for suggesting further searching for
the type, and to the herbaria that responded to my
inquiry for this. Tony Orchard, Canberra, confirmed
that the amendment to the locality on the type sheet
is in the hand of Allan Cunningham and assisted with
discussions. I also appreciate discussions with Gwilym
Lewis, Royal Botanic Gardens, Kew. Thomas Pink
and Liz Taylor, Royal Horticultural Society, London,
assisted with bibliographic and chirographic inquiries.

References

Ascherson, P.F.A. & Graebner, K.O.R.P.P. (1909). Synopsis der
Mitteleuropdischen Flora 6: 725, 726. (Wilhelm Engelmann:
Leipzig).

Bentham, G. (1864). Flora Australiensis: A Description of the
Plants of the Australian Territory 2: 214. (Lovell Reeve & Co.:
London).

Brown, R. (1849). Botanical appendix. In: C. Sturt, Narrative
of an expedition into Central Australia performed under the
authority of Her Majesty’s Government, during the years 1844,
5, and6[..i\ 2: 66-92. (T. & W. Boone: London).

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Australian collecting localities. (Australian Biological
Resources Study: Canberra). [Flora of Australia Supple¬
mentary Series 13].

Dampier, W. (1703). A voyage to New Holland &c in the year
1699. (James Knapton: London).

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467-468. (J.G. & F. Rivington [etc.]: London).

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The type of Sturt pea found

Ford, N. & Vickery, J.W. (1950). The correct name of Sturt’s
Desert Pea, Clianthus formosus (G. Don) comb. nov.
Contributions from the National Herbarium of New South
Wales 1:302-303.

George, A.S. (1999a). Willdampia , a new generic name for Sturt
pea. Western Australian Naturalist 22: 191—193.

George, A.S. (1999b). William Dampier in New Holland:
Australia's first natural historian. (Bloomings Books:
Hawthorn).

Government Gazette of South Australia no. 51, p. 1731, 23
November 1961.

Miller, H.S. (1970). The herbarium of Aylmer Bourke Lambert:
notes on its acquisition, dispersal, and present whereabouts.
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Murray, I. & Hercock, M. (2008). Where on the coast is that?
(Hesperian Press: Calisle, W.A.).

With the exception of images and other material protected by a trademark and subject to review by the Government of
South Australia at all times, the content of this publications is licensed under the Creative Commons Attribution 4.0 Licence
(https://creativecommons.Org/licenses/by/4.0/). All other rights are reserved.

© 2018 Board of the Botanic Gardens and State Herbarium (Adelaide, South Australia)

Orchard, A.E. (2014). The dispersal of Allan Cunningham’s
botanical (and other) collections. Telopea 17: 43-86.
Orchard, A.E. (2017). Kennedia speciosa A.Cunn., uncertain no
more. Australasian Systematic Botany Society Newsletter 170:
11-14.

Symon, D. & Jusaitis, M. (2007). Sturt pea: A most splendid
plant. (Board of the Botanic Gardens and State Herbarium:
Adelaide).

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genus Swainsona (Fabaceae) in New South Wales. Telopea 4:
1-5.

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(Fabaceae). Telopea 5: 427-581.

53

w

Swainsona 31:55-58 (2018)

© 2018 Board of the Botanic Gardens & State Herbarium (Adelaide, South Australia) ©0

Micarea kartana sp. nov. (lichenised Ascomycetes) from Kangaroo Island,
South Australia

Gintaras Kantvilas

Tasmanian Herbarium,Tasmanian Museum and Art Gallery, PO Box 5058, UTAS LPO, Sandy Bay, Tasmania 7005
Email: Gintaras.Kantvilas@tmag.tas.gov.au

Abstract: The new species, Micarea kartana Kantvilas & Coppins, is described from Kangaroo Island,
South Australia. A member of the M. prasina Fr. complex, it is characterised by a goniocyst-like thallus
that contains sedifolia -grey pigment and gyrophoric acid, grey to blackish apothecia, and 0-1-septate
ascospores, 10-14 x 4-5.5 pm. It is compared briefly to selected other species of the group.

Keywords: lichens, Pilocarpaceae, taxonomy, new species, South Australia

Introduction

The genus Micarea Fr. is a large and diverse assemblage
of crustose lichen taxa which is widespread in boreal,
austral, temperate and tropical regions, and occurs on a
wide range of substrata, including bark, wood, rock and
soil. Phylogenetic studies (Andersen & Ekman 2005)
indicate that the genus is highly variable and contains
infra-generic groups worthy of generic rank [e.g.
Brianaria (Ekman & Svensson 2014) and Szczawinskia
(Funk 1983)], as well as taxa with their closest affinities
in other genera, for example Psilolechia A.Massal.
(Coppins & Purvis 1987). In addition, several infra-
generic groups can be recognised within Micarea on the
basis of their anatomy, morphology and chemistry, and
may well be separated into distinct genera in the future
(Andersen & Ekman 2005).

In Australia, the genus remains poorly known.
Occasional new species have been published (Stirton
1875; Jatta 1911; Coppins & Kantvilas 1990;
McCarthy & Elix 2016a, b), and some chiefly Northern
Hemisphere taxa have been confirmed as being present
in the region (e.g. Rambold 1989; Coppins 2009).
However, the 24 species listed by McCarthy (2017) are
hardly representative of what is clearly a very species-rich
genus in Australia. Unpublished studies by the author
in Tasmania (in collaboration with B.J. Coppins) have
indicated in the order of a dozen additional, mostly
undescribed taxa there, and potentially many more are
to be found in the extensive unidentified collections
held in Australian herbaria.

The Micarea prasina group, of which the new species
is a member, includes the generitype and was first
recognised as such by Coppins (1983) for species with
a goniocyst-like thallus, strongly convex, immarginate
apothecia and mostly 0-1-septate, ellipsoid-ovoid
ascospores. The group has since been the focus of

chemical (Elix et al. 1984) and morphological and
phylogenetic studies (e.g. Czarnota & Guzow-
Krzeminska 2010) and, as a result, many species have
been recognised (e.g. Coppins & Tonsberg 2001; van
den Boom & Coppins 2001; Czarnota 2007; Coppins
2009; Czarnota & Guzow-Krzeminska 2010; Brand et
al. 2014; Guzow-Krzeminska etal. 2016; van den Boom
et al. 2017). The group is well represented in Australia
where it is especially abundant on rotting wood and
bark in forested habitats, but its taxonomy is poorly
resolved and with most collections ascribed either to
M. prasina Fr. itself, or to the related M. micrococca
(Korb.) Gams ex Coppins. However, additional taxa
are present and several unpublished entities, recognised
by Coppins and Kantvilas (unpubl. ms.) were listed by
Jarman & Kantvilas (2001) and Kantvilas & Jarman
(2012) in their study of wet eucalypt forest, a prime
Micarea habitat.

Here a further member of the M. prasina group is
described from Kangaroo Island, South Australia, and
compared to other members of the complex.

Material and methods

Anatomical and morphological observations were
undertaken using light microscopy, with thin hand-cut
sections mounted in water, 10% KOH, lactophenol
cotton blue, Lugols iodine after pretreatment
with dilute KOH, and ammoniacal erythrosin.
Ascospore measurements are presented in the format:
5 th percentile-^^m^-95 th percentile, with outlying
values given in brackets. Routine chemical analyses by
thin-layer chromatography follow standard methods
(Elix 2014). Comparative data on related species were
derived from the literature as cited. Nomenclature of
pigments follows Meyer & Printzen (2000).

Published online: 11 April 2018 • flora.sa.gov.au/swainsona

ISSN 2206-1649 (Print) • ISSN 2206-1657 (Online)

G. Kantvilas

Swainsona 31 (2018)

The species

Micarea kartana Kantvilas & Coppins sp. nov.

Micareae prasinae s.lat. affinis sed thallo acidum
gyrophoricum etpigmentum “sedifolia-^m^” continenti,
apotheciis sordidis vel denigratis, pigmentum singulare
olivaceum continentibus et ascosporis comparate
late ellipsoideis, 10-14 pm longis, 4-5.5 pm latis
distinguibilis.

Typus: Australia, South Australia: Kangaroo Island,
Grassdale Lagoon, 36°00’S 136°53’E, 20 m alt., on a
moist log of rotting eucalypt in dry sclerophyll forest,
4 Oct. 2013, G. Kantvilas338/15 (holo: HO 580610;
iso: AD, E).

Mycobank number: MB 824888.

Thallus minutely granular, generally bright green when
wet or dry, here and there with dull greyish patches,
composed of coralloid, rather isidioid goniocysts and
forming a brittle, thinly dispersed crust 0.2-0.3 mm
thick; goniocysts 30-45 pm wide, composed of
fascicles of photobiont cells in a mostly colourless gel
matrix with patches of greyish, sedifolia-gtey pigment,
K+ violet, C+ violet; photobiont a micareoid green
alga with ± globose cells 5-11 pm wide. Apothecia
scattered, dull grey to blackish, 0.2-0.6 mm wide,
c. 0.15-0.25 mm tall, plane to convex, sometimes very
strongly so, nestled among the goniocysts, immarginate
from the outset but occasionally a little less intensely
pigmented at the outer rim. Excipulum reflexed and ±
excluded, at most visible only in section in the youngest
apothecia as a layer c. 20 pm thick of loosely entangled,
branched and anastomosing hyphae. Hypothecium
40-80 (-170) pm thick, inspersed with oil droplets,
patchily olive-green, intensifying greenish to yellowish
green in both K and N, C+ yellowish brown and
fading. Hymenium 40-55 pm thick, mostly colourless
but overlain by a patchy layer of olive-green pigment
(as in the hypothecium) which forms vertical bands

between the asci. Paraphyses numerous, branched and
anastomosing, 0.8-1 pm wide, with the apices not
enlarged. Asci 8-spored, 40-52 x 10-15 pm, with a
well-developed, intensely amyloid tholus, pierced by an
indistinct channel that mostly lacks a darker staining
ring structure (approximating the Pilocarpaceae-type).
Ascospores ellipsoid to ovoid, 0-1-septate, hyaline but
with occasional older spores steeped in olive-green
pigment, (9-) 10-77.6-14 (-15) x 4-4.9- 5.5 (-6) pm
(n = 100). Pycnidia not found. Chemistry: gyrophoric
acid; thallus in squash C+ faintly reddish. Figs 1-2.

Etymology. The specific epithet is derived from the
name Karta (meaning island of the dead), given to
Kangaroo Island by Australia’s indigenous inhabitants.

Remarks. The diagnostic characters of Micarea
kartana are summarised in Table 1, where this species
is compared to a selection of other taxa from the
M. prasina group; of these, M. prasina, M. micrococca
and M. viridileprosa have all been recorded for Australia
(McCarthy 2017). Distinguishing members of this
group can be tricky (e.g. see Barton & Lendemer 2014),
and characters that have been usefully applied include
thallus chemistry, apothecial pigments, anatomy and
morphology, ecological and geographical distribution,
as well as DNA-sequence data. The new species has
several highly diagnostic characteristics. For example,
the presence of sedifolia-gtey pigment in the thallus
is commonly encountered in the M. prasina group,
but in such instances, one might expect that the dark
colouration of the apothecia would also be due to this
pigment. Instead, sedifolia- grey is absent in the apothecia
and replaced by an unusual olivaceous pigment that
intensifies greenish to yellowish green in both K and
N-, and reacts C+ yellowish brown. Furthermore, the
ascospores of M. kartana are noticeably wider and more
broadly ellipsoid than in other members of the group.

Fig. 1. Micarea kartana habit (holotype). Scale = 1 mm. Fig. 2. Micarea kartana asci and ascospores, with amyloid

parts stippled (holotype). Scale = 10 pm.

56

Swainsona 31 (2018)

Micarea kortana sp. nov. from Kangaroo Island

Three additional corticolous species of Micarea have
been recorded from Kangaroo Island (unpublished
data). None are common or widespread and all are
easily distinguished from M. kartana. These taxa
include M. prasina, whose distinguishing features are
summarised in Table 1, and two taxa, M. denigrata
(Fr.) Hedl. and M. globusella (Nyl.) Coppins, that both
contain gyrophoric acid and sedifolia-grey pigment, but
with the latter, significantly, confined to the apothecia.
Micarea denigrata differs further by having 0-1-septate
ascospores, 10-12 x 3-4 pm, whereas M. globusella
differs by having ascospores 1,3 (-7)-septate and 13-28
x 2.5-3 pm (Czarnota 2007; Coppins 2009).

Ecology and distribution. The new species is known
only from the type locality in dry sclerophyll forest,
where it grew on a rotting, charred eucalypt log on
the ground, sheltered by undershrubs. The habitat is
fairly typical for Micarea species although, in general,
such forests tend to be rather too open and dry for
these lichens. Thus, whereas the genus is species-rich in
moister, cooler, forested parts of Australia, on Kangaroo
Island, only three additional corticolous or lignicolous
taxa have been recorded to date (unpubl. data; see
above). Lichens associated with the new species
included Leptogium pecten F.Wilson, Carbonicola
foveata (Timdal) Bendiksby & Timdal and Hertelidea
pseudobotryosa R.C. Harris, Ladd & Printzen.

Additional specimen examined

SOUTH AUSTRALIA, Kangaroo Island: type locality,
2015, G. Kantvilas 337/15 (AD, HO).

Acknowledgements

I thank Brian Coppins for his initial comments on a
specimen of the new taxon, and Jean Jarman for the
photograph and for preparing the line drawing for
publication, and two anonymous referees who helped
to improve this paper.

References

Andersen, H.L. & Ekman, S. (2005). Disintegration of
the Micareaceae (lichenized Ascomycota): a molecular
phylogeny based on mitochondrial rDNA sequences.
MycologicalResearch 109: 21-30.

Barton, J. & Lendemer, J.C. (2014). Micarea micrococca and
M. prasina , the first assessment of two very similar species in
eastern North America. Bryologist 117: 223-231.

Brand, A.M., van den Boom, RP.G. & Serusiaux, E. (2014).
Unveiling a surprising diversity in the lichen genus Micarea
(Pilocarpaceae) in Reunion (Mascarenes archipelago, Indian
Ocean). Lichenologist A6: 413-439.

Coppins, B.J. (1983). A taxonomic study of the lichen genus
Micarea in Europe. Bulletin of the British Museum (Natural
History), Botany 11: 17-241.

Coppins, B.J. (2009). Micarea Fr. (1825). In: Smith, C.W.,
Aptroot, A., Coppins, B.J., Fletcher, A., Gilbert, O.L., James,
P.W. & Wolseley, P.A. (eds), The lichens of Great Britain and
Ireland, pp 583-606. (British Lichen Society: London).
Coppins, B.J. & Kantvilas, G. (1990). Studies on Micarea in
Australasia I. Four new species from Tasmania. Lichenologist
22: 277-288.

Table 1. Salient features of selected members of the Micarea prasina complex. All taxa have a goniocyst-like thallus.

thallus

chemistry

apothecia

ascospores

conidia

M. byssacea

(Th.Fr.) Czarnota
et all

green to olive-green,
with sedifolia- grey
pigment

methoxymicareic

acid

pallid to grey to
black, often mottled,
with sedifolia-grey
pigment

(6-) 8-12 (-13) x 2.7-3.5
(-4.2) pm,

0 (-l)-septate

mesoconidia:

(3.8-) 4.5-5.5X 1.2-1.5 pm;
microconidia:

5-7.5 (-8) x 0.8-1 pm

M. kartana

Kantvilas &
Coppins

bright green, with
sedifolia- grey pigment

gyrophoric acid

dull grey to blackish,
with unknown olive
pigment

9-13.5 (-15) X 4-5.5 (-6) pm,
0-1-septate

unknown

M. levicula

(Nyl.) Coppins 2

pale greenish,
becoming ± isidioid

gyrophoric acid

pallid

8-12x2.5-3.5 pm,

0-1-septate

unknown

M. micrococca

(Korb.) Gams ex
Coppins 1

bright green to
olive-green

methoxymicareic

acid

pallid, without
pigments

10-12 (-16) x 3-4.5 pm,

0 (-l)-septate

mesoconidia:

(3.8-) 4.5-5.5 x 1.2-1.5 pm;
microconidia:

5-7.5 (-8) x 0.8-1 pm

M. prasina Fr. 3

pale to dark grey-
green, sometimes
with sedifolia- grey
pigment

micareic acid

pale to dark grey or
blackish, sometimes
with sedifolia-grey
pigment

(7-) 8-12 (-14) X (2.3-) 3-4
(-5.5) pm,

0-1 (-3)-septate

mesoconidia:

(3.5-) 4-6 x 1-1.2 (-1.5) pm;
microconidia:

(5-) 5.5-8 x 0.5-1 pm

M. subviridescens

(Nyl.) Hedl. 3

pale grey-green
to bluish green,
sometimes with
sedifolia-grey pigment

prasinicacid

pallid to grey to
blackish, with
sedifolia-grey
pigment

10-18 x 4-6 pm, (0-)
1-3-septate

unknown

M. viridileprosa

Coppins &v.d.
Boom 2

bright green,
becoming ± sorediate

gyrophoric acid

pallid, without
pigments

8-12 (-14) x 2.5-4 pm,

0-1 (-2)-septate

mesoconidia:

4.5-6 x 1.3-2 pm

M. xanthonica

Coppins &
Tonsberg 4

pale yellow-green

thiophanic acid

pallid, without
pigments

9.3-14 (-14.3) x (3.5-) 3.8-4.2
(-4.7) pm,

(o-) 1 (-3)-septate

unknown

Data sources. 1 Czarnota & Guzow-Krzemiriska (2010); 2 van den Boom & Coppins (2001); 3 Coppins (2009); 4 Coppins &Tonsberg (2001).

57

G. Kantvilas

Swainsona 31 (2018)

Coppins, B.J. & Purvis, O.W. (1987). A review of Psilolechia.
Lichenologist 19: 29-42.

Coppins, B.J. & Tonsberg, T. (2001). A new xanthone-
containing Micarea from Northwest Europe and the Pacific
Northwest of North America. Lichenologist 33: 93-96.

Czarnota, P. (2007). The lichen genus Micarea (Lecanorales,
Ascomycota) in Poland. Polish Botanical Studies 23: 1-199.

Czarnota, P. & Guzow-Krzeminska, B. (2010). A phylogenetic
study of the Micarea prasina group shows that Micarea
micrococca includes three distinct lineages. Lichenologist 42:
7 - 21 .

Ekman, S. & Svensson, M. (2014). Brianaria (Psoraceae), a
new genus to accommodate the Micarea sylvicola group.
Lichenologist 46: 285-294.

Elix, J.A. (2014). A catalogue of standardized chromatographic
data and biosynthetic relationships for lichen substances. Third
Edition. (Published by the author: Canberra).

Elix, J.A., Jones, A.J., Lajide, L., Coppins, B.J. & James, P.W.
(1984). Two new diphenyl ethers and a new depside from
the lichen Micarea prasina Fr. Australian Journal of Chemistry
37: 2349-2364.

Funk, A. (1983). Szczawinskia, a new genus of the lichen¬
forming coelomycetes. Syesis 16: 85-88.

Guzow-Krzeminska, B., Czarnota, P., Eubek, A. & Kukwa, M.
(2016). Micarea soralifera sp. nov., a new sorediate species in
the M. prasina group. Lichenologist 48: 161-169.

Jarman, S.J. & Kantvilas, G. (2001). Bryophytes and lichens at
the Warra LTER site. I. An inventory of species in Eucalyptus
obliqua wet scleophyll forest. Tasforests 13: 193-216.

Jatta, A. (1911). Lichenes lecti in Tasmania a W. Weymouth.
Bolletino della Societci Botanica Italiana 1911: 253-260.

Kantvilas, G. & Jarman, S.J. (2012). Lichens and bryophytes in
Tasmanian wet eucalypt forest: floristics, conservation and
ecology. Phytotaxa 59: 1-31.

McCarthy, P.M. (2017). Checklist of the lichens of Australia
and its island territories. (Australian Biological Resources
Study: Canberra), http://www.anbg.gov.au/abrs/lichenlist/
introduction.html [Version 12 April 2017].

McCarthy, P.M. & Elix, J.A. (2016a). A new species of Micarea
(lichenized Ascomycota, Pilocarpaceae) from alpine
Australia. Telopea 19: 31-35.

McCarthy, P.M. & Elix, J.A. (2016b). Five new lichen species
(Ascomycota) from south-eastern Australia. Telopea 19:
137-151.

Meyer, B. & Printzen, C. (2000). Proposal for a standardized
nomenclature and characterization of insoluble lichen
pigments. Lichenologist 32: 571-583.

Rambold, G. (1989). A monograph of the saxicolous
lecideoid lichens of Australia (excl. Tasmania). Bibliotheca
Lichenologica 34: 1-345.

Stirton, J. (1875). Lichens British and foreign. Transactions of
the Glasgow Society of Field Naturalists 4: 85-95.

van den Boom, P.P.G. & Coppins, B.J. (2001). Micarea
viridileprosa sp. nov., an overlooked lichen species from
Western Europe. Lichenologist 33: 87-91.

van den Boom, P.P.G., Brand, A.M., Coppins, B.J. & Serusiaux,
E. (2017). Two new species in the Micarea prasina group
from Western Europe. Lichenologist 49: 13-25.

With the exception of images and other material protected by a trademark and subject to review by the Government of
South Australia at all times, the content of this publications is licensed under the Creative Commons Attribution 4.0 Licence
(https://creativecommons.Org/licenses/by/4.0/). All other rights are reserved.

© 2018 Board of the Botanic Gardens and State Herbarium (Adelaide, South Australia)

58

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Swainsona 31:59-80 (2018)

© 2018 Board of the Botanic Gardens & State Herbarium (Adelaide, South Australia) ©0

Notes on the taxonomy of Australian Lindernia subg. Didymadenia
(Linderniaceae)

W.R. Barker

State Herbarium of South Australia, GPO Box 1047, Adelaide, South Australia 5001
Email: Bill.Barker@sa.gov.au

Abstract: Names are formalised as a precursor to more detailed taxonomic and evolutionary
accounts of Lindernia subg. Didymadenia in Australia for five newly founded sections (Prolatae
W.R.Barker, Didymadenia (W.R.Barker) W.R.Barker, Scapigerae W.R.Barker, Heterandrae W.R.Barker and
Hemiarrhena (Benth.) W.R.Barker) and 22 new species (L prolata from the near-coastal regions of
central eastern Australia, L. cyanoplectra and L. porphyrodinea from the Kimberley region, Western
Australia, with L. dunlopii extending into Northern Territory, and L. atrata, L. murfetiana, L. dierythra,
L. scopularis, L petrensis, L tiwiensis, L leucochroa, L thyridostoma, L scutellata, L. pustulosa, L. brennanii,
L. lucrusmiana, L. venustula, L. enypniastina, L. acrandra, L. pronanthera, L. robyniae and L. calliandra from
northern Northern Territory). Some of the new species are established as a result of the typification
of several existing names through examination of types or type photographs. The misapplication of
L. lobelioides (F.Muell.) F.Muell. and the partial misapplications of L scapigera R.Br. and L plantaginea
(F.Muell.) F.Muell. are corrected, L. mitrasacmoides (O.Schwarz) W.R.Barker and L. grossidentata
O.Schwarz are resurrected as names of two common Northern Territory species and the separation of
L. tectanthera W.R.Barker from L lobelioides is clarified.

Keywords: Australia, new species, infrageneric classification, subg. Didymadenia, Lindernia,
Linderniaceae

Introduction

Linderniaceae in Australia, segregated relatively recently
from the now dismantled broadly circumscribed
traditional Scrophulariaceae (e.g. Rahmanzadeh et al.
2005), has been, until the last six years, represented by
two genera, the subshrub Artanema D.Don, represented
by a single north-east Australian species, and the genus
Lindernia All. In the 1800s, Brown (1810) initially
placed Australian species in Artanema, Lindernia and
Torenia L., but later authors (Bentham 1846, 1868;
Mueller 1859; Bentham & Hooker 1876) included the
Australian species, many of them today known to be
endemic, in globally widespread genera such as Bonnaya
Link & Otto, Vandellia P.Browne ex L. and, less often,
Llysanthes Raf. Ultimately Mueller (1867, 1882, 1889)
followed Brown’s lead and placed all Australian species,
except Artanema fimbriatum (Hook, ex Graham)
D.Don, in Lindernia.

In the 20 th century, the widespread genera Vandellia,
Bonnaya and Llysanthes were subsumed in a widely
circumscribed Lindernia as subgenera or sections in the
works of Pennell (1935, 1943a, 1943b) on north-east
American, Himalayan and New Guinean species, Philcox
(1968) on Malesian species, and Yamazaki (1978a,
1978b, 1980, 1981, 1985, 1990) on Asian species.

Hemiarrhena Benth. (1868), a monotypic Australian
genus of doubtful tribal affinities, was universally

maintained as a distinct genus. Originally described by
Bentham as a phyletically isolated member of the tribe
Rhinantheae Lam. & DC., a view adopted in other works
on the Scrophulariaceae (e.g. Wettstein 1891-1893),
it was always maintained in the Lindernia alliance by
Mueller (1889), who viewed it and other species of
Lindernia in the field and described it from his own
collections. In the late 1900s it was returned to Lindernia
in Australian works, starting with Barker (1983) and
Dunlop (1987), with Barker (1990) including it in his
new subg. Didymadenia , which he established for the
main Australian radiation of Lindernia.

A recent molecular study by Fischer et al. (2013)
reconstituted Lindernia , giving it a narrow circum¬
scription covering the type and allied species, and
resurrecting the globally widespread genera Bonnaya and
Vandellia. Apart from some wide-ranging tropical species
placed in Bonnaya , they removed all Australian species
from Lindernia to Vandellia, although their approach was
based on limited evidence (Biffin et al., in press).

In an analysis of new molecular sequence data (Biffin
et al., in press), the bulk of Australian species have
been returned from Vandellia to Lindernia. The
following genera of Linderniaceae are now recognised in
Australia: Artanema, Bonnaya, Lindernia, Torenia and an
unpublished genus established in Biffin et al. (in press).
Three subgenera of Lindernia are confirmed, the type
L. procumhens (Krock.) Philcox in subg. Lindernia, east

Published online: 2 May 2018 • flora.sa.gov.au/swainsona

ISSN 2206-1649 (Print) • ISSN 2206-1657 (Online)

W.R. Barker

Swainsona 31 (2018)

Australian L. hyssopioides (L.) Haines in a subgenus based
on the generic name Ilysanthes (Biffin et al., in press),
and the main radiation of Linderniaceae restored to
subg. Didymadenia W.R.Barker.

The establishment of a firm generic framework has
been critical to publishing the following taxonomic
changes and additions which further expand the known
diversity of Australian Linderniaceae and particularly its
subg. Didymadenia (Barker 1990, 1992a, 1992b, 1998,
2000; Wannan 2013, 2016). More detailed taxonomic
advances (Barker, in prep.) and an evolutionary study
(Barker & Biffin, in prep.) of the genus in Australia are
well-advanced towards publication.

Diagnostic attributes

Hair types

Glandular hairs 0.2-1.0 mm long with multicellular
uniseriate stalks are evident in many species across
the family. Their presence or absence and relative
length on vegetative parts and in the inflorescence can
be of diagnostic value at the species or infraspecific

Fig. 1. The subepidermal gland in Lindernia subg. Didy¬
madenia. A, B Transverse section of leaf of Lindernia mitra-
sacmoides [W.R. Barker 7045, AD), stained in Toluidine Blue O.
C View through top of leaf of Lindernia dunlopii ( R.M . Barker
284, AD), cleared with KOH and stained with Sudan IV. — Scale:
A, B = 100 pm; C = 50 pm.

level. Other hairs are of diagnostic value outside
the subgenus. These tiny hairs, terminated by four-
celled glands c. 13-30 pm in diameter, occur across
the family; such hairs have been seen, for example, in
samples of Bonnaya and the subgenera Lindernia and
Didymadenia of Lindernia. Apparently homologous
hairs occur widely across the families of the traditional
Scrophulariaceae (Barker, pers. obs.). Antrorse narrow
deltoid single-celled eglandular hairs occur on leaf, bract
and sepal margins in many genera of Linderniaceae;
they are absent from subg. Didymadenia although a few
species in sect. Hemiarrhena (Benth.) W.R.Barker bear
probably homoplasious soft antrorse eglandular hairs
along the sepal margins.

Subepidermal glands

Lindernia subg. Didymadenia is defined by 2-celled
glands, elliptic in outline, which are found midway
between upper and lower surfaces of the leaves (Fig. 1),
on stems and branches, bracts and sepals and sometimes
the corolla and capsule. While always embedded in the
cellular matrix below the cuticle, they are often evident
externally, through the cuticle. On occasions (e.g. in
L. dunlopii and L. pustulosa) the surface of green parts
of the plant (e.g. leaves, bracts and sepals) is raised
above each gland, or in other species these parts and the
capsule surface are ‘spotted’ arising from the visibility of
the glands through the epidermis. The descriptive term
‘blister glands’ is proposed for these instances.

Seeds

Seed morphology is a promising area of study for
establishing synapomorphies for defining genera and
infrageneric groups in the Linderniaceae (Biffin et al .,
in press).

While seeds in the newly erected sect. Didymadenia
have four longitudinal ribs (Barker 1990, 1992a),
they can be more variable in other sections of subg.
Didymadenia , with some species having up to 8-ribbed
seeds. The concavities between the ribs are generally
relatively deep furrows, but in some species of sect.
Hemiarrhena the seeds are almost circular in transverse
section. The ribs are bridged by finer transverse ridges,
varying in number from about four to 20, which are in
some species confined to the intervening furrows, while
in others they connect across the longitudinal ribs.

Scent

Some of the Australian species of subg. Didymadenia are
noticeably scented. The source of the scent is likely to
be the 2-celled subepidermal glands, although whether
they contain aromatic oil still has to be proven.

Herbarium material examined

Specimens from PERTH, DNA and AD form
the principle basis of this paper, supplemented by
observation of material on visits to BRI, MEL and
CANB and examination of specific specimens by
staff or associates in PERTH, MEL and NSW. The

60

Swainsona 31 (2018)

Notes on the taxonomy of Australian Lindernia subg. Didymadenia

specimen citations reference only duplicates that have
been examined for this study

The material of L. grossidentata O.Schwarz and
Ilysanthes mitrasacmoides O. Schwarz used in the
protologues (Schwarz 1927) was sent to Berlin by
Darwin-based collector F.A.K. Bleeser. Many of the
Berlin collections were destroyed in World War II,
including the B syntypes of these species, while Bleeser’s
own herbarium was destroyed after a cyclone (McKee
1963; Willis 1966; George 2009). Their absence from
the B collection has been confirmed by Dr R. Vogt
(pers. comm., 9 Apr. 2018). Herbarium web sites have
been searched for type material. Duplicates of Bleeser
collections survive, principally in MEL, NSW and
K, with some secondarily in DNA. The Australasian
Virtual Herbarium (2017) provides details of 230
Bleeser collections, mainly in MEL and NSW, while
an on-line search of the specimen image database
on JSTOR Global Plants (2017) revealed 27 types
collected by Bleeser, including some duplicates, mostly
in MEL and NSW, as well as K, B, BM and WIS, but
no syntype material, identifiable by annotations by
Schwarz, has been located. Fortunately, isosyntypes of
Ilysanthes mitrasacmoides are housed in NSW and DNA.

Taxonomy

Lindernia subg. Didymadenia W.R.Barker

J. Adelaide Bot. Gard. 13 (1990) 79. — Type: L. chryso-
plectra W.R.Barker

Hemiarrhena Benth., FI. Austral. 4 (1868, as 1869)
318; Benth., Hooker’s Icon. PI. 11 (Ser. 3: 1) (1870) 46,
pi. 1059. — Type: L. plantaginea (EMuell.) EMuell.

Bradshawia EMuell., Proc. Linn. Soc. New South
Wales, Ser. 2, 6 (1892) 473, nom. prov., based on
L. macrosiphonia (EMuell.) W.R.Barker.

Annuals, rarely perennial herbs; subepidermal glands
present in the stems, leaves, sepals and often capsule
wall. Vestiture: leaf, bract and calyx margins not
scabrous from antrorse robust aculeate eglandular hairs,
rarely with scattered soft eglandular hairs. Stamens: two
pairs inserted on the abaxial and adaxial side of the
corolla tube, sometimes one of the pairs lost or reduced

to staminodes; lower filaments bearing a lateral spur,
rarely the spur reduced to a knob or absent; anthers
with two confluent opposed, sometimes obliquely so,
cells, sometimes 1-celled through reduction or loss of a
cell. Stigma 2-lobed. Capsules narrow obloid-ellipsoid to
globular; seeds many, longitudinally 4-8-ribbed, with
a few to many transverse secondary ridges across the
intervening furrows.

Distribution. Mostly confined to northern and
northeastern tropical and subtropical Australia; two
species, L. suhulata and L. scapigera , extend to southern
New Guinea.

Note. Five sections within this subgenus are defined
below.

A. Lindernia sect. Prolatae W.R.Barker, sect. nov.

Typus: L. pro lata W.R. Barker

Annuals; stems and branches erect or narrowly
ascending, terminated by open erect racemes. Leaves
sessile and subulate or subpetiolate, narrow to broadly
ovate. Pedicels erect to widely spread. Corolla regular,
without internal longitudinal flaps; tube narrow,
elongated, conspicuous, as long or longer than the
limb; upper lip porrect, emarginate. Stamens 4, the
abaxial filaments with a cylindrical spur; anthers
2-celled. Capsules ovoid to subglobular; seeds with
4-8 longitudinal ribs and c. 6-20 secondary transverse
ridges.

Distribution. A section spread from the Dampier
Peninsula near Broome, Western Australia, across
tropical and subtropical northern Australia and eastern
Australia as far as the north coastal region of New
South Wales. This matches the geographical range of
Lindernia in Australia.

Note. The section includes L. subulata R.Br., L. beasleyi
Wannan, L. stantonii Wannan and the species newly
described here.

Etymology. The epithet of the type species (q.v.) was
chosen in part as the resultant sectional name reflects
the broad geographical distribution of the section.

Key to sections of Lindernia subg. Didymadenia

1. Perfect anthers 4

2. Anthers 2-celled

3. Corolla tube and hood not infolded, the stamens and style evident from below

4. Leaves entire, subulate and sessile or ovate and sessile or subsessile. A. sect. Prolatae

4: Leaves entire through sinuate to dentate, subpetiolate, ovate or broadly so. C. sect. Scapigerae

3: Corolla tube infolded, the adaxial filaments and style positioned above the folds . . . B. sect. Didymadenia

2: Anthers of adaxial pair 2-celled, of abaxial pair 1-celled.D. sect. Heterandrae

1: Perfect anthers 2,1 - or 2-celled, with or without a pair of abaxial staminodes.E. sect. Hemiarrhena

61

W.R. Barker

Swainsona 31 (2018)

Key to species of Lindernia sect. Prolatae

1. Inflorescence (rachis, bracts, pedicels and sepals) glabrous

2. Leaves ovate to ± circular. 1. L. prolata

2: Leaves narrow linear to subulate

3. Sepals red- to black-lined; middle lobe of the lower corolla lip white with two blue-purple blotches. . 2. L. atrata
3: Sepals not lined; corolla not as above

4. Pedicels more than 10 mm long

5. Corolla lobes spotted; ovary glandular pubescent. L subulata

5: Corolla lobes not spotted; ovary glabrous . L. beasleyi

4: Pedicels less than 10 mm long. L. stantonii

1: Inflorescence glandular hairy

6. Corolla white to blue without red spotting; lower corolla lip with a green blotch

7. Glandular hairs in inflorescence 0.01 -0.025 mm long. 3. L. cyanoplectra

7: Glandular hairs in inflorescence 0.1 -0.15 mm long . 4. L. murfetiana

6: Corolla yellow with red spots. 5. L. dierythra

1. L prolata W.R.Barker, sp. nov.

Holotypus: R.W. Jobson 1291, 1 Oct. 2011, Kingaroy,
Queensland (NSW 885850). Isotypi: AD 279775, BRI.
L. sp. Tingoora (A.R. Bean 10311) Queensland Herba¬
rium: N.Fechner in Bostock & A.E.Holland, Census
Queensland FI. 2007 (2007) 188.

Multi-branched Pannual herb, the main branches
c. 3.5 _ 19 cm long, decumbent to ascending, rooting
at basal nodes. Leaves ovate to ± circular, 3~10 mm
long, sessile, narrow cuneate, sometimes subpetiolate.
Inflorescences open erect terminal racemes; pedicels
erect, sometimes spreading in fruit, much longer than
the bracts. Corolla (4.0-) 6-8.5 mm long along upper
side, blue to blue-purple, rarely white; lower lip with
a white mid lobe, with dark blue-purple base and two
blotches. Stamens 4; abaxial spur exserted, cylindrical,
pale blue or bluish-white; anthers 2-celled. Capsule
obloid-ovoid to globular, 2.5 - 3.5 mm long, glabrous;
seeds {Johnstone 2355 & Orme) obloid-ellipsoid, rarely
broadly so, 0.35 - 0.45 (-0.5) mm long, with 6 sharp
longitudinal ribs.

Distribution. In eastern Australia in regions neighbouring
coastal south-eastern Queensland and north-eastern New
South Wales. In seasonally inundated areas.

Etymology. The adjectival epithet derives from the
Latin adjective prolatus , extended or elongated, alluding
to the very long pedicels at flowering time; also
influenced by resultant sectional name (q.v.).

Selected additional specimens

QUEENSLAND: 7.2 km along Dangore Mtn road, WSW
of Tingoora, 21 May 1996, A.R. Bean 10311 (AD); 33 km
NNE of Chinchilla, 1 Oct. 1993, M.E. Ballingall2733 (AD);
Hellhole Creek, near Auburn Road in Barakula Forestry
[Reserve], 6 Mar. 1984, V Hando s.n. (AD 98413060).

NEW SOUTH WALES: C. 2 km (direct) E of Coopernook
along Spring Hill Road, 30 Apr. 2008, R. Johnstone 2355 &

A.E. Orme, (NSW, as photo, AD, seeds only); 20 km W of
Wooli, 23 Nov. 1987, R. Bates 12879 (AD).

2. L. atrata W.R.Barker, sp. nov.

Holotypus: ID. Cowie 12020, 22 Apr. 2008, Limmen
National Park, c. 24 km S of Ranger Station, Northern
Territory (AD 246363). Isotypi: DNA {n.v.), B {n.v.).

Erect glabrous single-stemmed annual, 12-50 cm tall.
Leaves deltoid-linear to filiform, the longest on a stem
5~ 12 mm long, sessile, entire. Inflorescences open simple
terminal racemes of 1-7 or more flowers; pedicels
longest at lowest node, ascending, 5 _ 22 mm long,
much longer than the subulate bracts. Corolla 5 - 8 mm
long along the upper side, mid blue throughout,
sometimes with fine red-purple striations, with two
mid yellow blotches, one behind the other, on the lower
side of the mouth and throat and with two deep purple
blotches at base of lowest lobe. Stamens 4; abaxial spur
exserted, cylindrical, white; anthers 2-celled. Capsule
ellipsoid to globular, 2-2.3 mm long, glabrous; seeds
obloid, 0.25 - 0.28 mm long, c. (?5 - ) 6 longitudinal
sharp ribs, with 9~10 fine secondary ridges across
intervening furrows.

Distribution. In the Limmen region of Northern
Territory, on the west coast of the Gulf of Carpentaria,
south of Arnhem Land.

Etymology, from the Latin adjective atratus, dressed
in black, derived from ater, black, dark, and the
suffix -atus, meaning provided with, alluding to the
conspicuous red- to black-lined margins of the sepals.

Selected additional specimens

NORTHERN TERRITORY: Limmen National Park, c. 12
km SW of Nathan River Ranger Station, Site 70, 18 Apr.
2008, P.S. Short 5513 (DNA); Cox River Station, 30 June
1977, T.S. Henshall 1568 (DNA).

62

Swainsona 31 (2018)

Notes on the taxonomy of Australian Lindernia subg. Didymadenia

3. L cyanoplectra W.R.Barker, sp. nov.

Holotypus: B.J. Carter 679 , 6 Sep. 1993, Taylors
Lagoon, c. 76 km ENE of Broome, Western Australia
(PERTH 3234320). Isotypus: AD 270009.

Annual, 7“ 10 cm high, with erect to decumbent
branches, with the inflorescence covered by tiny
glandular hairs to c. 0.02 mm long. Leaves sessile,
subulate, 3 — 15 mm long, dilated at the base. Inflorescence
an open terminal raceme, with pedicels much longer
than the subulate bracts. Corolla 4.5 _ 6.5 mm long along
upper side, light to mid blue, the lower lip with a green
blotch. Stamens 4; abaxial spur long, exserted, blue;
anthers perfect, blue, 2-celled. Capsule broadly ellipsoid,
1.8-2.8 mm long, glabrous; seeds ( Carter 679) ellipsoid,
c. 0.3-0.33 mm long, with 4-3 (-?6) longitudinal ribs.

Distribution. South and east of the Dampier Peninsula,
northern Western Australia, in seasonally inundated areas.

Etymology. The adjectival epithet derives from
the Greek kyanos , dark blue, and plektron , a tool
for plucking or spur, alluding to the colour of the
conspicuous staminal spurs of this species.

Selected additional specimens

WESTERN AUSTRALIA: C. 75 km E of Broome, just S of
highway to Fitzroy Crossing, 26 June 2007, D. Coultas & B.
Taylor Opp 10 (PERTH); Edge of Nemila Creek near Blina
Swamp, 21 Mar. 2010, D. Murfet 6786 & A. Lowrie (AD);
55 km E of Broome, just S of highway to Fitzroy Crossing,
27 June 2007, Woodman & K. Greenacre Y1 SI 1-01 (PERTH).

4. L. murfetiana W.R.Barker, sp. nov.

Holotypus: W.R. Barker 9129 & K. Brennan , 19 May
2013, on vehicular track c. 250 m SE of Litchfield
Park Road, c. 1.2 km direct S of Finnis River
Crossing, Northern Territory, Darwin & Gulf District
(AD 268115). Isotypi: BRI, CNS, CANB, DNA,
PERTH.

L. sp. Hann River (M. Lazarides 9167) [W.R.Barker]:
R.A.Kerrigan & Alb., Checkl. N. Territory Vase. PI.
(2007).

Erect, rarely decumbent, annual herb, (10-) 20-25 cm
tall, glabrous but for the moderately dense to dense
glandular hairs in the inflorescence. Leaves sessile,
filiform to narrow ovate-caudate, near base of plant
3 _ 8 mm long but much shorter distally. Inflorescence a
terminal few-flowered raceme. Corolla 9~10.5 mm long
along the upper side, white, yellow or blue, with green
blotches on the lower lip. Stamens 4, with abaxial spurs
exserted from the corolla mouth, cylindrical, long;
anthers white, 0.8-1.0 mm long, 2-celled. Capsule
broad ellipsoid to globular, 2.3~2.5 mm long, glabrous;
seeds ellipsoid-obloid to obloid, 0.2-0.3 mm long, with
6-8 longitudinal ribs.

Distribution. Mainly in the Darwin to Litchfield area
of northern Northern Territory, with outliers further
west in Arnhem Land; in seasonally inundated areas.

Etymology. Named after Denzel Murfet, an avid
collector associated with the State Herbarium of South
Australia, whose many collections and associated
floral photographs from Northern Australia have
been valuable in the unravelling of the taxonomy of
Lindernia.

Selected additional specimens

NORTHERN TERRITORY: Howard Springs, S side of
ephemeral lagoon area (Sect. 3950), over fence bordering
Sections 3953 and 3954, ca. 1 km E of Dutchies Lagoon, c.
100 m N of Corella Avenue, 29 May 1997, W.R. Barker 7735
& R.M. Barker (AD); 300 m from Cox Peninsula Road and
Bynoe Harbour road, 6 July 2008, D.E. Murfet 6054 (AD);
Edge of McMinns Lagoon, 26 June 1968, D. Wheelwright 45
(AD, DNA); 19 km NNW of Twin Falls, 3 June 1980, M.
Lazarides 9167{CANB, DNA).

5. L. dierythra W.R.Barker, sp. nov.

Holotypus: W.R. Barker 7873 & K. Brennan, 16 Apr.
1999, near Lost City [precise locality withheld for
conservation reasons], Litchfield National Park,
Northern Territory (AD 268129). Isotypi: B, BRI,
CANB, CNS, DNA, K, MEL, MO, NSW, PERTH.

L. sp. Litchfield (I.D. Cowie 5725) [W.RBarker]: R.A.Kerri-
gan & Alb., Checkl. N. Territory Vase. PI. (2007).

Erect, finely glandular-pubescent, annual herb,
3.5“17 cm high, often a simple single stem. Leaves
sessile, narrow linear, longer at higher nodes, the longest
8-15 mm long. Inflorescence an open terminal raceme or
appearing paniculate when upper branches are present;
pedicels longer than the bracts, usually reflexed in fruit.
Corolla 6.5“8 mm long along upper side, bright yellow
with red spots on the lower lip. Stamens 4, the abaxial
spurs short, cylindrical, yellow; anthers 0.7-0.8 mm
long, 2-celled. Capsules broad ovoid, c. 3.5 mm long,
glandular-pubescent; seeds obloid or broadly so,
c. 0.4 mm long, with 4 blunt longitudinal ribs.

Distribution. Near The Lost City, Litchfield National
Park, Northern Territory in grassy herbfield on sand.

Note. As observed by I.D. Cowie (. 5725 ), this species
mimics taxa such as Uvedalia linearis R.Br. var. lutea
(Benth.) W.R.Barker & Beardsley in the rare attribute
of having yellow flowers with prominent red spotting
on the lower side of the corolla.

Etymology. The epithet dierythra is an adjective
meaning spotted or variegated with red (Brown 1956),
alluding to the red-freckled corolla mouth; it is derived
from the elision of the Greek dia-, a prefix for indicating
separation, and erythros, red.

Additional specimen examined

NORTHERN TERRITORY: Near Lost City, 16 Apr. 1995,
I.D. Cowie 5725 (AD, DNA).

63

W.R. Barker

Swainsona 31 (2018)

B. Lindernia sect. Didymadenia (W.R.Barker)

W.R.Barker, stat. nov.

Lindernia subg. Didymadenia W.R.Barker, J. Adelaide

Bot. Gard. 13 (1990) 79, basionym. — Type:

L. chrysoplectra W.R.Barker

Annuals or short-lived perennials; stems and branches
erect or narrowly ascending, terminated by open
erect racemes, or prostrate, procumbent or pendent
(from rock faces) with terminal inflorescences not
strongly demarcated through the foliose bracts. Leaves
subpetiolate, narrow to broadly ovate to obovate.
Pedicels erect or ascending, in fruit sometimes deflexed,
rarely elongating and extending into rock crevices.
Corolla regular or resupinate, invaginated on either side
into a pair of internal flaps; tube usually conspicuous,
rarely much shorter than the limb; upper lip porrect or
upturned, emarginate. Stamens 4, or 2 with 2 adaxial
(abaxial in resupinate flowers) staminodes; abaxial
filaments with a cylindrical or reduced spur, rarely

lacking; perfect anthers 2-celled. Capsules ovoid to
subglobular; seeds 4-furrowed, with c. 6-20 transverse
ridges.

Distribution. Most species occur in the Kimberley
region of northern Western Australia, but a few occur
in northern Northern Territory and north Queensland.
In wet situations, woodland, stony ground and on rock
faces.

Notes. The section includes the following newly
described species, as well as L. cleistandra W.R.Barker,
L. eremophiloides W.R.Barker, L. macrosiphonia (F.Muell.)
W.R.Barker, L. hypandra W.R.Barker, L. chrysoplectra
W.R.Barker, L. aplectra W.R.Barker, L. tectanthera
W.R.Barker and L. barkeri Wannan.

The flowers, fruits and seeds of L. macrosiphonia are
remarkably large.

Key to species of Lindernia sect. Didymadenia

1. Basal leaves subulate or narrow linear

2. Corolla 9-10 mm long; anthers 1.2 mm long; lower filaments golden-spurred. L. chrysoplectra

2: Corolla 5.5-7.3 mm long; anthers 0.4 mm long; lower filaments without spurs . L. aplectra

1; Basal leaves foliose, spathulate

3. Flowers subtended byfoliose bractsthroughoutthe inflorescence; plantfoliose, procumbent on ground

or pendent from vertical rock faces

4. Corolla tube 70-100 mm long. L. macrosiphonia

4: Corolla tube up to 15 mm long along the upper side

5. Leaves alternate; sepals fused for Va to Vs their length. L. eremophiloides

5: Leaves opposite; sepals free almost to base

6. Pendent perennial, growing on rock faces

7. Pedicels hardly lengthened in fruit, exposed; capsule 5.5-9.5 mm long. 6. L. scopularis

7: Pedicels turned back and elongated into dense foliage or deep into crevices;

capsule 3.5-5.5 mm long . L. cleistandra

6: Procumbent annual . 7.L. petrensis

3: Flowers subtended by narrow linear to subulate bracts, apart sometimes for the basal nodes of
the inflorescence; plant erect annual, with cluster of leaves at base, and stem with scattered leaves
reduced in size or a scape

8. Flower regularly oriented

9. Staminodes without a spur

10. Staminodes outcurved, conspicuous, yellow, terminated often with an obscure vestigial
anther often with a very short to short filiform white filament; robust, erect, scapiform
herb; leaves usually confined to a basal rosette, sometimes of reduced size at a few
nodes up the stem or main branches. L. tectanthera

10: Staminodes porrect, white; delicate erect to scandent herb; leaves broadest in middle

nodes of stem up to the node below the inflorescence. L. barkeri

9: Staminodes porrect to outcurved, with a terminal spur and a short to long filament arising

behind its tip. 8. L. lobelioides

8: Flower resupinate

11. Corolla tube narrowly dilated. L. hypandra

11: Corolla tube broadly dilated. 9.L. porphyrodinea

64

Swainsona 31 (2018)

Notes on the taxonomy of Australian Lindernia subg. Didymadenia

6. L scopularis W.R.Barker, sp. nov.

Holotypus: W.R. Barker 7080 & R.M. Barker,
5 June 1994, Bessie Springs, Northern Territory (AD
99818368). Isotypi: BRI, CANB, CNS, DNA, GH,
K, MEL, NSW, PERTH.

L. sp. Cliff lover (P.K Latz 10123) [WR.Barker]: R.A.Kerri-
gan & Alb., Checkl N. Territory Vase. PL (2007).

Perennial glabrous herb, usually with pendent branches
to c. 40 cm or more long, or, when smaller, forming
small cushions. Leaves spathulate, (1-) 2-4 (-6) cm
long, usually coarsely serrulate, sometimes entire.
Inflorescences terminal, leafy, with to 10 or more flowers,
with pedicels shorter than the leafy bracts, c. 1-10 mm
long, in fruit recurved but not elongating. Corolla
8-10 mm long along the upper side, usually white,
with fine blue striations extending from tube to lobes,
sometimes “pale lilac” or “pale mauve”. Stamens 2,
in adaxial position, the abaxial pair reduced to
staminodes; adaxial anthers 1-1.4 mm long, 2-celled;
abaxial staminodes obloid, yellow or white, tipped by a
very short lateral filament and vestigial anther. Capsule
ovoid, 5.5~9.3 mm long, glabrous; seeds narrow obloid,
often curved, 1.4-1.8 mm long, 4-angled, with 10-12
transverse ribs between.

Distribution. Inland from the west coast of the Gulf
of Carpentaria, Northern Territory, east-northwest and
southwest of Borroloola in the Nathan River station
area to the Abner Range (Bessie Springs).

Etymology. The adjectival epithet is derived from the
Latin scopulus , cliff, and the suffix -aris, belonging to,
alluding to the cliff face habitat of this species.

Selected additional specimens

NORTHERN TERRITORY: 28 km S Nathan River
homestead, 11 Sep. 1995, P.K. Latz 14547 (AD); McArthur
River area, 27 Jan. 1976, L.A. Craven 3413 (NT); 11 km S of
Balbarini Homestead, 10 Nov. 1988, P.K. Latz 11041 (AD,
DNA).

7. L. petrensis W.R.Barker, sp. nov.

Holotypus: W.R. Barker 7838 & K. Brennan, 5 Apr.
1999, on south face of Nourlangie Rock, c. 200 m
N and above Anbangbang rock art site, Northern
Territory (AD 279073). Isotypi: BRI, CANB, CNS,
DNA, PERTH.

L. sp. Kakadu (J.L. Egan 4819) [WR.Barker]: R.A.Kerrigan
& Alb., Checkl. N. Territory Vase. PI. (2007).

Procumbent, rarely scandent, much-branched, glan¬
dular-pubescent annual, with stem and main branches
3 _ 30 cm long. Leaves spathulate, 0.2-4 cm long,
serrulate, rarely entire. Inflorescence rarely solitary,
usually a few flowers in a leafy open raceme; bracts leaf¬
like or shorter, 0.3-2.5 cm long, the pedicels to 2.5 cm
long, deflexed in fruit. Corolla 6.5 - 9 mm long along
upper side, with white tube and mid blue-purple limb.
Stamens 2, adaxial perfect, the anthers 0.5 - 0.6 mm
long, 2-celled, deep blue; abaxial staminodes linear,

golden yellow, tipped by a vestigial anther. Capsule
globular, 2.5 _ 4.2 mm long, glabrous; seeds obloid,
0.8-1.3 mm long, 4-angled.

Distribution. Across northern Arnhem Land, Northern
Territory, and to the south and south-west on Bradshaw
Station. Between sandstone rocks in dense sorghum
over herbfield or in bare sandstone gravel.

Etymology. An adjectival epithet from the Greek petra,
rock, shelf or ledge or rock; petrensis, among rocks,
alluding to the habitat on flatfish ground in skeletal soil
on or between rocks.

Selected additional specimens

NORTHERN TERRITORY: Arnhem Land, c. 81 km SSW
of Maningrida, 23 Mar. 2000, ID. Cowie 8738 (DNA); East
of entry to Merl camping area, 21 Apr. 1999, W.R. Barker
7887 & I. Cowie (AD); Upper Liverpool River, Chester Ck
Kulnguki area, 20 Apr. 2009, KG. Brennan 7983 (DNA);
Bradshaw Station, near Fire Plot 3, 18 Feb. 1999, C. Michell
2183 (DNA).

8. L. lobelioides (F.Muell.) F.Muell.

Syst. Census Austral. PI. (1882) 97. — Vandellia
lobelioides F.Muell., Trans. Philos. Inst. Victoria
3 (1859) 61, basionym (“Sect. Bonnaya”) [non
V lobelioides Oliv., Trans. Linn. Soc. London 29 (1875)
120, nom. illeg. (= Craterostigma newtonii (Engl.) Eb.
Fisch., Schaferh. & Kai Miill.)]. — Ilysanthes lobelioides
(F.Muell.) Benth., FI. Austral. 4 (1868) 498. — Type
citation: “A companion of Vandellia clausa to which it
stands in close affinity. ... [From treatment of V clausa
referred to in the protologue] On sand-plains, subject
to occasional inundations, on the Victoria River and
its tributaries.” — Lectotypus (hie designatus):
F. Mueller s.n., s.dat., Victoria River, from the Depot
Creek to the main camp [the latter at c. 15°34'S,
130°22'E] / Dr M[ueller] s.n., May [18]56, Victoria
River, below Steep Head, [c. 15°33'S, 130°55'E]
(MEL 1552811; Fig. 2). Isolectotypus: Dr M[ueller]
s.n., s.dat., Victoria River (K 859763; Herb. Hooker)
(“Bonnaya (Ilysanthes) lobelioides ferd. Mueller” in
Mueller’s hand). Syntypus possibilis sed exclusus:
F.Muell. s.n., s.dat., Victoria River (MEL 1552812;
“Vandellia lobelioides ? / Lindernia” in Mueller’s hand).

Typification. The lectotype was collected between 6 th
and 9 th May 1856. During this period, the Gregory
exploring party returned to the Bynoe Range on the
Gregory River where Steep Head is located (Birman
1979).

The MEL lectotype is the better of the two syntypes; it
has a clearly discordant label (the mid in the right-hand
group of three) with handwritten notes and a sketch
by Mueller showing unilocular anthers matching
those of L. clausa (F.Muell.) F.Muell., which Mueller
collected on the same expedition (MEL 1552814,
MEL 1552815).

The collection MEL 1552812 has also been considered
a potential syntype. However, it is not this species; the
capsules are elongated as evidenced by the distinctly
narrow septum.

65

W.R. Barker

Swai/isonaS 1 ! (2018)

SYN-

TYPE

VuJlfelW lc>Lcl'cicJjL'5> EM

MEL! 552811

BOTANICAL MUSEUM OF MELBOURNE.

•py' ^y P' ■' si - 'P

PEKD. MI KIJ.KK, I'll. * MJ).

NATIONAL HERBARIUM OF
VICTORIA (MEL), AUSTRALIA

Fig. 2. Lectotype of Vandellia
lobelioides F.Muell. at the
National Herbarium ofVictoria
(MEL 1552811; reproduced
with permission from the
Royal Botanic Gardens
Victoria).

<x \obe\v6l<jgS
S"ffiL"v\AO<ijLCV oQ (j[£.fo.c^N®A.

(d.r&.viA T-W-ZfcV^

Fig. 3. Staminodes in lectotype of Vandellia lobelioides F.Muell. (MEL 1552811). Del. Neville Walsh (published with his permission).

66

Swainsona 31 (2018)

Notes on the taxonomy of Australian Lindernia subg. Didymadenia

Notes. The descriptions of L. tectanthera by Barker (1990,
1992a) encompass both L. lobelioides and L. tectanthera
except in the description of the staminal spurs.

L. lobelioides and L. tectanthera are both widespread in
the Kimberley and are very similar in almost all respects,
in floral shape and coloration, and in habit, although the
latter is rosulate with multiple scapes, the former usually
erect with larger leaves above the ground, and a single
terminal inflorescence. The staminodes, however, are
very different morphologically, as presented in the above
key. Results of DNA analysis (Barker & Biffin, in prep.)
show the two taxa to be phyletically well separated.

9. L porphyrodinea W.R.Barker & M.D.Barrett, sp. nov.
Holotypus: M.D. Barrett4647&KW. Dixon, 20 Mar.
2015, near Bachsten Creek [precise locality withheld
for conservation reasons], Western Australia (PERTH
8926859). Isotypi: AD, BRI, CANB, DNA, K.

Erect glabrous annual herb to c. 20-35 cm high. Leaves in
rosette at plant base, sessile or very shortly subpetiolate,
obovate, 10-20 mm long, along stem subulate, 2-5 mm
long. Inflorescences floriferous open racemes terminating
the branches or in upper leaf axils, the pedicels much
longer than the bracts, widely spreading in fruit. Flower
resupinate, the corolla bilabiate, 6.5 mm long along
the upper (adaxial) side, purple with the throat marked
with a white longitudinal band along the lower side
and a pair of white striations on each side. Stamens 2
functional, positioned abaxially, anthers c. 0.8-1 mm
long; staminodes 2, positioned adaxially, stout, yellow.
Capsules globular, 2.5 mm long, glabrous; seeds (2 seen)
narrow obloid, 0.4-0.5 mm long, finely 4-5 angled.

Distribution. In the Prince Regent River watershed,
western Kimberley, Western Australia. On sand flats in
mixed herbfield or damp clay in closed grassland.

Note. A patch of hundreds of plants, most in full
flower, co-occurred with L. hypandra which was only
just coming into flower {M.D. Barrett 671, AD).

Etymology. The adjectival epithet is derived from the
Greek porphyro -, purple, dinos, cup, and -eus, noted
for (Stearn 1966, p. 267), alluding to the distinctively
open-mouthed purple-limbed corolla.

Selected additional specimen

WESTERN AUSTRALIA: Blyxa Creek, Prince Regent
River Reserve, 19 Aug. 1974, A. S. George 12432 (CANB,
PERTH).

C. Lindernia sect. Scapigerae W.R.Barker, sect. nov.
Typus: L. scapigera R.Br.

Annuals; stems and branches erect or narrowly
ascending to procumbent, terminated by open racemes.
Leaves subpetiolate, narrow to broadly ovate to obovate.
Pedicels erect or ascending, in fruit sometimes deflexed,
hardly elongating. Corolla regular, lacking internal
longitudinal flaps; tube conspicuous, as long or longer
than the limb; upper lip porrect or slightly upturned,
emarginate. Stamens 4, with the abaxial filaments
bearing a lateral cylindrical spur; anthers 2-celled.
Capsules ovoid to subglobular; seeds longitudinally
4-8-ribbed, with 5~9 (~?10) transverse ridges.

Distribution. From the Kimberley, Western Australia,
to north and east Queensland.

Note. The section includes L. alsinoides R.Br.,
L. pubescens (Benth.) F.Muell. and L. scapigera R.Br.
Three further species that have been confused with
L. scapigera are separated.

Key to species of Lindernia sect. Scapigerae

1. Corolla to c. 6 mm long along upper side. L. alsinoides s. lat.

1: Corolla greater than 6 mm long along upper side

2. Stems and leaves glandular hairy. L. pubescens

2: Stems and leaves glabrous

3. Corolla blue to blue-purple

4. Scandent, often multi-stemmed annual; leaves entire through undulate to coarsely serrulate
within a population; corolla blue, purple or blue-purple with white tube, rarely white, with

deeper blue-purple blotches on either side of the lowest lobe. 10. L. scapigera

4; Erect, free-standing annual; leaves entire to coarsely serrate within a population; corolla mid

blue with lower lip splashed with white. 11 . L. grossidentata

3: Corolla yellow

5. Corolla light to deepish yellow throughout, often with a few red streaks in the mouth; hood,

1.3-2.5 mm long; adaxial staminal spur 1.6-2.4 mm long .12. L. tiwiensis

5: Corolla white inside, yellow in throat extending onto lower lip, with outside of tube above

yellow to brown; hood 0.8-1.5 mm long; adaxial staminal spur 0.6-1.5 mm long .... 13. L. leucochroa

67

W.R. Barker

Swainsona 31 (2018)

10. L. scapigera R.Br.

Prodr. (1810) 441. — Vandellia scapigera (R.Br.)
Benth. in DC., Prodr. 10 (1868) 415. — Tittmannia
scapigera (R.Br.) Spreng., Syst. Veg., ed. 16, 2 (1825)
800. — Ilyogeton scapigera Benth. in DC., Prodr. 10
(1868) 415, pro syn. (this combination in Ilyogeton
Endl. did not appear in Walp., Rep. Bot. Syst. 3: 297,
as attributed by Bentham). — Type citation: “(T.)
v.v.“ [from the tropical region of Australia, seen living
in situ]. — Lectotypus (Philcox 1968: “holotype”):
R. Brown Iter Australiense 2696, lect. 20 Dec. 1802,
desc. 21 Dec. 1802, Carpentaria Islands h, cfr
Vandelioides a facing Island. / North Coast, Lindernia
gracilis. / R. Brown s.n.. Northern Territory, North
Island (island h), Sir Edward Pellew Group (BM
1040723; Fig. 4). Isolectotypi: R. Brown s.n., 20 Dec.
1802, Northern Territory, North Island (island h),
Sir Edward Pellew Group (BM 1040724); R. Brown
Iter Australiense 2696, s. loc. (“Lindernia scapigera,
Vandellia scapigera”; K 859768, photo); [R.Brown]
s.n., s.dat., Carpentaria (ex Herb. R. Brown 8/80). /
“Vandellia scapigera, Bth [sic!]” (K 859770).

Typification. The name L. scapigera has been applied
to several taxa of similar appearance across northern
Northern Territory. High quality photographs of
the types available on-line through JSTOR Global
Plants enable confirmation that the type belongs
with a species that extends from south of Darwin to
the Sir Edward Pellew Group of islands in the Gulf
of Carpentaria. This is the only species in the east of
the range of the complex (Barker, in prep.). Brown
collected and immediately described the collection in
detailed notes. The collection is represented by three
sheets (syntypes), necessitating a lectotypification. The
specimen BM 1040723 is the best collection as it has
multiple labels by Robert Brown and has several plants
in flower and fruit. This is the syntype annotated with
the species identity by Philcox (1968) for his revision
of Malesian Lindernia, in which he cited the BM
specimen as a holotype: “Carpentaria Island Dec.,
R. Brown ‘2696’ (Holotype BM, isotype K)”. The
ICN (Art. 9.9: McNeill et al. 2012) indicates Philcox’s
published designation of holotype is to be interpreted
as a lectotypification.

The syntypes in BM and K are clearly from the same
collection from the annotations and the similarity of
the specimens; they are also in good condition.

L. scapigera 2: Biffin et al.. Austral. Syst. Bot. 31 (2018), in
press.

Distribution. In Northern Territory, from south of
Darwin across northern Arnhem Land to Vanderlin
Island in the Sir Edward Pellew Group islands in the
Gulf of Carpentaria.

Selected additional specimens

NORTHERN TERRITORY: Angularli Creek, Coburg
Peninsula, 23 Aug. 2009, D.E. Murfet 6570 (AD, DNA);
Hades Flat Meteorological] Station, 1 km W of Oenpelli
road, 17 km from Arnhem Highway, 6 May 1983, R.M.
Barker 458 (AD); Gulungul Creek crossing on Arnhem
Highway, c. 600 m W of turnoff to Jabiru Airport, 16 May
1994, W.R. Barker 6881, R.M. Barker &M.C. O’Leary (AD);

Arafura Swamp, east side, c. 17 km SE of Ramingining,
10 Dec. 1998, I.D. Cowie 8129 & R.K. Harwood (AD);
5 km NNW of Lake Eames, Vanderlin Island, 24 July 1988,
P.K. Latz 10747 (AD); Near Gulbuwangay River on track to
Mirrnaja, 11 Oct. 2009, D.E. Murfet 6616 (AD; voucher for
Biffin et al., in press).

11. L grossidentata O.Schwarz

Repert. Spec. Nov. Regni Veg. 24 (1927) 96. — Type
citation: “9 mi E of Darwin, wet patch of ground:
Bleeser 186”. — Neotypus (hie designatus): W.R.
Barker 7050 & R.M. Barker, 1 June 1994, Howard
Springs Park Estate, pending subdivision; c. 400 m
down Parakeet Road from Bronzewing Road; 50 m
N of road, Northern Territory (AD 99818363).
Isoneotypi: CANB, DNA, NSW.

Typification. No type material has been located despite
a search of on-line resources (see p. 61). The species
to which the name is applied here, and to which the
neotype belongs, matches the protologue closely,
particularly in the aniseed scent and often coarsely
toothed leaves (“margine remote et acute grossidentata”;
Schwarz 1927).

L. sp. MountBundey (C.R. Dunlop 8840) [N.T. Herbarium]:
R.A.Kerrigan & Alb., Checkl. N. Territory Vase. PL (2007).

Distribution. In Northern Territory, in the vicinity of
Darwin, extending to the Fish River area.

Note. Schwarz (1927) provides a detailed description of
the species.

Selected additional specimens

NORTHERN TERRITORY: Mount Bundey Quarry,
12 Apr. 1991, Dunlop 8840 & Cowie (DNA); Charles Darwin
National Park, 6 Mar. 1998, PS. Short 4667 & C.R. Dunlop
(AD, DNA); Douglas Hot Springs road, 17.6 km from old
Stuart Highway, 29 Apr. 1983, R.M. Barker 360 (AD).

12. L. tiwiensis W.R.Barker, sp. nov.

Holotypus: I.D. Cowie 8462, 16 Feb. 2000, Melville
Island, c. 7 km SE of Garden Point, Northern Territory
(AD 279773). Isotypi: DNA, BRI, CANB, K, MEL,
NY, PERTH.

L. sp. Melville Island (C.R. Dunlop 4609) [W.R.Barker]:
R.A.Kerrigan & Alb., Checkl. N. Territory Vase. PI.
(2007), partly.

Erect to scandent, glabrous, annual herb, simple or
with up to 5 stem-like branches, these 8~40 cm long.
Leaves in spaced pairs, spathulate, 5 - 25 mm long,
entire to coarsely serrate. Lnflorescences open terminal
racemes of up to 10 flowers; pedicels much longer than
the subulate to linear bracts, 1 ~25 mm long, deflexed
in fruit. Corolla 6-6.7 mm long along the upper side,
yellow, sometimes with red streaks; upper lip a porrect
hood, 1.3~2.5 mm long. Stamens 4, perfect, with
abaxial spur narrow cylindrical, straight or distally
curved outwards and/or upwards, 1.6-2.4 mm long;
anthers 2-celled, white. Capsule narrow obloid-ovoid to

68

Swainsona 31 (2018)

Notes on the taxonomy of Australian Lindernia subg. Didymadenia

LrnUemin

PLANTS OF AUSTRALIA

COLLECTED BY ROBERT BROWN
1 KOI -1 J»0 5

Fig. 4. Lectotype of Lindernia
scapigera R.Br. at the Natural
History Museum, London (BM
001040723; image by Natural
History Museum, CC-BY 4.0).

ovoid, 3.2-4 mm long; glabrous; seeds c. 0.5 - 0.6 mm
long, with c. 3 _ 6 longitudinal ribs.

Distribution. Confined to the Tiwi Islands, Northern
Territory.

Etymology. The adjectival epithet alludes to the
restriction of this species to the Tiwi Islands.

Selected additional specimens

NORTHERN TERRITORY: Melville Island, 8 Sep. 1977,
C.R. Dunlop 4609 (AD, DNA); northern part, headwaters of
Dudwell Ck, 28 Oct. 2014, /. Cowie 13702 &N. Cuff (DNA);

Bathurst Island, 30 Apr. 1998, C. Michell 1288 & R.K.

Harwood (DNA); Melville Island, 10 km NW Pickertaramoor,

7 Jun. 1987,/ Russell-Smith 2496&D. Lucas (DNA).

13. L. leucochroa W.R.Barker, sp. nov.

Holotypus: W.R. Barker 9116, R.M. Barker & K
Brennan, 12 May 2013, c. 250 m towards Jabiru from
Nourlangie Rock turnoff on the Kakadu Highway to
Pine Creek, Northern Territory (AD 268116; voucher
for Biffin et al., in press). Isotypi: CANB, DNA,
PERTH.

69

W.R. Barker

Swainsona 31 (2018)

L. sp. Melville Island (C.R. Dunlop 4609) [W.R.Barker]:
R.A.Kerrigan & Alb., Checkl. N. Territory Vase. Pl.
(2007), partly.

L. scapigera 1: Biffin et al., Austral. Syst. Bot. 31 (2018), in
press.

Scandent to suberect, glabrous, annual herb, with stem
and main branches 12-40 cm or more long. Leaves
spathulate, distally 0.5 _ 1.8 cm long, entire to coarsely
serrulate. Inflorescence an open terminal raceme, the
pedicels reflexed in fruit, much longer than the tiny
bracts. Corolla 8-11 mm along the upper side, in front
view with the lower lip white and the throat yellow,
with the outer surfaces pale yellow, the hood sometimes
flushed brown and then sometimes with its extremities
flushed with blue. Stamens 4, perfect, with abaxial spur
narrow cylindrical, yellow; anthers 0.8-1 mm long,
white. Capsules ellipsoid to broad ovoid, 3-4.5 mm
long, glabrous; seeds obloid to ellipsoid, 0.3-0.4 mm
long, longitudinally 5 - 8-ribbed.

Distribution. From Daly River to central Arnhem
Land plateau, Northern Territory, in herbfield or
woodland.

Etymology. An adjective, formed by the elision of
the Greek adjectives leucos , white, and ochros, yellow,
alluding to white face of the flowers with conspicuous
yellow encompassing the throat and mouth and often
at the back of the limb.

Selected additional specimens

NORTHERN TERRITORY: Nigalaye Brook, 30 km WNW
of Cannon Hill Ranger Station, 30 May 1980, M. Lazarides
9099 (AD, DNA ex OSS); On Jabiru-Oenpelli road, on
jump up at NE end of crossing of Magela Creek floodplain,
23 May 1994, W.R. Barker 6991 & R.M. Barker (AD);
Margins of Flying Fox Creek, c. 50 m W of bridge, 12 May
2013, W.R. Barker 9123 & R.M. Barker (AD); Noonamah,
Jenkins Road near creek, 12 July 2008, D.E. Murfet 6060
(AD).

D. Lindernia sect. Heterandrae W.R.Barker, sect. nov.

Typus: L. thyridostoma W.R.Barker

Annuals; stems and branches erect or narrowly
ascending terminated by open scapiform racemes.
Leaves subpetiolate, broadly ovate to obovate. Pedicels
erect or ascending, in fruit sometimes deflexed. Corolla
regular, lacking internal longitudinal flaps; tube much
shorter than limb; upper and lower lips widely spread
forming an open throat. Stamens 4, with the abaxial

filaments with the lateral spur reduced to a swelling;
adaxial anthers 2-celled, abaxial with a single perfect
cell. Capsules narrow ovoid to subglobular; seeds
4-ribbed, with 6-9 transverse ridges.

Distribution. Confined to northern Northern
Territory, this section comprises the following two
species.

14. L. thyridostoma W.R.Barker, sp. nov.

Holotypus: W.R. Barker 7812, K. Brennan & R.M.
Barker, Edge of hill c. 3 km NW of Nabarlek air
strip, Northern Territory (AD 269127). Isotypi: BRI,
CANB, CNS, DNA, K, PERTH.

L. sp. Open throated (J. Russell-Smith 5581) [W.R.Barker]:
R.A.Kerrigan & Alb., Checkl. N. Territory Vase. PI.
(2007).

Erect, rarely scandent, usually rosulate, delicate annual
herb, 1.5 - 30 cm high, with a single stem or up to 15
stem-like branches, with long flexuous glandular hairs
on the leaves and in the inflorescence. Leaves spathulate,
0.5 - 10 cm long. Inflorescences usually open racemes,
rarely few-branched panicles, with pedicels much
longer than the subulate bracts. Corolla 3.5 - 4 mm long
along upper side, darkish blue with the mouth marked
by a yellow spot on the lower side, often with brown-
purple speckling on either side, and a translucent spot
on either side of the dilated rear of the tube. Capsule
ovoid, 3~4.5 mm long, glabrous or with tiny glandular
hairs; seeds 0.3-0.4 mm long, with 4 longitudinal
angles.

Distribution. Widespread across northern Northern
Territory, in caves or sheltered places on rock faces.

Etymology. The adjectival epithet derives from the
Greek, thyris, -idos, small door or window, and stoma,
throat, alluding to the pale translucent blotches on the
upper side of the corolla throat.

Selected additional specimens

NORTHERN TERRITORY: Tributary of Fitzmaurice
River, 23 Feb. 1994, G.J. Leach 4203 (AD, DNA); Litchfield
National Park south, Tableland Creek Gorge, 14 Feb. 1996,

I. D. Cowie 6167 & R. Booth (AD, DNA); Radon Springs
[Upper Baroalba Creek, Mt Brockman], 13 May 1987, B.S.
Wannan & C.J. Quinn UNSW 20271 (AD); UDP Falls,
Waterfall Creek Nature Reserve, 9 May 1983, R.M. Barker
496 (AD); Gulungul, Kakadu National Park, 23 Mar. 1995,

J. Russell-Smith 10214 (DNA); Limmen National Park, St
Vidgeons block SW corner, c. 65 km from St Vidgeons ruins,
24 Apr. 2009, D.L. Lewis 1106 (AD).

Key to species of Lindernia sect. Heterandrae

1. Corolla blue-purple and often red-spotted, with a gullet mouth, the very back dilated, distally with

parallel sides and with two translucent patches at the rear. 14. L. thyridostoma

1: Corolla mid blue through pink to white, with a widely dilated dish-like mouth with widely spread

sides . 15. L scutellata

70

Swainsona 31 (2018)

Notes on the taxonomy of Australian Lindernia subg. Didymadenia

15. L. scutellato W.R.Barker, sp. nov.

Holotypus: W.R. Barker 9108, R.M. Barker, K Brennan
& S. Ranger, 9 May 2013, c. 300 m direct NNW from
crest of road over Koongarra Saddle at head of Baroalba
Creek valley, on W side of creekline (AD 268119).
Isotypi: CNS, DNA, PERTH.

L. sp. Small whitish corolla (I.D. Cowie5661) [W.R.Barker]:
R.A.Kerrigan & Alb., Checkl. N. Territory Vase. PI.
(2007).

Erect to scandent delicate annual herb, 6~70 cm high,
with a single stem to several stem-like branches arising
from ground level, sometimes with tiny glandular hairs
in the inflorescence. Leaves often subrosulate, largest
near ground level, spathulate, 7 ~35 mm long, entire
to irregularly coarsely undulate. Inflorescence an open
terminal raceme or panicle, with pedicels much longer
than the subulate bracts. Corolla like an open saucer,
2-4 mm long along upper side, mid blue to white, with
a yellow tube and a yellow spot on the lower side of
the mouth. Capsule ovoid or narrowly so, 2.7 _ 4.3 mm
long, covered by sparse tiny glandular hairs, rarely
glabrous; seeds obloid, c. 0.4-0.5 mm long, with 4
blunt longitudinal ribs.

Distribution. Confined to the northwest and western
Arnhem Land escarpment and plateau in the Northern
Territory.

Etymology. The adjectival epithet derives from the
diminutive of scutum , shield, and scuta flat tray, platter,
scutella, a small flat dish or plate, alluding to the saucer
shaped corolla.

Selected additional specimens

NORTHERN TERRITORY: Myra Falls vicinity, Tin Camp
Creek, 29 May 1973, T.G. Hartley 73<W(CANB); c. 700 m
WSW of the East Alligator River, c. 3.4 km S of Cahill’s
Crossing, c. 23.5 km NNW of Jabiru Airstrip, 20 Apr. 1999,
W.R. Barker 7885 & I. Cowie (AD); 22 km SE of Jabiru
airstrip, Magela Creek upper catchment, 12 Apr. 1995, /.
Cowie 5661 & K. Brennan 3122 (AD, DNA); 1 km direct
S of Koongarra Saddle, 20 May 1994, W.R. Barker 6928, K.
Brennan &R.M. Barker (AD).

E. Lindernia sect. Hemiarrhena (Benth.) W.R.Barker,
comb, et stat. nov.

Hemiarrhena Benth., FI. Austral. 4 (1868) 518,

basionym. — Type: L. plantaginea (F.Muell.) F.Muell.

Annual, rarely perennial, herbs; stems and branches
erect or narrowly ascending to procumbent. Leaves
subpetiolate, narrow to broadly ovate to obovate.
Inflorescences open racemes, often scapiform, or dense
scapiform heads. Pedicels short, erect or ascending, in
fruit sometimes deflexed, hardly elongating. Corolla
regular, lacking internal longitudinal flaps; tube
conspicuous, as long or longer than the limb; upper lip
porrect or sharply reflexed upwards, emarginate or lobed.
Stamens 2, the abaxial pair absent or present as a pair of
staminodes; anthers 1-celled. Capsules narrow obloid to

subglobular; seeds 4-angled, shallowly furrowed, with c.
5 - 9 fine, sometimes obscure, transverse ridges.

Distribution. Apart from L. clausa, L. plantaginea and
L. dunlopii, which extend to northern Western Australia,
and L. mitrasaemoides, which extends more widely in
northern Northern Territory, the group is confined to
western and northern margins of the Arnhem Land
plateau, Northern Territory, where it is very diverse.

Note. The species have narrow ecologies in the overall
range of the section, which includes savannah woodland,
herbfields, skeletal soil on damp sandstone pavements,
in caves in sandstone cliffs, on top of outcrops, and, in
at least one instance, in a gorge with remnant rainforest.

16. L. mitrasaemoides (O.Schwarz) W.R.Barker, comb,
nov.

Ilysanthes mitrasaemoides O.Schwarz, Repert. Spec.
Nov. Regni Veg. 24 (1927) 96, basionym. — Lindernia
mitrasaemoides O.Schwarz, nom. inval. : Dunlop et al .,
Checkl. Vase. PL N. Terr. (1987) 74. — Type citation:
“Port Darwin, 10 miles E (Bleeser no. 206, flor.),
4 miles N (Bleeser no. 315, fruct.); bruised plant smells
strongly of aniseed.” — Lectotypus (hie designatus):
F.A.K. Bleeser 315, Apr. 1927, Darwin (NSW 50225;
Fig. 5). Isolectotypus: DNAA0030367.

Nomenclatural note. Previous instances of publication
of this combination in Lindernia are invalidly
published. After 1 Jan. 1953, a new combination
requires the basionym to be clearly indicated and
directly referenced (McNeill et al. 2012: Art 41.5).
Typification. The syntype material seen by Schwarz
is lost (see p. 61). The NSW and DNA specimens are
isosyntypes, the NSW specimen being referred to as
such (“duplicate of a syntype of Ilysanthes mitrasaemoides
O.Schwarz”) by McKee (1963). In keeping with the
citation of the syntypes in the protologue, the NSW
specimen is largely, but not entirely, in fruit, and bears
a faint number “315” written on a newspaper rectangle,
similar to the collector number labels on other Schwarz
types, e.g. those of Acacia pellita O.Schwarz, Capparis
citrifera O.Schwarz and Calophyllum ramiflorum
O.Schwarz, seen on-line (JSTOR Global Plants 2017).
Like the NSW Lindernia specimen, dated “4/1927”,
these three examples were collected in 1927 and sent
to Berlin in time for Schwarz’s publication later in the
same year. The NSW specimen is the better quality
of the two isosyntypes and therefore the choice for
lectotype. Furthermore, from copied annotations, the
DNA specimen has been segregated from the NSW
material, which was presumably amongst the many
collections sent directly by Bleeser to Blakely in Sydney
(McKee 1963).

Ilysanthes lobelioides auct. non F.Muell: Specht, Rec. Amer.-
Austral. Sci. Exped. Arnhem Land 3 (1958) 298, partly.
— L. lobelioides auct. non (F.Muell.) F.Muell.: Dunlop
et al ., Checkl. Vase. PI. N. Terr. (1995) 102.

Distribution. In the northern mainland margin of the
Northern Territory between the Darwin region and
north-west Arnhem Land.

71

W.R. Barker

Swainsona 31 (2018)

Key to species of Lindernia sect. Hemiarrhena

1. Anthers 2-celled

2. Aniseed-scented annual; leaves, bracts and sepals smooth-surfaced, the leaves undulate to coarsely

serrulate; inflorescence a moderately dense head; capsules ovoid or narrowly so.16. L. mitrasacmoides

2: Flyspray-scented annual; leaves, bracts and sepals pustulate, the leaves usually pinnatifisect to

pinnatifid, rarely coarsely serrate; capsules broad ovoid.17. L. pustulosa

1: Anthers 1-celled

3. Filaments behind the anther connective terminated by an obloid swelling. L. clausa

3: Filaments evenly narrow to the anther connective

4. Flowers in dense heads; pedicels hardly longer than subulate bracts

5. Corolla hood porrect; abaxial staminodes absent; anthers awned by the long distal attenuation

6. Perennial herb with woody rootstock; leaf blades narrow elliptic to narrow obovate, entire

to shallowly coarsely serrate.18. L. plantaginea

6: Annual, with undeveloped rootstock; leaf blades narrowly to broadly ovate to obovate,

entire to minutely sinuolate, rarely denticulate.19. L. dunlopii

5: Corolla hood with a distal upturned bilobed lip; abaxial staminodes present; adaxial anthers

acute distally.20. L. brennanii

4: Flowers in open racemes; pedicels longer than subulate bracts

7. Corolla hood well-developed, coloured; anthers included in tube, under hood or against the

front of the hood
8. Corolla hood with recurved lip
9. Upper corolla lip glabrous

10. Perennial, with long scandent branches. 21. L. lucrusmiana

10: Annual, with erect branches.22. L. venustula

9: Upper corolla lip eglandular hairy.23. L. enypniastina

8: Corolla hood porrect; anthers under tip of hood, distally awned.24. L. acrandra

7: Corolla hood reduced, hardly coloured; anthers exposed, yellow, projected well in front of
corolla hood

11. Anthers horizontal, in line with the filament, with dehiscence slits facing lower side of
mouth

12. Anthers 1.6-1.8 mm long .25. L. pronanthera

12: Anthers 0.6-1 mm long .26. L. robyniae

11: Anthers vertical, at right angles to the filament, with dehiscence slits facing forward;

anthers 1.1 -1.35 mm long .27. L. calliandra

Note. Schwarz (1927) provides a detailed description of
the species.

Selected additional specimens

NORTHERN TERRITORY: Charles Darwin National
Park, 4 Sep. 1998, P.S. Short 4750 & C.R. Dunlop (AD);
Howard Springs Park Estate, pending subdivision, c. 400 m
down Parakeet Road from Bronzewing Road, 50 m N of
road, 1 June 1994, W.R. Barker 7051 & R.M. Barker (AD);
12.5 km S of Cannon Hill Ranger Station, 28 May 1980,
L.A. Craven 6018 (AD); c. 9 km by road towards Jabiru from
Nourlangie Rock turnoff on the Kakadu Highway from Pine
Creek, 12 May 2013, W.R. Barker 9117, R.M. Barker & K.
Brennan (AD); 19 km NNW of Twin Falls, 3 June 1980,
L.A. Craven 6298 (DNA).

17. L pustulosa W.R.Barker, sp. nov.

Holotypus: W.R. Barker 7833, R.M. Barker &
K Brennan, 4 Apr. 1999. Beside vehicular track,
c. 1.0 km direct S of Koongarra Saddle, Northern
Territory (AD 268130). Isotypi: CNS, CANB, DNA,
PERTH, MEL.

L. sp. Dissected leaf (N.B. Byrnes 1520) [W.R.Barker]:
R.A.Kerrigan & Alb., Checkl. N. Territory Vase. PI.
(2007).

Robust, erect, often branched, annual 12-35 cm high,
with the scent of flyspray, stem often branched from
the base, with short glandular hairs in the inflorescence,
the surface of the leaf segments, bracts and sepals
prominently pustulate. Leaves pinnatisect or pinnatifid
to lobulate, 3~16 mm long. Inflorescences open terminal
floriferous racemes, the pedicels 5 - 15 mm long, much
exceeding the narrow deltoid to linear bracts. Corolla
10-12 mm long along the upper side, pale pink to blue,

72

Swainsona 31 (2018)

Notes on the taxonomy of Australian Lindernia subg. Didymodenia

B

A

NSW

LOAN:

Fig. 5. Lectotype of llysanthes mitrasacmoides O.Schwartz. A Herbarium sheet from the National Herbarium of New South Wales
(NSW 50225), seedpacket folded upwards and not shown; the label at the bottom left of the lectotype has a faint, largely erased
number not discordant with "315". B Label in packet with annotation by H.S. McKee. C Contents of packet. (Reproduced with
permission of the Royal Botanic Gardens Sydney).

with long tube and closed mouth. Stamens: the adaxial
pair perfect, with anthers c. 0.7 mm long, 2-celled;
the abaxial staminodes terminated by a reduced empty
anther, with a blue or yellow linear oblong spur. Capsules
broad ovoid, 3 _ 3.8 mm long; seeds 0.25 - 0.4 mm long,
mid brown, sharply 4-angled.

Distribution. Known from northern and western
margins of the Arnhem Land plateau, Northern Territory,
on exposed sandstone rock sheets prone to inundation.

Etymology. The epithet is adjectival, deriving from
the Latin pustula, blister, bubble, and the suffix -osus,

denoting abundance, alluding to the heavily blistered
leaf segments, bracts and sepals.

Selected additional specimens

NORTHERN TERRITORY: Spencer Range, 43 km E of
Oenpelli Mission, 17 Feb. 1973, L.G. Adams 3014 (CANB);
Edge of hill c. 3 km NW of Nabarlek air strip, 3 Apr. 1999,
W.R. Barker 7815, K. Brennan & R.M. Barker (AD); Mt
Brockman, 26 Mar. 1995,/. Egan 4474 (DNA); 17 km SE of
Twin Falls, 20 Mar. 1988, R. Fensham 886 (DNA); Waterfall
Creek, 1 m[ile] above falls, 2 Apr. 1969, N. Byrnes 1520
(CANB, DNA).

73

W.R. Barker

Swainsona 31 (2018)

18. L plantaginea (F.Muell.) F.Muell.

Fragm. 6 (1867) 102 (at least as to the type “ad
flumen Victoriae F.M.”); R.A.Kerrigan & Alb.,
Checkl. N. Territory Vase. PL (2007), partly. —
Vandellia plantaginea F.Muell., Trans. Philos. Inst.
Victoria 3 (1859) 62 (“Sect. Bonnaya”), basionym. —
Hemiarrhenaplantaginea (F.Muell.) Benth., FI. Austral.
4 (1868) 518, partly (as to Mueller and Marten
collections). — Type citation: “In moist meadows near
Macadam Range”. — Lectotypus (hie designatus):
Dr M[uell.] s.n., s.dat., Bonnaya ? plantaginea n.sp.
/ A rare plant, [moist] but ... between McAdam
Range & Providence Hill (K 859761, Herb. Hooker;
Fig. 6a). Isolectotypus possibilis: Ferd. Mueller,
Between Providence Hill & McAdam Range. / Ferd.
Mueller, Sept [18] 55, Ad ostium fluminis Victoriae /
FM, 1855, Arnhem’s Land. Hemiarrhena plantaginea
Benth. (MEL 1552813; Fig. 6b).

Typification. In the protologue, Mueller indicates:
“This species is extremely rare, and the only flowering
specimen which was found is deposited in Sir Wm.
Hooker’s herbarium, at Kew.” A possible syntype MEL
1552813, identifiable by one of two more specific
locality labels, has a single individual plant. It is in poor
condition by comparison with the two K plants on the
lectotype sheet; it lacks flowers and has apparently been
detached from its root base. A second label referring
to the mouth of the Victoria River appears to be
discordant; the type location between Providence Hill
and the Macadam Range, appearing on the first label,
is some distance to the north. Rather than the Victoria
River reference being a general regional statement, it is
possible that it signifies that the MEL material is from
a second distant collection.

Another collection ( F. Mueller s.n., 10 Jul. 1856,
Elsey’s Creek, MEL 2255630), possibly of L. aplectra
W.R.Barker, is annotated Hemiarrhena plantaginea by
Mueller. However, as it is not annotated with the name
“Bonnaya plantaginea” or mentioned in the protologue
by citation of its locality, it is not considered a syntype.

L. sp. Long-leaved(J.L. Egan 5112) [W.R.Barker]: R.A.Kerri¬
gan & Alb., Checkl. N. Territory Vase. PL (2007).

Note. L. plantaginea differs from the following species,
with which it has been confused to this time, by its
perennial rootstock, alluded to in the protologue and
early publications, and its consistently narrow leaves.

Selected additional specimens

WESTERN AUSTRALIA: Blyxa Creek, Prince Regent
River Reserve, 19 Aug. 1974, A.S. George 12419 (PERTH);
Drysdale River crossing, 25 km SE of Carson River Station,
1 July 1997, K.F. Kenneally 11880 (PERTH).

NORTHERN TERRITORY: Keep River National Park, SW
of Jarrnarm, 19 May 1995,/. Egan 5112 (AD, DNA).

19. L. dunlopii W.R.Barker, sp. nov.

Holotypus: R.M. Barker 468 & C. Dunlop, 7 May
1983, Little Nourlangie Rock, 12.2 km from Jim Jim
road, turnoff 21 km from Arnhem Highway, Northern
Territory (AD 98504060). Isotypi: AD 98504061,
CANB, CNS, DNA, K, MEL, NSW, PERTH.

L. plantaginea auett. non (F.Muell) F.Muell.: R.A.Kerrigan
& Alb., Checkl. N. Territory Vase. Pl. (2007), partly;
Biffin et al., Austral. Syst. Bot. 31 (2018), in press,
partly.

Erect, scented, rosulate, annual, 10-62 cm high,
with non-woody rootstock, glandular hairy in upper
parts. Leaves spathulate, to 4.3 cm long, entire to very
shallowly sinuolate. Inflorescence a dense floriferous
terminal head, sometimes with secondary inflorescences
from upper stem nodes, with bracts narrow elliptic-ovate
to -obovate, 1-2.5 mm long, longer than the pedicels.
Corolla 7.8-10.5 mm long along the upper side, pale
blue; hood porrect, auriculate, attenuating into entire
obtuse apex. Stamens an adaxial pair, the abaxial pair
absent, slightly exserted; anthers distorted-obovoid,
0.5“ 1 mm long, mid to deep blue, 1-celled, awned.
Capsule broad ovoid to almost globular, 2.5~3 mm long,
covered by blister glands and dense glandular hairs;
seeds obloid-ellipsoid, c. 0.3-0.35 mm long, 4-angled.

Distribution. Widespread from the Kimberley, Western
Australia, to Arnhem Land, Northern Territory, in
herbfield and woodland understorey.

Etymology. Named in recognition of Clyde Dunlop’s
major contribution to floristic knowledge of northern
Australia and to the infrastructure and community
understanding of plant systematics in the Northern
Territory.

Selected additional specimens

WESTERN AUSTRALIA: 2.3 km by road SE of Mt
Elizabeth Homestead, on southward vehicular track, 17
May 2000, W.R. Barker 8027 & R.M Barker (AD); Outside
mess hut, Amax Capsite, Mitchell Plateau, 26 June 1976,
KF. Kenneally 5356 (PERTH); Above the E bank of Wonga
Creek c. 25 km W of Kalumburu Mission, 31 May 1996,
A.A. Mitchell4366 (AD).

NORTHERN TERRITORY: Litchfield National Park,
c. 750 m direct E of car park at north end of Lost City,
on vehicular track above first jump-up, 19 May 2013,
W.R. Barker 9127 & K. Brennan (AD; voucher for Biffin et
al., in press); 1 km direct S of Koongarra Saddle, 20 May
1994, W.R. Barker 6930, K. Brennan & R.M. Barker (AD);
Katherine Gorge National Park, 21 June 1975, C. Dunlop
3775 (AD).

20. L. brennanii W.R.Barker, sp. nov.

Holotypus: W.R. Barker 6936, K. Brennan & M.
O’Leary, 21 May 1994, On Arnhem Land plateau,
c. 32 km ESE of Jabiru, Northern Territory (AD
99843024). Isotypi: CANB, CNS, DNA, MEL,
NSW.

L. sp. Cleft hood (C.S. Robinson 686) [WR.Barker]: R.A.Kerri-
gan & Alb., Checkl. N. Territory Vase. Pl. (2007).

Erect rosulate, scapose annual, 17 - 40 cm high, usually
simple, sometimes with up to 14 erect stem-like
branches, sometimes branched at upper nodes, perceived
by some as scented, minutely glandular hairy in upper

74

75

Fig. 6. Vandellia plantaginea F.Muell. A Lectotype at the Kew Herbarium (K 859761; copyright of the Board of Trustees of the Royal Botanic Gardens,
Kew). B Possible syntype at the National Herbarium of Victoria (MEL 1552813; reproduced with permission from the Royal Botanic Gardens Victoria).

W.R. Barker

Swainsona 31 (2018)

parts. Leaves spathulate, 0.8-4.5 cm long, entire to
irregularly shallowly coarsely sinuate. Inflorescences
dense floriferous heads, with bracts deltoid, 2-2.5 mm
long, longer than the pedicels. Corolla 8-12 mm long
along the upper side (to the point of recurvature of the
hood), blue, the hood distally recurved into a deeply
emarginate, 2-lobed lip. Stamens in 2 pairs; adaxial pair
fertile, with anthers ellipsoid-obloid, 0.7 - l mm long,
blackish-purple, 1-celled, with obtuse to subacute, non-
aristate ends; abaxial pair reduced to a pair of oblong
to linear spurs. Capsule obovoid to ellipsoid or broadly
so, 2.2~3 mm long, covered by tiny glandular hairs
c. 0.02 mm long; seeds obloid, 0.4-0.5 mm long, with

4 sharp to blunt longitudinal ridges.

Distribution. Northern Territory, confined to
the Arnhem Land plateau or north and west on
surrounding watershed plains, in open, seasonally
inundated situations.

Etymology. Named after Kym Brennan, long-time
field botanist in northern Northern Territory, whose
knowledge of the flora is held by few others, for his
guidance in seeking out the diversity of Lindernia, and
for his invaluable floral photographs.

Selected additional specimens

NORTHERN TERRITORY: On E side of Murgenella Creek
crossing, on Oenpelli road, 26 May 1994, W.R. Barker 7019,
R.M. Barker 826 &M. O’Leary (AD); Rum Bottle Creek, 28
May 1972, D.E. Symon 7936 (AD); Edge of hill c. 3 km NW
of Nabarlek air strip, 3 Apr. 1999, W.R. Barker 7816, R.M.
Barker & K. Brennan (AD); At creek crossing c. 1 km direct

5 of Koongarra Saddle, 20 May 1994, W.R. Barker 6929, K.
Brennan & R.M. Barker (AD); Site 70, 10 km N of Twin
Falls, 28 May 1980, L.A. Craven 6056 (AD); above Gunlom
Falls, 11 Apr. 1993, K. Brennan 2219 (DNA ex OSS).

21. L. lucrusmiana W.R.Barker, sp. nov.

Holotypus: D. Lucas 118, 16 June 1988, Steep
boulder gorge, Kakadu, Northern Territory (DNA
43216). Isotypus: BRI (n.v.).

L. sp. Robust branched (D. Lucas 118) [W.R.Barker]:
R.A.Kerrigan & Alb., Checkl. N. Territory Vase. PI.
(2007).

Procumbent to erect, glabrous, perennial herb, with
main branches 20-50 cm or more long, rooting at
lowest nodes. Leaves spathulate, 1.5~7 cm long, entire
to coarsely serrulate. Inflorescence a simple erect terminal
raceme of a few flowers; pedicels at lower nodes in fruit
8-10 mm long, variously spreading in fruit. Corolla
8.5“9.5 mm long along the upper side, purple or bluish
purple, with a bi-lobed recurved upper lip, greatly
exceeded by an elongated decurved lower lip 7“ 8 mm
long. Stamens: adaxial pair perfect, the abaxial missing;
anthers ellipsoid, c. 1 mm long, 1-celled, borne under
upper corolla lip. Capsules narrow ellipsoid, c. 4.5 mm
long; seeds ellipsoid-obloid, 0.45“0.5 mm long, 4
angled.

Distribution. On the north-western escarpment of the
Arnhem Land plateau, Northern Territory. Known from
two collections only from the same general location.

Etymology. The name concatenates the initial letters
of the surnames of Jeremy Russell-Smith and Di Lucas,
long-time Arnhem Land ecologists, in recognition of
their continued substantial contribution to knowledge
of the biodiversity of the region. They are the plants
only collectors and are ecological authorities on its
rainforest habitat (e.g. Russell-Smith et al. 1993).

Additional specimen examined

12 km E of Mudginberri Homestead, 7 Jan. 1991, Russell-
Smith 8387 & Lucas (DNA, MEL n.v.).

22. L. venustula W.R.Barker, sp. nov.

Holotypus: K. Brennan 3131, 14 Apr. 1995, Jabiru
Dreaming, East Alligator River, Northern Territory
(AD 99530166). Isotypi: DNA, CANB, CNS, NSW,
PERTH, K.

L. sp. Recurved hood (I.D. Cowie 1171) [WR.Barker]:
R.A.Kerrigan & Alb., Checkl. N. Territory Vase. PI. (2007).

Erect or decumbent annual, 3~40 cm tall, with stem
usually branched, rarely simple, glabrous but for minute
antrorse eglandular hairs lining the sepals. Leaves
spathulate, 0.3~6 cm long, the longest entire to coarsely
serrulate. Inflorescences usually simple, sometimes
branched open racemes, with pedicels in fruit spreading
to greatly deflexed. Corolla 2-10 mm long along
the upper side, mid blue, rarely white or pink; upper
lip recurved into two obtuse lobes divided by deep
cleft. Stamens 2 in adaxial position, with no abaxial
pair; anthers 0.4-0.95 mm long, usually mid blue,
sometimes brown. Capsule cylindrical, 3.5 - 6.5 mm
long, glabrous; seeds 0.3-0.35 mm long, with 4 weak
longitudinal angles.

Distribution. Confined to the western escarpment and
plateau of Arnhem Land, Northern Territory, in skeletal
soil associated with rock faces, at their base, in caves, or
on their summit.

Etymology. The adjectival epithet derives from the
Latin venusta , Venus-like, elegant, graceful, and the
diminutive -ulus, indicating a tendency, alluding to
the curvaceous flowers noticeable in the type and other
longer limbed variants of this species.

Selected additional specimens

NORTHERN TERRITORY: Ubirr Aboriginal Art Site, c. 30
m E of Main Gallery, c. 250 m N of car park, 19 May 1997,
W.R. Barker 7717, R.M. Barker &J.A. Barker (AD); c. 5.5 km
E of Mt Howship, c. 27 km SSW of Nabarlek, 3 Apr. 1999,
W.R. Barker 7817 & R.M. Barker (AD); Headwaters of East
Alligator River, 13 May 1997, G. Leach 4605 (AD); N facing
wall in central part of Mt Brockman, 23 Feb. 1973, L.A.
Craven 2360 (DNA); Upper Baroalba Creek, 17 Apr. 1995,
K. Brennan 3143 (AD); near Kurundie Creek, 20 Apr. 1990,
/. Cowie 1167, 1171 & G. Leach (AD, DNA).

76

Swainsona 31 (2018)

Notes on the taxonomy of Australian Lindernia subg. Didymadenia

23. L. enypniastina W.R.Barker, sp. nov.

Holotypus: K Brennan 3182, 26 Apr. 1995, Narridj
Creek area, Northern Territory (AD 99528336).
Isotypi: BRI, CANB, CNS, DNA, K, MEL, NSW,
PERTH.

L. sp. Narridj Creek (R.K. Harwood 981) [W.R.Barker]:
R.A.Kerrigan & Alb., Checkl. N. Territory Vase. PI.
(2007).

Much-branched, delicate, glabrous or eglandular
hairy annual, usually forming leafy clumps, rarely an
unbranched stem, 8-30 cm high. Leaves spathulate,
8-40 mm long, coarsely serrulate, rarely ± entire.
Inflorescence a terminal raceme of up to 15 flowers;
pedicels 2~5.5 mm long, deflexed in fruit. Corolla deep
blue purple, c. 9 mm long along the upper side; hood
with 2 upturned rounded lobes; lower lip decurved,
5~6 mm long. Stamens the adaxial pair only; anthers
obovoid, 1 mm long, 1-celled, positioned in front of tip
of corolla hood, acute at the lower end. Capsules narrow
ellipsoid, 6-6.5 mm long; seeds obloid-ellipsoid,
0.3-0.4 mm long, with 4 rounded longitudinal angles.

Distribution. Known from two sites on the north
Arnhem Land escarpment, Northern Territory, in the
north of Jabiru, and about 150 km to the west in the
Cadell River watershed.

Etymology. An adjectival epithet from the Greek,
enypniastes, m., a dreamer, and the suffix -inus,
indicating possession, in recognition of the first
Australians and their continuous spiritual ties to the
land and its flora encompassing at least 65,000 years,
evidence for which has been recently firmly established
with an archaeological site found on the same plateau
outlier as the type locality (Clarkson et al. 2017).

Selected additional specimens

NORTHERN TERRITORY: Foot of small gap high on
outlier W of main escarpment, c. 250 m S of Narradj Creek
plain, 8 May 2013, W.R. Barker 9096, K. Brennan, Ambrose
Djandjul &R.M. Barker (AD); c. 2 km NW of Korlobidahda,
Plot 2358, 13 Apr. 2000, R.K Harwood981 (AD, DNA).

24. L acrandra W.R.Barker, sp. nov.

Holotypus: W.R. Barker 7810, K. Brennan & R.M.
Barker, 3 Apr. 1999, Edge of hill c. 3 km NW of
Nabarlek air strip, Northern Territory (AD 279774).
Isotypi: BM, CANB, CNS, DNA, MEL, NSW.

L. sp. Nabarlek (K. Brennan 3156) [W.R.Barker]: RA.Kerri-
gan & Alb., Checkl. N. Territory Vase. PL (2007).

Delicate, erect, eglandular hairy, annual herb, 6~40 cm
high. Leaves spathulate, 0.7 - 8.5 cm long, shallowly
coarsely serrate. Inflorescence a terminal floriferous
raceme; pedicels in fruit usually reflexed to spreading
down, sometimes upward spreading, 5 ~7 mm long.
Corolla 9-12.7 mm along the upper side, deep blue
purple, with the hood porrect, the lower lip broad,
flat, spreading, 6.5 - 8 mm long. Stamens 2, adaxial,
projected in front of the corolla hood; anthers joined,
1-celled, obloid, 1.4-1.7 mm long, white or pale

yellow. Capsule narrow obloid-ovoid, 5-7 mm long,
sparsely eglandular pubescent, obloid to ellipsoid, often
broadly so, 0.25“0.3 mm long, almost round in cross
section, 4 angled, the cross-walling obscure.

Distribution. Known from north-west Arnhem Land
in Northern Territory.

Etymology. An adjectival epithet, from the Greek aero-,
at the apex, and andros, androecium, alluding to the
anthers projected well in front of the corolla hood.

Selected additional specimens

NORTHERN TERRITORY: Nabarlek, 8 Apr. 1989, R Hinze
514 (DNA); c. 3 km SE of Mt Howship, c. 28 km SSW of
Nabarlek, 3 Apr. 1999, W.R. Barker 7822 (AD).

25. L. pronanthera W.R.Barker, sp. nov.

Holotypus: W.R. Barker 7814, K. Brennan & R.M.
Barker, 3 Apr. 1999, Edge of hill c. 3 km NW of
Nabarlek air strip, Northern Territory (AD 268131).
Isotypi: BRI, CANB, CNS, DNA, K, MEL, NSW,
PERTH.

L. sp. Brennans showy anthers (K. Brennan 3155) [WRBarker]:
RA.Kerrigan & Alb., Checkl. N. Territory Vase. PI.
(2007).

Floriferous, fragile to robust, erect, rarely scandent,
eglandular hairy, annual herb 8~35 cm high. Leaves
spathulate, the upper or penultimate pair largest,
1.5“3.5 cm long, coarsely serrulate to serrate in the
distal Vi- 2 /?,. Inflorescences terminal, open, floriferous
racemes or panicles; pedicels 6-15 mm long, in fruit
probably hardly elongated, narrowly spreading upward
to almost vertically deflexed. Corolla 1.8-2 mm long
along the upper side as far as the tip of the mid yellow
tube; hood more or less absent; lower lip 4.5“5 mm
long, mid yellow proximally, distally blue-purple.
Stamens glabrous: abaxial pair absent; adaxial pair
with anthers single-celled, ellipsoid, 1.6-1.8 mm
long, mid yellow, directed forwards. Capsules narrow
ellipsoid, 4-5 mm long, glabrous; seeds (Barker 7814
et al.) oblong to ellipsoid, 0.3-0.35 mm long, with 4
longitudinal angles, shallow furrows between traversed
by c. 9 fine ridges.

Distribution. Confined to north-west Arnhem Land in
Northern Territory.

Etymology. The epithet is an adjective derived from the
elision of the Latin adjective pronus, inclined forward,
facing downward, and noun, anthera, in epithets also
used adjectivally, alluding to the disposition of the
prominent anthers.

Selected additional specimen

NORTHERN TERRITORY: Kakadu National Park,
northern outliers S of Fire Plot 143, 12 July 2008, K.
Brennan 7751 (DNA).

77

W.R. Barker

Swainsona 31 (2018)

26. L. robyniae W.R.Barker, sp. nov.

Holotypus: W.R. Barker 7859 & K. Brennan, 14
Apr. 1999, Cliffs below Mary River Ranger Station,
Northern Territory (AD 268126). Isotypi: CANB,
CNS, DNA, NSW, PERTH.

L. sp. Robyns showy anthers (R.M. Barker 497) WRBarker,
ined.

L. sp. Robyns showy anthers (L.A. Craven 6234) R. A. Kerrigan
& Alb., Checkl. N. Territory Vase. PL (2007).

Erect to narrowly spreading, simple or branched,
glabrous annual, 4.5“35 cm high. Leaves spathulate,
0.3-4.5 cm long; entire through sinuolate to irregularly
coarsely dentate. Inflorescences open racemes, with
pedicels deflexed to broadly downwardly spreading
in fruit, much longer than the narrow-linear minute
bracts. Corolla 2-3 mm long along the upper side,
blue throughout, with yellow patch at base of lower lip
behind anthers; lower lip spreading 2-4.8 mm long.
Stamens 2, in the adaxial position, with abaxial pair
absent; anthers projected well in front of hood, almost
horizontal, ellipsoid-ovoid, 0.9 _ 1.0 mm long, yellow,
1-celled. Capsule narrow ovoid-obloid, 3.8-7 mm long;
seeds obloid to ellipsoid, 0.35 _ 0.45 mm long, with 4
longitudinal angles.

Distribution. On south-western Arnhem Land
escarpment, Northern Territory, and Mount Douglas
not far to the west, in skeletal soil in rocky areas.

Note. While generally correctly applying the phrase
name to this species using my manuscript keys, the
Northern Territory Herbarium chose as the voucher
in their collection a specimen of the following species
{L. calliandra ).

Etymology. The epithet is a Latin noun in the possessive
case, in recognition of Robyn Barker, who has been
a participant in my investigations into Australian
Lindernia , having reported on its remarkable diversity
before my own experiences.

Selected additional specimens

NORTHERN TERRITORY: Mt Douglas, W side, 6 July
1999, I.D. Cowie & R.K Harwood 8358 (AD); Ikoymarrwa
Lookout, Kakadu National Park, 21 Mar. 2011, D.E. Murfet
7191 & A. Lowrie (AD); c. 400 m upstream from Ferny
Gully picnic area (at end of entrance track), 15 Apr. 1999,
W.R. Barker 7865 (AD); UDP Falls, 108 km NE of Pine
Creek, 9 May 1983, R.M. Barker 497 (AD).

27. L calliandra W.R.Barker, sp. nov.

Holotypus: K. Brennan 3139, 14 Apr. 1995, Little
Nourlangie Rock, Northern Territory (AD 99528344).
Isotypi: CNS, DNA, PERTH.

L. sp. Showy anthers (L.A. Craven 2393) [W.R.Barker]:
R.A.Kerrigan & Alb., Checkl. N. Territory Vase. PI.
(2007).

Erect to decumbent or scandent, glabrous herb,
3.5-36 cm high. Leaves spathulate, 0.4-6.5 cm long,
entire to serrate. Inflorescences open, terminal, rarely

subpaniculate racemes, with pedicels in fruit spreading
widely to strongly deflexed, much longer than the
narrow linear-subulate bracts. Corolla 1.5“2.5 mm long
along the upper side, yellow proximally on the tube
and upper lip, which is blue-purple distally. Stamens:
the abaxial pair absent; the adaxial pair with anthers in
front of and almost at right angles to the corolla tube,
facing forward, ellipsoid, 1.1-1.3 (-1.35) mm long,
1-celled, mid yellow. Capsule narrow ovoid-obloid
glabrous 3~8 mm long; seeds obloid or irregularly so,
0.3-0.4 mm long, with 4 longitudinal angles.

Distribution. Confined to the west margins of the
Arnhem Land plateau, Northern Territory, associated
with rock faces and boulders, in sand, often in shade.

Etymology. The adjectival epithet derives from
the Greek prefix colli-, beautiful, and andros, male,
masculine, alluding to the showy anthers.

Selected additional specimens

NORTHERN TERRITORY: Site 64, 14.5 km NE of Jabiru
East, 26 May 1980, L.A. Craven 5951 (DNA); in small gorge
c. 0.5 km SW of summit of Little Nourlangie Rock, on
low sloping rock sheet, c. 50 m NW of edge of rock where
vehicular track turns to Anbangbang Billabong, 20 May
1997, W.R. Barker 7719, 7719A, R.M. Barker & J.A. Barker
(AD); Magela Creek upper catchment, 11 Apr. 1995, /. Cowie
5599&K. Brennan (AD, DNA); Twin Falls, c. 72 km SSW of
Jabiru, Site 86, 1 June 1980, L.A. Craven 6234 (AD, DNA).

Acknowledgements

In relation to typifications in this paper I gratefully
acknowledge the assistance of Pina Milne, Wayne
Geberts and Taryn Ellis (MEL) for arranging high
quality photographs of Muellerian types, of Neville
Walsh for the examination of flowers on type material
of L. lobelioides, of Richard Jobson and Shelley James
(NSW) for arranging examination of the type of
L. mitrasaemoides and gaining improved material of
L. prolata, of Chelsea Novice (AD) for photography
of the L. mitrasaemoides type, of Matt Barrett and
Julia Percy-Bower (PERTH) for a higher quality type
of L. porphyrodinea, and of Carolyn Ricci (AD) for
assistance with anatomical studies. Russell Barrett, an
unnamed referee, Jurgen Kellermann and Robyn Barker
are thanked for reviewing the manuscript, together
with John McNeill who advised on the nomenclatural
options for the establishment of a homotypic section
based on subg. Didymadenia.

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With the exception of images and other material protected by a trademark and subject to review by the Government of
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Swainsona 31:81-91 (2018)

© 2018 Board of the Botanic Gardens & State Herbarium (Adelaide, South Australia) ©0

Antrelloides atroceracea, a new genus and species in the Pezizaceae
(Pezizales) from Australia

Pamela S. Catcheside ab & David E. A. Catcheside b

a State Herbarium of South Australia, GPO Box 1047, Adelaide, South Australia 5001
Email: Pam.Catcheside@sa.gov.au

b College of Science and Engineering, Flinders University, PO Box 2100, Adelaide, South Australia 5001
Email: David.Catcheside@flinders.edu.au

Abstract: A new genus, Antrelloides, and a new species, Antrelloides atroceracea P.S.Catches. &
D.E.A.Catches. (Ascomycota, Pezizales, Pezizaceae) from Australia are described and illustrated.
Phylogeny within the Pezizaceae is discussed and an overview of exothecial, cleistothecial and
apothecial members of the Pezizaceae given. Morphological and phylogenetic comparisons are made
within the Pezizales.

Keywords: Fungi, Pezizaceae, Pezizales, Antrelloides

Introduction

A curious Ascomycete, first found in 2007, appears
as small cushion-like black lumps (Figs 1A & IB)
amongst laterite nodules alongside and on sandy
tracks of Flinders Chase National Park, Kangaroo
Island (Fig. IE). It has also been collected in grey sand
on a track between Denmark and Walpole, Western
Australia. The stipe is buried in the soil and, on
excavation, the whole fruit body is seen to be turbinate,
the obconic base capped by a thin, black, convex disc.
In section, the stipe consists of irregular, knobbly
columns radiating upwards and outwards from the base.
The ascoma resembles a miniature cavern or grotto of
continuous columns of stalactites and stalagmites, but
gathered at the base and capped by the hymenium
(Figs 1C & ID). When first observed, the black lumps
looked like small pieces of discarded patent black shoe
leather, hence the fungus was given the informal name
‘Shiny Black Shoe Leather. The hymenial surface
of later collections was dull, so the tag name became
‘Black Shoe Leather’—an appellation we still use, as it
is helpful when describing to other potential collectors
what to look for. However, the texture is firm and waxy
but brittle, certainly not leathery nor, obviously, has it
anything to do with shoes.

flinders Chase National Park covers an area of
approximately 325 km 2 of largely undisturbed remnant
vegetation. Its habitats include eucalypt woodland,
sandy heath, rugged coastal scrub and dunes. Since we
have been surveying in Flinders Chase we have found
rare and under-collected fungi such as Amanita arenaria
(O.K.Mill. & E.Horak) Justo (Catcheside & Catcheside
2005) and the species described in this paper, both of
which have also been found in Western Australia.

, Kangaroo Island, Australia

Materials and methods

Habitat and associated plant communities were noted
in the field. Collection locations were recorded by GPS,
geodetic datum WGS84/GDA94 (Garmin GPS 12)
and in situ and section photographs were taken with
a Nikon E4500 or, for Pig. ID, a Lumix DMC-GX7
and for Pig. IE, the habitat shot, an Olympus TG2.
Macroscopic characters were described directly from
fresh material. Colours are designated using the Royal
Botanic Gardens Edinburgh Colour Chart (1969),
given as colour descriptor and number, e.g. violaceous
black 38, and in general terms. For the more variable
colouration in the stipe tissue, colours are given
according to 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 1000FD).

Sections of fresh material and dried specimens were
hand-cut and mounted in various media. For the
amyloid reaction, fresh material was stained with
Melzer’s reagent and dried material was rehydrated in
5% NH 4 OH before staining. Water mounts were used
to determine colour of context.

Measurements were made using an Olympus BH-2
microscope at 400x or lOOOx with a calibrated ocular
micrometer. Spore dimensions are given as: length
range x width range (n = 40) and Q ratio (spore length/
spore width). Dimensions of asci are given as length
range x width range (n = 20). A Nikon 4500 camera
was used to photograph microscopic characters. For
scanning electron microscopy (SEM) a small piece
of hymenial tissue was immersed in 2.5% KOH for
3 mins and rinsed in demineralised water to release

Published online: 28 Sep. 2018* flora.sa.gov.au/swainsona

ISSN 2206-1649 (Print) • ISSN 2206-1657 (Online)

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

Swainsona 31 (2018)

spores. A drop of the resulting material was mounted
on aluminium stubs with double-sided tape, dried
and then sputter-coated with platinum at Adelaide
Microscopy. Specimens were viewed under 1 OkV
in a JEOL Neoscope JCM 5000 SEM at the State
Herbarium of South Australia.

Descriptions of Antrelloides atroceracea are based on
the type collection, P.S. Catcheside PSC 2710 ( AD -
C 55811), unless otherwise stated, with outlying
measurements for other collections given in brackets.
Photographs of fruit bodies and microscopic characters
are from the type collection, unless stated otherwise
in the figure captions. All South Australian collections
have been accessioned into the State Herbarium of
South Australia (AD). AD numbers (AD-C #####)
are given in the Taxonomy section together with the
Collector’s number (PSC ####); in other sections only
the Collector’s number is used.

DNA extraction, amplification and processing were
as described in Catcheside et al. (2016); primers ITS1
and LR5 were used for amplification and ITS1, ITS4,
LR0R, and LR5 for sequencing (White et al. 1990).
Sequences were manipulated with the Geneious 8.1.9
and 11.0.3 suites of programmes using MUSCLE for
alignment and MrBayes 3.2.6 and RAxML 8.2.11
plug-ins for tree building.

Antrelloides atroceracea 28S sequences were aligned
with the top 100 blastn hits from GenBank on 7 Feb.
2017, and also 28S sequences from Boudiera dennisii
Dissing & Sivertsen, Cazia flexiascus Trappe (Trappe
1989), Eremiomyces echinulatus (Trappe & Marasas)
Trappe & Kagan-Zur, Pachyella adnata (Berk. &
M.A.Curtis) Pfister, P. hahingtonii (Berk. & Broome)
Boud., P. habrospora Pfister, P. punctispora Pfister,
P. violaceonigra (Rehm) Pfister, Ruhlandiella peregrina
Lantieri & Pfister and Sphaerosoma trispora McLennan
& Cookson. Redundant sequences were removed
from the alignment and a phylogram constructed,
rooted to Ascobolus carbonarius P.Karst. MrBayes used
the HKY85 substitution model, 4 heated chains at a
temperature of 0.2 for 1,100,000 iterations including
a burn-in of 100,000 that was discarded. Following
burn-in, trees were sampled every 200 iterations. The
average standard deviation of split frequencies reached
0.011. RAxML used the GTR GAMMA nucleotide
model with rapid bootstrapping for 1000 iterations and
a search for the best scoring maximum likelihood tree.

Taxonomy

Antrelloides P.S.Catches. & D.E.A.Catches., gen. nov.

Typ e: Antrelloides atroceracea P.S.Catches. &

D.E. A. Catches.

Mycobank number: MB825520.

Apothecial ascomata up to 40 mm diam. Hymenium
convex; firm, waxy; black, violaceous-black. Basal
stipe substantial; obconic; a chambered base of knobby

columnar ridges radiating from base; fragile, waxy. Asci
amyloid; cylindrical; operculate; 8-spored. Ascospores
ellipsoid; smooth under light microscope. Paraphyses
longer than asci; septate; tips swollen and tending to
aggregate, encrusted with brown amorphous matter;
branching occasionally near tips. Medullary excipulum
of several layers of interwoven hyphae, grading into
ectal excipulum. Ectal excipulum of large globose to
subglobose cells with interwoven hyphae. Stipe tissue of
textura globulosa and chains of subglobose cells.

Etymology. From the Latin antrellum , a small grotto or
cavern, and the Greek -oides, like.

Antrelloides atroceracea P.S.Catches. & D.E.A.Catches.
sp. nov.

Holotype: South Australia. Flinders Chase National
Park, Kangaroo Island: On soil surface in slight
depressions on side of path in lateritic sandy soils,
35° 56’ 25”S, 136° 43’ 58”E, alt. c. 65 m, heath with
Banksia marginata Cav., Leptospermum continentale
Joy Thomps., Melaleuca gibbosa Labill., Isopogon
ceratophyllus R.Br., Petrophile multisecta F.Muell.,
Lepidosperma semiteres F.Muell. ex Boeck., Hakea
mitchellii Meisn., 6 July 2007, PS. Catcheside PSC
2710&D.E.A. Catcheside (AD-C 55811).

Mycobank number: MB825521.

Apothecia solitary, scattered; occasional to frequent;
shiny to dull; black, violaceous-black 38; forming
circular to irregular cushion-like mounds above soil
surface or amongst laterite nodules (Figs 1A & IB).
Whole fruit body broadly obconic, turbinate; diameter
(10—) 15-40 mm, height above ground 6-15 mm,
total height 15—30 mm. In cross section, the whole
fruit body resembles a miniature cavern of continuous
columns like stalactites and stalagmites fused at the base
and radiating upwards and outwards to the capping
hymenium (Figs 1C & ID). Disc convex, irregularly
domed; black, violaceous-black 38; shiny to matt;
smooth to wrinkled, undulating, occasionally pitted;
waxy, brittle; margin smooth to irregularly lobed.
Flesh 0 .6-1 mm thick; exterior black, lower layers pale
grey. Stipe continuing below soil surface; diameter
17-38 mm, depth 16-24 mm; deeply and irregularly
lacunose; forming a series of knobbly columns fused at
the base and radiating out from base; height of columns
15-28 mm, diameter of columns 2.5-5 mm; brown-
black-grey 5F3, 6F3-5 at top, grading to grey 7E3, 6D3
at base (dark red-brown 6F5-7F5 in WA specimens);
texture waxy, fragile. Asci cylindrical-clavate (250—)
272—316 (—340) x (8.5—) 11—15 pm, average (264—)
287.8 (-297) x 9.14 (-13) pm (Figs 2B & 2C);
8 -spored; amyloid over entire length but more strongly
amyloid at apex (Figs 2A & 2D); operculate; tips
rounded; base forked, arising from croziers (Fig. 2E).
Ascospores long ellipsoid; (16.8-) 17.6-22.4 (-24) x
6.4-8.8 pm, average (18.29—) 20.33 (-21.2) x (7.12-)
7.74 (-8.09) pm; Q range 2.4-2.8 (-3.0), Q average
(2.35-) 2.61 (-2.76); appearing smooth under light
microscope (x 1000) (Fig. 2H), but densely verrucose
when viewed with SEM (Fig. 21); thick-walled; often

82

Swainsona 31 (2018)

Antrelloides atroceracea, a new genus and species in the Pezizaceae

Fig. 1. A-E Antrelloides atroceracea. A Two fruit bodies in situ-, B single fruit body in situ (collection FMK1145); C section of fruit body
(holotype); D section of fruit body (collection PSC 4292); E habitat of Antrelloides atroceracea in heath. F Section of Ruhlandiella
berolinensis (collection PSC 4322). Scale = 10 mm. Photos: D.E.A. Catcheside.

83

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

Swoinsono 31 (2018)

Fig. 2. Antrelloides atroceracea. A Hymenium showing asci, ascospores, paraphyses and excipular tissue (in Melzer's solution);
B hymenium showing asci, ascospores, paraphyses (in 5% KOH); C hymenium showing asci, ascospores, paraphyses (in water);
D asci (in Melzer's solution); E asci (in Congo Red); F bifurcating paraphyses (in Congo Red); G tips of paraphyses (in water);
H ascospores (in 5% KOH); I scanning electron micrographs of ascospores; J ectal excipulum (in water); K stipe tissue (in Congo
Red). Scale: A-F 100 pm; G, H, J, K 10 pm; I 5 pm Photos: P.S. Catcheside.

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Swainsona 31 (2018)

Antrelloides atroceracea, a new genus and species in the Pezizaceae

with two largish globules; mostly in upper part or with
occasional ascospore in lower half of ascus; uniseriate,
irregularly uniseriate, biseriate near top in some asci.
Paraphyses straight; longer than asci; septate; tips slightly
swollen 5-8 pm at tip; tips brown and encrusted
with brown amorphous matter (Fig. 2G), brown
pigment occasionally extending down into upper part
of paraphysis; tips separate or clumped; occasionally
bifurcate at tip (Fig. 2F). Subhymenium 25-40 pm
thick, of densely woven septate hyphae and small cells to
6 pm in length. Medullary excipulum a mixture of four
types of tissue: i. densely interwoven hyphae of diameter
5-10 pm and not swollen at septa; ii. hyphae composed
of cells swollen at septa and appearing ‘knuckled’; iii.
chains of ‘balloon’ cells joined by short hyphal cells; iv.
regularly and irregularly globose and subglobose cells
15-45 pm diameter. Medullary excipulum grading into
the larger cells of the ectal excipulum. Ectal excipulum
(Fig. 2J) of globose, subglobose, sub-polygonal cells
approx 20-65 (-80) pm diameter; cells thin- to thick-
walled, contents clear to brown-pigmented, some cells
encrusted. Stipe tissue mostly of globose, subglobose
cells 20-60 (-70) pm diameter but intermixed with
chains of small, irregular globose cells and hyphae
swollen at septa and appearing ‘knuckled’ (Fig. 2K).

Habitat. Occurring on sandy and sandy lateritic soils.

Etymology. Latin ater, black, ceracea , waxy.

Additional specimens examined

WESTERN AUSTRALIA. North of Bow Bridge, Willmott
Forest Block on Roe Road, in grey sand on track, 34° 58’S,
116° 57’E, 4 June 2008, Eucalyptus marginata Sm. (jarrah),
Andersonia caerulea R.Br., Astartea fascicularis (Labill.) DC.,
Melaleuca sp., Taxandria parviceps (Schauer) J.R.Wheeler
& N.G.Marchant, Katrina Syme & Julie Fielder KS2094/08
(AD-C 56009).

SOUTH AUSTRALIA. All collections from: Kangaroo Island,
Flinders Chase National Park, Platypus Waterholes Walk in
similar soils; on soil surface or in slight depressions on side
of path in lateritic sandy, soils, heath with Banksia marginata
Cav., Leptospermum continentale Joy Thomps., Melaleuca
gibbosa Labill., Isopogon ceratophyllus R.Br., Petrophile multisecta
F.Muell., Lepidosperma semiteres F.Muell. ex Boeck., Hakea
mitchellii Meisn. 35° 56’ 25”S, 136° 43’ 58”E, alt. c. 60 m,
4 June 2008, PS. Catcheside PSC 2899 & D.E.A. Catcheside
(AD-C 59830); 35° 56’ 26”S, 136° 43’ 58”E, alt. c. 60 m,

27 June 2008, PS. Catcheside & Katrina Syme (FMKI 145;
AD-C 55361); 35° 56’ 25.5”S, 136° 43’ 58”E, alt. c. 60 m,

28 June 2010; PS. Catcheside PSC 3422 & D.E.A. Catcheside
(AD-C 57285); 35° 56’ 5.6”S, 136° 44’ 4l.6”E, alt. c. 65 m,
30 June 2011, PS. Catcheside PSC 3600 & D.E.A. Catcheside
(AD-C 57335); 35° 56’ 5.7”S, 136° 43’ 4l.8”E, alt. c. 65 m,
23 June 2013, PS. Catcheside PSC 3745 & D.E.A. Catcheside
(AD-C 58504); 35° 56’ 8”S, 136° 43’ 45.7”E, alt. c. 65 m,
23 June 2013, PS. Catcheside PSC 3751, D.E.A. Catcheside &
H.P. Vonow (AD-C 58378); 35° 56’ 23.5”S, 136° 44’ 57.5”E,
alt. c. 65 m, 24 June 2015, PS. Catcheside PSC 4292, D.E.A.
Catcheside&H.P. Vonow (AD-C 60144); 35° 56’ 23.5”S, 136°
44’ 57.5”E, alt. c. 65 m, 24 June 2015, PS. Catcheside PSC
4386, D.E.A. Catcheside &A. Winston (AD-C 60145).

Results of molecular analysis

A 1566bp sequence for Antrelloides atroceracea
(GenBank MH722261), encompassing 18S part, ITS1,
5.8S, ITS2, 28S part, was obtained from PSC 2710
and PSC 3600, which were identical in sequence, and
showed no heterozygosity. Bayesian and maximum
likelihood phylogenetic analysis places A. atroceracea
in the Pezizaceae with the closest known relatives
being Peziza natrophila and Lepidotia hispida (Fig. 3).
Antrelloides atroceracea differs by 4.8% over 835bp in
snps, indels and other mismatches in 28S ribosomal
sequences from P. natrophila (AF335152.1 [isotype]
and AF335153.1) and L. hispida (as P. quelepidotia
AY640959.1 and KT869021.1 in GenBank),
which are identical over the available 28S sequence.
Lepidotia hispida (as P. quelepidotia KT869020.1 in
GenBank) and A. atroceracea differ by 5 snps in their
5.8S ribosomal genes and diverge by 31% in ITS1 and
51% in ITS2.

Discussion

Morphological form of Antrelloides atroceracea

When we first found ‘Black Shoe Leather’ we were
puzzled about its placement within the Pezizales.
Antrelloides atroceracea is clearly a member of this order:
its ascomata are apothecial and its asci are operculate,
paraphyses are present and its ascospores are non-septate.
However, its unusual morphological characteristics pose
particular challenges when determining its taxonomic
affiliations. Antrelloides atroceracea has individual
morphological characters similar to those of taxa from
a number of genera, but the fungus as a whole does not
conform to any described genus that we know.

With its above-ground disc and its buried stipe
Antrelloides atroceracea presents problems as to how to
classify its form. Some members of the Pezizales are
epigeous, others hypogeous and the ascomata exhibit
very varied forms. Weber et al. (1997), Moreno et al.
(2014) and Frey et al. (2016) have proposed terms,
including apothecium, stereothecium, pulverothecium,
cleistothecium and ptychothecium, as well as the less
common exothecium, to describe the various fruit body
forms. Stereothecia, pulverothecia and cleistothecia
are closed structures containing indehiscent asci. The
asci may be scattered but they are not organised in a
definite hymenium. Stereothecia and pulverothecia are
macroscopic and usually hypogeous, while cleistothecia
are minute and may be on dung, plants, animals or
other fungi. A ptychothecium is hypogeal, has an
internal but organised hymenium which may be
unfolded or folded and may have one or more openings
and its asci are generally indehiscent.

Apothecia at maturity are epigeous with an open
exposed hymenium and active ascospore expulsion,
are usually discoid or cupulate and may be sessile or
stipitate. A few of the Pezizales have exothecia. These

85

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

Swainsona 31 (2018)

comprise a strongly convex external hymenium with
paraphyses longer than the asci that often form a tissue,
an epithecium, covering the asci.

The ascomata of Antrelloides atroceracea are not closed
hypogeal structures, so cannot be considered to be
stereothecia, pulverothecia or cleistothecia. The sterile
base of Antrelloides atroceracea is distinct from the
hymenium, therefore the fruit body cannot be described
as a ptychothecium. Nonetheless, its chambered base
does bear some resemblance, albeit without asci, to
the convoluted and folded inner structure of hypogean
fungi such as Mycoclelandia Trappe & G. Beaton Trappe
(Beaton & Weste 1982; Trappe & Beaton 1984; Hansen
et al. 2001 ), Peziza whitei (Gilkey) Trappe (Korf 1973;
Trappe 1975; Beaton & Weste 1982; Trappe & Claridge
1975; Hansen et al 2001) and 7? ellipsospora (Gilkey)
Trappe (Hansen et al. 2001; Desjardin et al. 2015).

The exposed hymenium of Antrelloides atroceracea
suggests the apothecial form, one of the most common
forms amongst the Pezizales. However, its convex above¬
ground hymenium bears similarities to the exothecium
of Ruhlandiella (Fig. IF).

Other taxa: similarities and differences

The definition of an apothecium may be extended to
genera such as the stipitate Helvella and Gyromitra with
their often convoluted pilei. Fungi in these genera show
some similarities with Antrelloides atroceracea. Its convex
disc is not convoluted as are the pilei of many species of
Helvella and Gyromitra but its chambered base or stipe is
reminiscent of the stipe of some species of those genera,
particularly those of Helvella umbraculiformis Seaver
(family Helvellaceae) (Seaver 1942) and Gyromitra
californica (W. Phillips) Raitv. (family Discinaceae)
(Phillips 1880; Seaver 1942; Kuo 2012; Mykoweb;
Beug et al. 2014). Neither taxon is known to occur in

Fig. 3. A Bayesian tree based on
sequences of the 28S ribosomal
gene showing phylogenetic
relationship of Antrelloides
atroceracea with representative
species from genera in
the Pezizaceae. Posterior
probabilities of > 0.999, > 0.99
and > 0.95 are shown by 1, **,
and * respectively above nodes
and maximum likelihood values
greater than 70% are shown
below the nodes. The sequence
for PSC 4110 Sphaerosoma
trispora was derived in this
study (GenBank MH722262),
other numbers identify
GenBank sequences. Truffle like
taxa are indicated by -T.

AY500526.1 Ascobolus carbonarius

95

- AY500546.1 Peziza gerardii

-PSC4110 Sphaerosoma trispora

AY500538.1 Marcelleina psendoanthracina

- AF335114.1 Boudiera dennisii

99

92

100

100

-±J~
ioo bAi
ioo 1

97 *- HM996916.1 Aquapeziza gl obispora

- AF335123.1 Pachyella babingtoni

I— AF335124.1 Pachyella punctispora

' - AF335123.1 Pachyella babingtoni

JN121353.1 Amylascus sp. -T
KP027405.1 Pachyphlodes depressus -T
- AY500544.1 Pachyphlodes carneus -T
KT899973.1 Pachyphlodes pfisteri -T
PSC2710 & PSC3600Antrelloides atroceracea

85

AF335152.1 Peziza natrophila
AY640959.1 Lepidotia hispida
AF335129.1 Peziza apiculata

FJ228463.1 Calongea priegnensis -T
JF343549.1 Ruhlandiella peregrina
AF335175.1 Ruhlandiella berolinensis
AF335168.1 Peziza whitei -T
1 |— GQ231743.1 Mycoclelandia arenacea -T
oo 1 GQ231747.1 Mycoclelandia bulundari -T
GQ231750.1 Ulurua nonparaphysata -T
U42694.1 Cazia Jlexiascus-T
HQ698057.1 Terfezia leptoderma -T
HQ698054.1 Terfezia arenaria -T
HQ698055.1 Terfezia claveryi -T
AF335139.1 Peziza ellipsospora
AF335132.1 Peziza badiofusca
AF335147.1 Peziza limnaea
AF335177.1 Tirmania nivea -T
AF335178.1 Tirmania pinoyi -T
AY232725.1 Eremiomyces echinulatns
_ — AY789328.1 Peziza phyllogena
*j- AY500533.1 Hapsidomyces venezuelensis
AY500553.1 Plicaria anthracina
AY500554.1 Plicaria trachycarpa

* ■- AY544668.1 Sarcosphaera crassa

-Kps'-AY500555.1 Sarcosphaera coronaria

100 L AF335115.1 Hydnotryopsis setchellii - T
■ AF335148.1 Peziza luteoloflavida
AF335117.1 Iodowynnea anriformis

- AY500534.1 Iodophanus carneus

— GQ231734.1 Elderia arenivaga -T
GQ231755.1 Mattirolomyces terfezioides -T
GQ231740.1 Mattirolomyces mulpu -T
HQ660382.1 Mattirolomyces spinosus -T
- AY544659.1 Peziza proteana
AY500548.1 Peziza lobidata

42

ljTT

HO,

AF335126.1 Peziza ammophila
AF378367.1 Peziza vesiculosa
AY789391.1 Peziza varia
9Y1— AF335138.1 Peziza echinospora
AF335137.1 Peziza domiciliana

0.05

86

Swainsona 31 (2018)

Antrelloides atroceracea, a new genus and species in the Pezizaceae

Australia. Both species have convoluted pilei and the
surface of the columns or ribs of the above-ground stipe
are dry, relatively smooth and whitish. The internal
structure of the erect columnar fruit bodies of the
Australian species Underwoodia beatonii Rifai (family
Helvellaceae) (Rifai 1968) is also somewhat similar to
the South Australian and Western Australian collections
of Antrelloides atroceracea. The columns of U. beatonii
are divided internally by longitudinal ridges separated
by alveolar cavities but the ridges are smooth, whitish
and not as brittle as those of A. atroceracea. A major
difference with taxa in the families Helvellaceae and
Discinaceae is that they have inamyloid asci (Frey et al.
2016), in contrast to the amyloid asci of A. atroceracea.

Two families within the Pezizales have amyloid asci,
the Pezizaceae (Hansen et al. 2001; Hansen et al. 2005;
Hansen & Pfister 2006; Laessoe & Hansen 2007;
Hansen et al. 2013) and the Ascobolaceae (Hansen
& Pfister 2006; Hansen et al. 2013; Frey et al. 2016).
Antrelloides atroceracea may be excluded from the
Ascobolaceae since its asci do not protrude at maturity,
its opercula are small not large and its spores do not
darken on maturity. The asci of Antrelloides atroceracea
are amyloid along their length but more strongly
amyloid at the tips, conforming most closely with type
(i) asci (Hansen et al. 2001).

The Pezizaceae is a large and very diverse family with
an estimated thirty-two genera (Frey et al. 2016). It
was possible that Antrelloides atroceracea was a member
of one of these genera. We looked for similarities with
genera with apothecia within the Pezizaceae. Such
genera include Pachyella Boud., Scabropezia Dissing
& Pfister, Boudiera Cooke, Hapsidomyces Krug &
Jeng, Iodophanus Korf, Iodowynnea Medel, Guzman
& Chacon, Sarcosphaera Auersw., as well as the more
common genera Peziza Fr. and Plicaria Fuckel. We did
not discount genera that had not been recorded from
Australia since there is a lack of extensive collections of
Ascomycetes in this country (Rifai 1968; Hyde 2001).

Pachyella, Plicaria and Scabropezia (Hansen & Knudsen
2000 ) have epigeal, sessile fruit bodies, hence differing
from Antrelloides atroceracea with its substantial
subterranean base. While species of Pachyella and
Plicaria are known from Australia, there are no species
of Scabropezia in the recorded data of the Atlas of
Living Australia.

Iodowynnea is a monotypic genus which was described
from Africa and tropical America (Medel et al. 1996).
Iodowynnea auriformis (Pat. ex Le Gal) Medel, Guzman
& Chacon does have a hypogeous stipe but this
bears caespitose clumps of individual apothecia. The
ellipsoid spores are ornamented with warts arranged
in longitudinal bands differing from the spores of
A. atroceracea whose verruculose ornamentation can be
detected only with scanning electron micrography.

The coprophilous Hapsidomyces venezuelensis
J.C.Krug & Jeng and species of Iodophanus differ

from A. atroceraea not only in their ecology but in
their size: apothecia seldom reach more than 3 mm
in diameter. Moreover the spores of Hapsidomyces
venezuelensis are globose and reticulate (Krug & Jeng
1984), those of species of Iodophanus, though ellipsoid,
are covered with a mucilaginous sheath bearing callose-
pectate ornamentation (Kimbrough et al. 1969). Only
Iodophanus carneus (Pers. ex Pers.) Korf apud Kimbrough
& Korf has been described from Australia (ALA).

The apothecia of Sarcosphaera are hypogeous during
development, closed at first but split open in a stellate
manner when mature to expose a smooth, whitish to
pale violaceous hymenium, characters differing from
A. atroceracea. Asci are amyloid and spores are ellipsoid.
The genus has not been recorded in Australia.

Although some truffle-like genera share the amyloidity
of asci and smooth, ellipsoid spores, their hypogeal
habit and internal hymenium separate them from
Antrelloides atroceracea.

While the above-ground, broadly convex disc of
Antrelloides atroceracea may resemble discs of either
Peziza or Plicaria, its distinctive below-ground,
chambered base is not like any pseudostipe of any
species of those genera. They share the amyloid
character of asci but Plicaria species have globose spores,
thus separating species in that genus from Antrelloides
atroceracea with its ellipsoid spores, a character they
share with species of Peziza.

Exothecial taxa

The few genera considered to have exothecial fruit
bodies are Ruhlandiella P. Henn. (Lantieri et al. 2012 ;
Laessoe & Hansen 2007; Frey et al. 2016), Sphaerozone
Zobel (Beaton & Weste 1978; Hansen et al. 2001;
Laessoe & Hansen 2007) and Sphaerosoma Klotzsch.
Dissing & Korf (1980) discussed relationships amongst
these genera and also Boudiera Cooke, separating
genera whose spores are actively discharged, Boudiera
and Sphaerosoma, from Ruhlandiella and Sphaerozone
whose asci are indehiscent. All are epigeous.

The genus Ruhlandiella is considered native to Australia
(Lantieri et al. 2012). However, the type species, the
mycorrhizal Ruhlandiella berolinensis Henn. occurs not
only in Australia but in plantings of Eucalyptus in Spain
(Galan & Moreno 1998), Greece (Agnello & Kaounas
2010), the Canary Islands and California (Dissing & Korf
1980). The tiny fruit bodies from our single collection of
R. berolinensis were draped over the soil surface (Fig. IF).
Although the hymenium has a similar form to that of
Antrelloides atroceracea, with paraphyses forming a semi¬
cover over the asci in both taxa, the sterile lower surface of
R. berolinensis is miniscule when compared with the large
chambered base of A. atroceracea. Moreover, the walls of
the asci fragment and release the ascospores within the
hymenium while the walls of the asci of A. atroceracea
are not evanescent. Its asci are amyloid, as are those of
R. berolinensis and R. reticulata (P.H.B.Talbot) E.Rubio,
Tena, Ormad & A.Suarez, but asci of R. peregrina

87

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

Swainsona 31 (2018)

Lantieri & Pfister and R. truncata are inamyloid (Warcup
& Talbot 1989; Galan & Moreno 1998; Rubio et al.
2010; Lantieri et al. 2012).

Sphaerozone has amyloid asci (Dissing & Korf 1980;
Laessoe & Hansen 2007); the genus is monotypic,
the type species being S. ostiolatum (Tul. & C. Tul.)
Setch. Zhang & Minter (1989) transferred two
species, Sphaerozone echinulatum G.W.Beaton and
S. ellipsosporum Cribb that had previously been included
in the genus to the new genus Gymnohydrotrya, the
new combinations being Gymnohydrotrya echinulata
(G.W.Beaton) B.C.Zhang & Minter and G. ellipsospora
(Cribb) B.C.Zhang & Minter. Sphaerozone ostiolatum
has globose, ornamented spores and its asci are
indehiscent (Dissing & Korf 1980; Pegler et al. 1993),
characters that separate it from A. atroceracea , though its
subhypogeous and convoluted and infolded fruit body
bears some slight resemblance to that taxon. Species of
Gymnohydrotrya have a subglobose to irregularly lobed
ascoma with several internal chambers and canals. The
hymenium is external but often extends internally.
They differ from Antrelloides atroceracea in their totally
hypogeous, globose ascomata, inamyloid asci and
ornamented ascospores, though the infolded chambers
bear some resemblance to the chambered base of
A. atroceracea.

Dissing & Korf (1980) recognised that Sphaerosoma
is a particularly problematic genus. Antrelloides
atroceracea bears little resemblance to the two species
that they accepted: S. fuscescens Klotzsch and S. trispora
McLennan & Cookson (McLennan & Cookson
1923). Both have inamyloid asci and globose, highly
ornamented spores, while A. atroceracea has amyloid
asci and ellipsoid spores. Ascomata of the South
Australian collections examined were olive-brown to
black, irregularly cushion- to saucer-shaped, sessile
and attached to the soil surface by a central soil pad
(Catcheside 2012). This under-collected genus requires
further investigation, though the cryptic nature and
probable rarity make this problematic.

Phylogenetic associations of Antrelloides atroceracea

Based on the studies of Hansen et al. (2001, 2003),
Hansen & Pfister (2006), Lsessoe & Hansen (2007)
and Hansen et al. (2013), Frey et al. (2016) divide the
class Pezizomycetes sensu O.E. Erikss. & Winka, Order
Pezizales J. Schrot. into four lineages with lineage A /
suborder Pezizinae Rifai (asci often amyloid) divided
into two families: Ascobolaceae with asci protruding
from the hymenium when mature and Pezizaceae with
non-protruding asci. Lineage B covers the Morchella-
Helvella clade, lineage C the Pyronemataceae s.l.
and lineage D the suborder Sarcoscyphinae Rifai.
Sphaerosoma is amongst genera incertae sedis. The
phylogenetic data presented here include taxa from all
four lineages.

There are some genera which lie in lineages other
than the Pezizaceae that have infolded ascomata
bearing some resemblance to the chambered base of

Antrelloides atroceracea. These include Hydnotrya Berk.
& Broome in lineage B, the Morchella-Helvella clade
(Trappe 1979; O’Donnell et al. 1997; Frey et al. 2016).
Hydnocystis Tub, Gene a, Vittad. and Geopora Harkn.
are in Lineage C/Pyrenomycetaceae s.l. (Trappe 1979;
Ltessoe & Hansen 2007; Moreno et al. 2014; Frey et
al. 2016). However, as well as being hypogean, they are
phylogenetically distant from Antrelloides atroceracea

(Fig. 3).

Phylogenetic analysis of Antrelloides atroceracea shows
that it is nested within the Pezizaceae (Fig. 3). It is
most closely related to Lepidotia hispida (Quel.) Boud.
(Norman & Egger 1999; Hansen et al. 2001, 2005;
Hansen & Pfister 2006; van Vooren et al. 2015) and
Peziza natrophila A.Z.M. Khan (Hansen et al. 2001;
van Vooren et al. 2015). Morphologically however,
although sharing the amyloidity of asci and globose
excipular tissue, they differ in structure and colour.
Lepidotia hispida is yellow-green, stipitate but its stipe
is simple, not chambered (Korf 1973; van Vooren et
al. 2015). Peziza natrophila is olive-brown, later dark
brown to black, shallow-cupulate to discoid and sessile
(Nowsher & Khan 1976). The habitats of both taxa,
Lepidotia hispida in peat bogs or swampy areas (Korf
1973; van Vooren et al. 2015) and Peziza natrophila
known mostly from plots treated with sodium and
potassium carbonates in pine plantations but also from
an old peat bog (Korf 1973; O’Donnell & Beneke
1973; Hansen et al. 2001; van Vooren et al. 2015)
differ substantially from that of A. atroceracea , which,
although fruit bodies may be subject to inundation, is
found in dry sandy to sandy-clay soils. It is possible that
Lepidotia hispida and Peziza natrophila are con-specific
(Hansen et al. 2001; van Vooren et al. 2015).

The 4.8% difference of the 28S ribosomal gene
sequence of Antrelloides atroceracea to that shared by
Peziza natrophila and Lepidotia hispida , coupled with
the very large differences in ITS1 and ITS2 sequence
(31% and 51% respectively) from that of L. hispida,
argue for the erection of the new genus Antrelloides.

Taxa in adjacent branches of the phylogenetic tree
include species of Pachyphlodes, Amylascus, Boudiera,
Aquapeziza and Pachyella. The truffle-like Pachyphlodes
and Amylascus have globose and echinulate spores,
differing from the epigean Antrelloides atroceracea
with its ellipsoid, faintly verruculose spores. Species of
Boudiera, Aquapeziza and Pachyella lie in neighbouring
branches. The latter genus has sessile ascomata thus
differing from the stiped A. atroceracea, Boudiera has
tiny apothecia which seldom reach more than 3 mm
diameter and spores are spiny or reticulate. The
monotypic Aquapeziza globispora D.M.Hu, L.Cai &
K.D.Hyde (Hu et al. 2012) has sessile, white apothecia
and smooth, globose spores. Its habitat on submerged
wood in freshwater streams excludes it from Antrelloides.
Peziza apiculata Cooke is sessile and has ellipsoid spores
with needle-like apicules at each end. It grows on rotten
wood or moist soil (Moravec 1977).

88

Swainsona 31 (2018)

Antrelloides atroceracea, a new genus and species in the Pezizaceae

General discussion

The form of the fruit body and the habitat of
Antrelloides atroceracea are interesting. Over a limited
area fruit bodies are not uncommon. They are almost
always on the edges of slightly raised, bare tracks and
are subject to inundation, with standing water at least
up to the margins of the fruit bodies. One population
of immature fruit bodies was found on sticky, sandy
clay soil in a slight depression where it was obvious
that the water level had reached the margins of the
discs. The substantial ribbed base anchors the fruit
body in the soil, preventing it from being washed away.
Species of Boudiera and Pachyella grow in swampy
ground and have ascomata that are able to withstand
flooding. Boudiera species are mostly on sand, Pachyella
species on water-soaked wood or decaying plant
debris (Pfister 1973; Hansen et al. 2001). Species of
both genera are sessile, lacking the chambered base of
A. atroceracea which anchors that taxon in the ground.
The hymenium of species of Boudiera occupies the
upper surface or covers the whole exposed surface
of the apothecium and its asci protrude beyond the
paraphyses when mature (Seaver 1942; Eckblad 1968),
possibly facilitating spore dispersal in wet conditions.
Pachyella species have gelatinous tissues, an adaptation
to its wet habitat.

Placement of new fungal taxa always presents
challenges. When the new fungus has no obvious
characters that fit it into any group it becomes more
problematic. Phylogenetic analysis is essential,
especially in these circumstances. However, molecular
data are often not available, especially for cryptic and
thus often, in Australia, under-collected taxa such as the
Discomycetes. Its genetic characters and the amyloidity
of the asci enable Antrelloides atroceracea to be placed
firmly in the Pezizaceae. Molecular sequencing and
phylogenetic analyses enable relationships between
species and genera to be understood but demonstrate
that, as in this case, there is little relationship between
morphological and genetic characters. In addition, Rifai
(1968) commented on the lack of extensive collections
of Australian Pezizales. From our experience, this has
changed little since Rifai’s observation.

Additional species examined

Sphaerosoma trispora

SOUTH AUSTRALIA: Mays Cottage to Platypus Walking
Track, Flinders Chase National Park, Kangaroo Island,
35° 56’S, 136° 44’E, 27 June 2011, PS. Catcheside PSC
3570, D.E.A. Catcheside, T.P. Bridle & H.P. Vonow (AD-C
56992); 35° 56’S, 136° 43’E, 28 June 2014, PS. Catcheside
PSC 3583, D.E.A. Catcheside, H.P. Vonow, T.P. Bridle & P.
Bridle (AD-C 56993); Mount Rescue Conservation Park
near Tintinara, 35° 55’S, 140° 17’E, 17 Aug. 2014, PS.
Catcheside PSC4110 & D.E.A. Catcheside (AD-C 58768)
(GenBank MH722262); Meningie, on ground, 29 June
1961, L.D. Williams 1173 (AD-C 47697); Meningie, on
ground, 7 Sep. 1961, L.D. Williams 1246 (AD-C 47693);
Wood’s Well, on ground, 27 Aug. 1961, L.D. Williams 1236
(AD-C 47691).

Ruhlandiella berolinensis

SOUTH AUSTRALIA: Kaiserstuhl Conservation Park,
S34° 34’, E139° 1’, 22 Aug. 2016, PS. Catcheside PSC 4322,
D. Catcheside & members of Adelaide Fungal Studies Group
(AD-C 59899).

Acknowledgements

We thank Dr Karen Hansen and an anonymous referee
for their thoughtful, constructive and very helpful
comments. We are grateful to Katrina Syme for kindly
providing her collection and for her comments on the
paper, Carolyn Ricci for her help with scanning electron
microscopy, Bob Baldock for his help in preparing
slides, the State Herbarium of South Australia (AD)
for provision of equipment and materials. We thank
Lisa O’Donovan at Adelaide Microscopy for advice on
preparation of specimens and to Adelaide Microscopy
for doing sputter-coating of the specimen. We also
wish to thank Frank Kutsche for assistance in obtaining
relevant scientific collecting permits for South Australia
and Rangers of Conservation and National Parks for
their assistance and interest. With Trish Mooney we
enjoyed our first encounter with ‘Black Shoe Leather’.
We are indebted to Helen Vonow, Collections Manager
at AD, for her keen ability to spot apothecia of ‘Black
Shoe Leather’. We are very grateful to the State
Herbarium of South Australia and Flinders University
for their continual support.

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Swainsona 31:93-100 (2018)

© 2018 Board of the Botanic Gardens & State Herbarium (Adelaide, South Australia) ©0

The Ptilotus murrayi species group: synonymisation of P. petiolatus
under P. murrayi and description of the new Western Australian species
P. unguiculatus (Amaranthaceae)

Timothy A. Hammer

School of Biological Sciences, Faculty of Science, The University of Western Australia, 35 Stirling Highway,

Crawley, Western Australia 6009

Email: timothy.hammer@research.uwa.edu.au

Abstract: The Ptilotus murrayi F.Muell. species group encompasses the morphologically similar
P. murrayi, P. gomphrenoides F.Muell. ex Benth. and P. petiolatus Farmar. These three species overlap in
distribution in Western Australia, where they are easily confused with one another. A critical examination
of specimens shows that there are no consistent morphological characters to justify the separation of
P. murrayi and P. petiolatus, and therefore the latter is formally synonymised under the former. Ptilotus
sp. Edaggee Station (T.E.H.Aplin 3208) is found to be a distinctive new species and described as Ptilotus
unguiculatus T.Hammer. Updated descriptions are given for P. murrayi and P. gomphrenoides.

Keywords: Amaranthaceae, Australia, new species, Ptilotus, taxonomy

Introduction

Ptilotus R.Br. (Amaranthaceae) is a genus of c. 120
species native to Australia, mostly occurring in arid or
semi-arid regions of Australia, with the highest diversity
in Western Australia (Hammer et al. 2015). A recent
molecular phylogeny by Hammer et al. (2015) resolved
several molecular clades within the genus, including
the large clade informally named ‘clade B’. Within
clade B, Ptilotus murrayi F.Muell. and P gomphrenoides
F.Muell. ex Benth. were resolved as sister species. The
morphologically similar P. petiolatus Farmar was not
included in the phylogeny (Hammer et al. 2015).
Together these species comprise the P. murrayi species
group.

Ptilotus murrayi was described by Mueller (1863) based
on a type from Will’s Creek (the Diamantina River,
incorrectly noted in the protologue as ‘Cooper’s Creek’)
near the border of South Australia and Queensland,
collected by J. Murray on the Howitt expedition
(Fig. 1A). Specimens of P. murrayi from Western
Australia were not available to Mueller, but he did
mention having seen similar specimens from Western
Australia that had pedunculate inflorescences:

Vidi etiam specimina manca plantae persimilis ad
montes Hamersley Range sub expeditione Francisci
Gregorii collecta, spicis pedunculatis praecipue
divellenda.

Mueller (1863: 145)

Bentham (1870) formally described P. gomphrenoides
based on these pedunculate specimens from Gregory’s

expedition in the Hamersley Range (Fig. IB). Farmar
(1905) erected P. petiolatus from specimens collected
by E. Clement in north-western Western Australia in
1897. He separated P. petiolatus from P. murrayi on
the basis that the former has inflorescences with acute
apices, shorter staminal filaments and larger anthers:

[...] quod flores P. murrayi simillimus sed spicis acutis
filamentis brevioribus antheris majoribus differt

Farmar (1905: 1089)

He went on to say that P. murrayi was incompletely
known, and that only a few fragments (‘an inch in
length’) were available at K, however he felt confident
that these differences were sufficient to erect the new
species.

Farmar (1905) also erected P. roseo-albus Farmar within
this species group, differing from P. gomphrenoides in
having shortly pedunculate spikes (as opposed to long-
pedunculate) and a denser abaxial sepal indumentum.
Within P. roseo-albus he described var. conglomeratus
Farmar, which differed from the typical variety in
having cylindrical inflorescences in clusters, as opposed
to conical and loosely arranged inflorescences. The
two varieties of P. roseo-albus were recombined under
P. gomphrenoides by Beni (1962) as P. gomphrenoides
var. roseo-albus (Farmar) Beni and var. conglomeratus
(Farmar) Beni, stating that the characters that Farmar
used were common characteristics of P. gomphrenoides.
Hammer & Davis (2017) recently synonymised these
varieties under the typical P. gomphrenoides , as these
varieties were found to intergrade imperceptibly into
one another. Black (1923) described P. murrayi var.
major J.M.Black from the Diamantina River, South

Published online: 28 Sep. 2018* flora.sa.gov.au/swainsona

ISSN 2206-1649 (Print) • ISSN 2206-1657 (Online)

T.A. Hammer

Swainsona 31 (2018)

Fig. 1 . Comparative morphology of flowering inflorescences. A Ptilotus murrayi; B P. gomphrenoides. Photos: A K.R. Thiele; B R. Davis.

Australia, on the basis of its larger leaves, longer
stems and inflorescences, and slightly longer sepals.
This variety was synonymised by Bean (2008), who
recognized that the characters attributed to P murrayi
var. major were within the typical variability of
var. murrayi. In a manuscript by Beni (unpubl.),
P. petiolatus subsp. limbatus Beni MS was included
as a manuscript name based on a specimen from
Edaggee Station, Western Australia (Fig. 2), which he
primarily differentiated from subsp. petiolatus by the
new subspecies having clawed sepals. While the new
taxon was never formalised, it was included under the
phrase name Ptilotus sp. Edaggee Station (T.E.H.Aplin
3208) at the Western Australian Herbarium awaiting
evaluation of its taxonomic status.

The geographic distributions of taxa in the P. murrayi
species group overlap significantly in Western Australia,
and this, coupled with the strong morphological
similarities, has led to some confusion between them.
Ptilotus murrayi has a disjunct distribution, occurring
in Western Australia from Carnarvon, Gascoyne,
Murchison, Pilbara and Dampierland (Interim
Biogeographic Regionalisation for Australia, IBRA;
Department of the Environment 2018) regions and
in south-western Queensland and north-eastern South
Australia in the Mitchell Grass Downs and Channel
Country IBRA regions. It has not been recorded
from the Northern Territory or north-western South
Australia (Fig. 3A). Ptilotus petiolatus and P. murrayi are
sympatric throughout the range of the former. Ptilotus
petiolatus is known from 15 specimens, ten from

Dampierland and five from the Pilbara IBRA regions.
Ptilotus gomphrenoides is also almost entirely sympatric
with P. murrayi in Western Australia, occurring in
the Carnarvon, Gascoyne, Pilbara, and northern
Murchison bio regions (Fig. 3B). Ptilotus sp. Edaggee
Station (T.E.H.Aplin 3208) is known from a single
collection on Edaggee Station, approximately 70 km
SSE of Carnarvon, Western Australia and 75 km SW
of the southwestern-most record of P. murrayi , which
occurs near the Gascoyne River (Fig. 3B).

During the construction of an online identification
key to all species in Ptilotus for the Flowering
Plants of Australia project on Key Base (available at
http://keybase.rbg.vic.gov.au/keys/show/ 6609),
P. murrayi , P. gomphrenoides and P petiolatus were
found to be difficult to discriminate due to unclear
morphological boundaries, which was also noted by
Burbidge (unpubl.) in a key constructed for Ptilotus. A
thorough examination of the morphology of specimens
for each species was conducted to critically evaluate
the species boundaries within the group. This showed
that specimens of P. murrayi and P. petiolatus could not
be reliably separated, including those that Farmar had
originally used to erect P. petiolatus. As a result, the later
name of P. petiolatus is here formally synonymised under
P. murrayi. Ptilotus sp. Edaggee Station (T.E.H.Aplin
3208) is also assessed and found to be a very distinctive
species. The morphological differences between the
two accepted species in the P. murrayi species group
(i.e. P. murrayi and P. gomphrenoides) are clarified and
descriptions are given for both.

94

Swainsona 31 (2018)

The Ptilotus murrayi species group

Material and methods

This study was based on the examination of dried
specimens of 7? gomphrenoides, P. murrayi , P. petiolatus
and P. sp. Edaggee Station (T.E.H.Aplin 3208)
housed at PERTH and specimens of P. murrayi on
loan from BRI and AD, covering the geographic
range of all species. Scans of type specimens were
assessed through JSTOR Global Plants (https://plants.
jstor.org/). The specimens were critically evaluated
irrespective of current determinations and matched
to the protologues and type specimens, and when
necessary, specimens were re-determined as a result of
this study. Care was taken to evaluate the specimens
based on their likely natural posture in the field,
and when available, label data were used to infer the
natural habit of the specimens. A distribution map of
the species was produced using georeferenced records
retrieved from the Australasian Virtual Herbarium
(https://avh.chah.org.au/). The terminology used in
this paper for the perianth of Ptilotus is different from
what is conventionally used within Australia. The term
‘sepal’ is used here in place of tepal’, as recent studies
support the calycine origin of the uniseriate perianth
for all Caryophyllales, the corolla having been lost (see
Ronse De Craene 2013; Vrijdaghs et al. 2014).

Results and discussion

The examination revealed that P. sp. Edaggee Station
(T.E.H.Aplin 3208) is clearly a distinctive species
within the genus. Beni (unpubl.) intended to describe
this taxon as a subspecies of P. petiolatus. While it is
understandable that he believed the two entities were
related, given their superficial similarities (e.g. habit,
leaf size and shape, inflorescence size and shape;
Fig. 2), P. sp. Edaggee Station (T.E.H.Aplin 3208) is
significantly different from P. petiolatus in important
floral characters. It may not be closely related to the
P. murrayi species group. The sepals of P. sp. Edaggee
Station (T.E.H.Aplin 3208) are conspicuously clawed
(i.e. narrow at the base and dilated toward the apex),
and more so in the outer sepals, where the dilated apex
is 0.7-0.9 mm wider than the base. Ptilotus petiolatus ,
P. murrayi and P. gomphrenoides do not have clawed
sepals. In other Ptilotus species, this trait is consistent
within a species and is often shared between closely
related species. Within clade B, several species groups
possess clawed sepals, such as the species group
including P. latifolius R.Br. and P. chamaecladus Diels
and the species group including P. conicus R.Br.
and P. corymbosus R.Br. Ptilotus sp. Edaggee Station
(T.E.H.Aplin 3208) also differs from P. petiolatus ,
by possessing crisped nodose hairs on the inner
margins of the sepals, which often become tangled
and obscure the ovary and staminal cup. Sepals in
Ptilotus petiolatus , P. murrayi and P. gomphrenoides are
adaxially glabrous (see Fig. 1). Ptilotus sp. Edaggee
Station (T.E.H.Aplin 3208) has longer staminal
filaments (0.7-0.9 mm) and style (0.6-0.8 mm) than

P. petiolatus (0.3-0.6 and 0.3-0.5 mm, respectively),
which ensure the anthers and stigma are raised above
these obscuring hairs. Furthermore, the bracteoles
of 7? sp. Edaggee Station are much longer and wider
than the bracts (by c. 1 mm), while the bracts and
bracteoles of the other species are similar in size. These
differences are sufficient to recognise 7? sp. Edaggee
Station (T.E.H.Aplin 3208) as a distinct species, and
therefore 7? unguiculatus T. Hammer is erected to
accommodate it (see below).

Ptilotus murrayi and 7? petiolatus can both be segregated
from 7? gomphrenoides in Western Australia by the latter
species having pedunculate inflorescences; peduncles
are short but present on some specimens (e.g. those
previously included under 7? roseo-albus). Ptilotus
gomphrenoides could additionally be segregated from the
other two species by examination of the midrib region
of the sepals (i.e. the thickened prominent midrib
flanked by two lateral veins that is clearly differentiated
from the scarious region surrounding it). All Ptilotus
species have five sepals that are arranged with two
outer sepals and three inner sepals. Sometimes the
outer and inner sepals are morphologically similar, but
in the 7? murrayi species group they are conspicuously
different. The outer sepals in the 7? murrayi species
group are conspicuously different to the inner sepals
in that they enclose the inner sepals and possess larger
and more conspicuous scarious margin, particularly
towards the apex. The midrib region and margin of
the sepal usually vary in colour in these species, being
either dark pink or green around the midrib and pink
on the margin (Fig. 1). The outer sepal has a prominent
and concaved midrib (thus allowing it to tightly
enclose the inner sepals in bud), but the midrib region
is much narrower and less starkly coloured. Ptilotus
gomphrenoides has a much wider sepal midrib region
(0.5-0.8 mm), which is usually pink to dark pink,
whereas the 7? murrayi and 7? petiolatus consistently
have a narrow (0.2-0.3 mm) and green sepal midrib
region. The outer sepals in 7? gomphrenoides have a
longer scarious margin above the midrib (0.8-1.5 mm)
and rounded apices (Fig. IB), whereas 7? murrayi and
7? petiolatus have a shorter scarious margin above the
midrib (0.5-1.0 mm) and more acute apices (Fig. 1A).
Using the characters outlined above, specimens of
7? gomphrenoides could be confidently segregated from
7? murrayi and 7? petiolatus.

The differences Farmar stated as separating
7? petiolatus and 7? murrayi do not withstand scrutiny
with the specimens currently available. All collections
of 7? petiolatus have inflorescences with an acute apex,
while collections of P. murrayi can have either an
acute or truncated apex; it is likely that Farmar had
only access to specimens of 7? murrayi with the latter
character. Farmar indicated that staminal filament
length and anther length were both important in
splitting 7? petiolatus from 7? murrayi. This study
found, however, that throughout the geographic
ranges of both species, they overlap in staminal
filament length (0.3—0.6 mm) and anther length

95

T.A. Hammer

Swainsona 31 (2018)

(0.3-0.5 mm). Beni (1979) suggested that 7 ?petiolatus
could be identified based on its unusually short style,
but 7? murrayi and 7? petiolatus were found to have
the same range of style length (0.3-0.5 mm) with the
specimens currently available.

Beni (1971) used leaf size to split 7? murrayi from
7? petiolatus and 7? gomphrenoides in his key. Leaf size
in these species imperceptibly intergrades, and is not
correlated with geography and any other characters,
rendering the use of this as a distinguishing character
ineffectual. Beni (1971) furthermore noted a difference
in phyllotaxis of 7? petiolatus and 7? gomphrenoides,
considering the former to be subopposite (‘annahernd
gegenstandig’, p. 172) and the latter to be alternate.
However, leaf arrangement in these species is not
consistent across all specimens available. Each of the
three species has specimens that have an alternate leaf
arrangement, but some specimens can be found to have
an apparent opposite or subopposite arrangement.

Bentham (1870) described 7? gomphrenoides as
apparently erect’ and 7? murrayi as apparently
prostrate’ (pp. 243-244). All three species were found
to have prostrate specimens (this character being more
common in 7? murrayi than the other species), as well
as specimens described on labels as ascending or erect.
Habit could not be correlated with other characters
(e.g. leaf size or the size of floral parts), to reliably
discriminate between the species. Several specimens in
7? gomphrenoides were found to exhibit the characters
attributed by Farmar (1905) and Beni (1971) to
7? petiolatus , such as slightly smaller flowers, leaves
and an overall more petite habit (e.g. A.C.Beauglehole
48962), which may indicate that these characters
attributed to 7? petiolatus by previous authors are within
the normal range variation for both 7? gomphrenoides
and 7? murrayi. Furthermore, no differences between
the habitats of the three species could be discerned, all
typically occurring on flat floodplains or on the fringes
of ephemeral wetlands, creeks or rivers, with clayey red
or brown soils.

Given that there are no consistent characters with
which to separate 7? petiolatus from 7? murrayi, the
former encompassing the normal variation from
throughout the range of 7? murrayi (including in
specimens available from eastern Australia), 7 ?petiolatus
is here synonymised under 7? murrayi. No significant
differences were found between the specimens of
7? murrayi from Western Australia and eastern Australia,
leaving the question open as to the significance of the
large geographic disjunction in the records between
Western Australia and eastern Australia (see Fig. 3A),
which may be more appropriately investigated with
an in-depth molecular study. Full descriptions for the
modern concepts of 7? murrayi and 7? gomphrenoides are
given below. A key to all accepted species of Ptilotus in
Western Australia, including those described here, is
currently under review by the journal Nuytsia and is
expected to be forthcoming.

Taxonomy

Ptilotus unguiculatus T.Hammer, sp. nov.

Type: Edaggee Station, Western Australia [precise
locality withheld for conservation reasons], 3 July
1970, T.E.HAplin 3208 (holo: PERTH 04073819!,
Fig. 2 ; iso: M 0241497 image!).

Ptilotus sp. Edaggee Station (T.E.H.Aplin 3208) Western
Australian Herbarium, in FloraBase, https://florabase.
dpaw.wa.gov.au/ [accessed: 9 May 2018].

Decumbent annual herb to 8 cm high, 20 cm wide. Stems
terete, ribbed, with dense nodose hairs on young growth
and the hairs becoming sparse with age. Basal leaves not
seen. Cauline leaves alternate, lanceolate to ovate, entire,
8-30 mm long, 5-11 mm wide, glabrous; base narrowly
attenuate or petiolate, 5-10 mm long; apex mucronate,
mucro c. 0.1 mm long. Inflorescences solitary or in clusters
of 2-3, terminal, sessile, spiciform, conical, 5-12 mm
long, 7—8 mm wide, white, subtended by leaves; apex
acute. Bracts broadly ovate, 1.8-1.9 mm long, 1.4—

1.5 mm wide, translucent, glabrous; midrib obscure;
apex acute. Bracteoles broadly ovate, 3.0-3.1 mm long,

2.4- 2.5 mm wide, translucent, glabrous; midrib obscure;
apex acute. Outer sepals spathulate, strongly clawed, 3.5-

3.6 mm long, 0.4-0.5 mm wide at base, 1.1-1.4 mm
wide at dilated apex; midrib region prominent, green,

1 .5- 2.5 mm long, 0.1-0.3 mm wide; basal outer surface
with long, wavy, nodose hairs, not exceeding sepal apex;
inner surface glabrous apart from crisped nodose hairs
on margins becoming dense and tangled within; apex
acute, the scarious margin 1.0-1.2 mm long, glabrous,
translucent, serrate, in-rolled. Inner sepals narrowly
spathulate, clawed, 2.5-3.1 mm long, 0.3-0.4 mm wide
at base, 0.6-0.8 mm wide at the dilated apex; midrib
region prominent, green, 1.7-2.2 mm long, 0.2-0.3 mm
wide; basal outer surface with long, wavy, nodose hairs at
base, becoming densely woolly with age, not exceeding
sepal apex; inner surface glabrous apart from crisped
nodose hairs on margins becoming dense and tangled
within; apex acute, the scarious margin 0.3-0.7 mm
long, glabrous, translucent, serrate, in-rolled. Fertile
stamens 5; filaments 0.7-0.9 mm long; anthers 0.4-
0.5 mm long, 0.2-0.3 mm wide, yellow. Staminal cup
0 .3-0.4 mm long, unlobed. Ovary globular, 0.6-0.7 mm
long, 0.6-0.8 mm wide, glabrous. Style straight, 0.6-
0.7 mm long, centrally placed on the ovary apex. Stigma
unlobed, capitate. Fruit not seen. Seed not seen.

Diagnostic characters. Ptilotus unguiculatus can be
distinguished from other members of the genus by
the following combination of characters: decumbent
habit, dense pubescence on young stems, glabrous
and petiolate leaves, sessile and terminal inflorescences
(subtended by leaves), bracts that are c. 1 mm shorter
and narrower than bracteoles, conspicuously clawed
(dilated at the apex) sepals, crisped nodose hairs on the
abaxial margin of the sepals that obscure the ovary, base
of sepal adaxial surface with long silky nodose hairs
that do not exceed the sepal apices, five fertile stamens,
staminal cup appendages absent, a central style centrally
placed on the ovary apex, and a glabrous ovary.

96

Swainsona 31 (2018)

The Ptilotus murrayi species group

Western Australian Herbarium

in ij> mi mil! in i in mill fim inn uni um m in nj n

PERTH 04073819

TEMPORARY SORTING SLIP - W.A. Herbarium (PERTH,

Please do not base « conlirmavil on this label.

sp, ( T e-H ■not')

Signed -2 qf/lfa/r'

tjr

. c OUPUC

by o t - s T5i5
i vf I

I I

Ptilotus patiol.g-'rus Jar mar

subsp. limbatus Beni

det.—rev.-G. BENL, MQnchen 198 5

WESTERN AUSTRALIAN HERBARIUM, PERTH
Flora of Western Australia

Ptilotus

Loc.

Precise locality withheld
for conservation reasons.

Lat. Long.

Coll T.E.H.Aplin 3208 3 July 19 70

Fig. 2. Scan of Ptilotus sp. Edaggee Station (T.E.H.AIpin 3208), the proposed holotype of P. unguiculatus (PERTH 04073819). Insert:
Close-up of inflorescence.

Phenology. The only examined specimen was collected
as flowering, but not fruiting, in early July. It is likely
that this species has a similar flowering and fruiting
time to other annual Ptilotus species in region, which is
typically from June to September.

Distribution and habitat. Ptilotus unguiculatus is
currently known only from Edaggee Station, in the
Carnarvon bioregion of Western Australia, where it was
collected from a disturbed area. No habitat description
was included with the specimen. Inspection of the

97

T.A. Hammer

Swainsona 31 (2018)

locality from aerial satellite images indicates it to be a
plain with open shrubs.

Conservation status. Ptilotus unguiculatus is listed
as Priority 1 under Conservation Codes for Western
Australian Flora, under the name P. sp. Edaggee Station
(Smith & Jones 2018). It is only known from the type
specimens, collected in 1970 from a disturbed area on
Edaggee Station, Western Australia.

Etymology. The specific epithet is from the Latin
unguiculatus (having claws), referring to the
conspicuously clawed sepals of the new species.

Notes. This new species may be potentially confused
with P. villosiflorus F.Muell., which is also a small
herb with greenish-white flowers, but which differs
by typically occurring on coastal dunes, having sepals
that are not clawed and abaxial sepal hairs that exceed
the sepal apex. Ptilotus latifolius and P. chamaecladus
are species closely related to P. villosiflorus that do have
clawed sepals. Ptilotus latifolius is a large rounded,
upright herb or subshrub with tangled branches that
has bracteoles that are longer than the sepals. Both
P. chamaecladus and P. latifolius have staminal cup
lobes alternating with the stamens, while the new
species has an unlobed staminal cup. The fruit of
P. villosiflorus , P. chamaecladus and P. latifolius is hard
and indehiscent, while those of the P. murrayi species
group are membranous and irregularly dehiscent. Due
to the similarity of the new species to the P. latifolius
species group and despite Beni (unpubl.) considering
the taxon to be closely allied with the P. murrayi species
group, I could not confidently place the new species in
either group. The difference in the fruit between these
two groups is an important diagnostic character, but
there was no fruit available for P. unguiculatus.

Ptilotus murrayi F.Muell.

Fragm. 3: 145 (1863). — Type: ‘From the flooded
tracts of Will’s Creek [=Diamantina River], beyond
desert’ [state unknown], 1861, J.Murray s.n. (syn.:
BM 000895594 image!, K 000357020 image!, MEL
2235243 image!, MEL 2235244 image!, PERTH
01558226!).

Ptilotus murrayi var. major JM.Black, Trans. & Proc. Roy.
Soc. S. Austral. 47: 368 (1923). — Type: ‘Between
Herrgott [=Marree] and Innamincka, South Australia,
Jun. 1916, RCockburn s.n. (holo.: AD 97747822A
image!).

Ptilotus petiolatus Farmar, Bull. Herb. Boissier ser. 2, 5:
1089 (1905), syn. nov. — Type: ‘North West Australia,
Between the Ashburton and De Grey rivers’, Western
Australia, [purchased] Aug. 1900, E.Clement s.n.
(syn.: K 000349177 image!, K 000349178 image!, K
000349179 image!).

Prostrate mat-forming or decumbent annual herb
5-20 cm high, 20-100 cm wide. Stems terete, ribbed,
glabrous. Basal leaves not seen. Cauline leaves alternate or
subopposite, narrowly lanceolate to broadly lanceolate
or spathulate, entire, 5-30 mm long, 3-9 mm

wide, glabrous; base narrowly attenuate or petiolate,
5-10 mm long; apex mucronate, mucro c. 0.1 mm long.
Inflorescences solitary or in clusters, axillary or terminal,
sessile, spiciform, cylindrical, 5-30 mm long, 4-8 mm
wide, white; apex acute or truncate. Bracts ovate, 1.0—
1.8 mm long, 0.8-1.1 mm wide, translucent, glabrous;
midrib obscure; apex rounded. Bracteoles ovate, 1.5—
2.1 mm long, 0.7-1.0 mm wide, translucent, glabrous;
midrib obscure; apex acute. Outer sepals narrowly
lanceolate, 2.0-3.3 mm long, 0.4-0.7 mm wide;
midrib region prominent, green, 1.5—2.5 mm long,
0.1-0.3 mm wide; basal outer surface with long, wavy,
nodose hairs at base, becoming densely woolly with age;
inner surface glabrous; apex acute, scarious margin 0.5-
1.0 mm long, glabrous, white. Inner sepals lanceolate,
1.8—2.8 mm long, 0.3—0.7 mm wide; midrib region
prominent, green, 1.7—2.2 mm long, 0.2—0.3 mm wide;
basal outer surface with long, wavy, nodose hairs at
base, becoming densely woolly with age; inner surface
glabrous; apex acute, scarious margin 0.3-0.7 mm long,
glabrous, pink. Fertile stamens 5; filaments 0.3-0.6 mm
long; anthers 0.3-0.5 mm long, 0.1-0.2 mm wide,
yellow. Staminal cup 0.3-0.5 mm long, unlobed. Ovary
globular, 0.7-1 mm long, 0.5-0.9 mm wide, glabrous.
Style straight, 0.3-0.5 mm long, centrally placed on the
ovary apex. Stigma unlobed, capitate. Fruit smooth,
membranous, irregularly dehiscent. Seed round, black,
glossy, c. 0.9 mm long, c. 0.7 mm wide. Fig. 1A.

Diagnostic characters. Ptilotus murrayi may be
distinguished from all other members of the genus by
the following combination of characters: a prostrate or
decumbent annual herb with glabrous stems and leaves,
sessile axillary inflorescences, sepals not clawed, outer
sepals with acute apices, a glabrous adaxial sepal surface,
densely woolly hairs on the base of the abaxial sepal
surface, 5 fertile stamens, staminal cup appendages
absent, a central style centrally placed on the ovary
apex, and a glabrous ovary.

Phenology. Ptilotus murrayi typically flowers from April
to August throughout its range, extending into late
October in Queensland and South Australia.

Distribution and habitat. Ptilotus murrayi occurs in
the Carnarvon, Gascoyne, Murchison, Pilbara and
Dampierland IBRA regions in Western Australia,
and in south-western Queensland and north-eastern
South Australia in Mitchell Grass Downs and Channel
Country IBRA regions, with several outlying collections
in the Simpson-Strzelecki Duneflelds (Fig. 3A). The
typical habitat is a flat floodplain or on the fringes of
ephemeral wetlands, billabongs or rivers with silty or
more commonly clayey red or brown soils.

Conservation status. Ptilotus murrayi is not listed as of
conservation concern in any state in which it occurs.

Selected specimens examined

QUEENSLAND: 5.6 km W of turnoff to Birdsville along
Windorah to Bedourie road, 27 June 2010, D.Halford
QM116 (BRI); Elizabeth Springs Conservation Park,

98

Swainsona 31 (2018)

The Ptilotus murrayi species group

24 Oct. 2010, R.Moore 1304 (BRI); Carlo, 16 June 2010,
J.Silcock JLS263 (BRI); 18 km E of Mount Leonard Station
Homestead, 9 June 2010, G.P.Turnpin GPT1333 (BRI).

SOUTH AUSTRALIA: Cordillo Downs on track near
S. boundary c. 1 km S of Bluebush Dam, 6 May 2011,
D.C.Bickerton, P.J.Lang &D.J.Duval 76 (AD).

WESTERN AUSTRALIA: Liveringa Station near Fitzroy
River, 20 Apr. 1985, T.E.HAplin 179 (PERTH); track behind
station to Yanarrie River within 200 m of homestead Yanrey
Station, 6 May 2004, G.Byrne 943 (PERTH); Maitland
Road, Bullgarra Cell, Karratha, 20 Aug. 1986, KGlennon
239 (PERTH); on the side of North West Coastal Highway
on a mound in a recendy burnt flood plain, 2 km W of Yule
River, 17 Aug. 2015, T.Hammer, S.Dillon & KThiele TH25
(PERTH); c. 800 m ENE of Balmoral Caravan Park, Balmoral
Road, Nickol, Karratha, 14 Apr. 2011, V.Long VLK tall-
11 (PERTH); c. 20 km E of Nanutarra Station homestead,
c. 200 km W of Paraburdoo, 15 May 199 7, A.A.Mitchell4735
(PERTH); 8.7 km NNW of Whim Creek, Mallina Station,
Pilbara, 19 Apr. 2006, S.van Leeuwen PBS 5816 (PERTH).

Ptilotus gomphrenoides F.Muell. ex Benth.

FI. Austral. 5: 244 (1870). — Type: ‘Hamersley Range,
N.W. coast, F. Gregory’s Expedition’, F.Mueller s.n.
(syn: K 000357029 image!, MEL 2281815 image!).

Ptilotus roseo-albus Farmar var. roseo-albus. Bull. Herb.
Boissierser. 2, 5: 1090 (1905). — Ptilotus gomphrenoides
var. roseo-albus (Farmar) Beni, Mitt. Bot. Staatssamml.
Miinchen 4: 277 (1962). — Type: ‘N.-W. Division, W
Australia, E. Clement, 1897’ [between the Ashburton
and De Grey rivers, Western Australia, 1897, E. Clement
s.n.] (syn: K 000357024 image!, K 000357025 image!,
K 000357026 image!, K 000357027 image!).

Ptilotus roseo-albus var. conglomeratus Farmar, Bull. Herb.
Boissier ser. 2, 5: 1090 (1905). — Ptilotus gomphrenoides
var. conglomeratus (Farmar) Beni, Mitt. Bot. Staatssamml.
Miinchen 4: 278 (1962). — Type: ‘N.-W. Division, W
Australia, E. Clement, 1897’ [between the Ashburton
and De Grey rivers, Western Australia, 1897, E. Clement
s.n.] (holo: K 000357028 image!).

Erect, decumbent or less-commonly prostrate annual
herb (2-) 10-30 cm high, 35-100 cm wide. Stems
terete, ribbed, glabrous or with sparse nodose hairs.
Basal leaves not seen. Cauline leaves alternate, narrowly
lanceolate to broadly lanceolate, entire, 5-30 mm
long, 1-5 mm wide, glabrous; base narrowly attenuate
or petiolate, 3-10 mm long; apex mucronate, mucro
0.1-0.2 mm long. Inflorescences solitary or in clusters,
axillary or terminal, pedunculate (peduncle 1-10 mm
long) or rarely sessile, spiciform, cylindrical, 4-20 mm
long, 6-7 mm wide, pinkish white; apex acute or
truncate. Bracts ovate, 1.1-1.4 mm long, 0.8-0.9 mm
wide, translucent, glabrous; midrib obscure; apex
rounded. Bracteoles ovate, 1.2-1.5 mm long, 0.8-
0.9 mm wide, translucent, glabrous; midrib faint or
obscure; apex acute. Outer sepals narrowly lanceolate,
2.0—3.0 mm long, 0.5—0.8 mm wide; midrib region
prominent, pink or rarely greenish, 1.2—1.5 mm long,
0.4—0.8 mm wide; basal outer surface with long, wavy,
nodose hairs at base, becoming densely woolly with
age; inner surface glabrous; apex rounded, scarious

Fig. 3. Occurrence of herbarium records. A Ptilotus murrayi; B
P. gomphrenoides (circles) and P. unguiculatus (triangle).

margins 0.8-1.5 mm long, glabrous, white. Inner sepals
lanceolate to broadly lanceolate, 1.8-2.5 mm long, 0.7-
1.2 mm wide; midrib region prominent, pink or rarely
green, 1.0-1.5 mm long, 0.5-0.8 mm wide; basal outer
surface with long, wavy, nodose hairs at base, becoming
densely woolly with age; inner surface glabrous; apex
rounded, scarious margin 0.5-0.8 mm long, glabrous,
pink. Fertile stamens 5; filaments 0.3-0.7 mm long;
anthers 0.2-0.4 mm long, 0.2-0.3 mm wide, yellow
or pink. Staminal cup 0.1-0.2 mm long, unlobed.
Ovary globular, 0.6-0.7 mm long, 0.5-0.7 mm wide,
glabrous. Style straight, 0.2-0.4 mm long, centrally
placed on the ovary apex. Stigma unlobed, capitate.
Fruit smooth, membranous, irregularly dehiscent. Seed
round, black, glossy, c. 0.8 mm long, c. 0.7 mm wide.
Fig. IB.

Diagnostic characters. Ptilotus gomphrenoides can
be distinguished from other species in the genus by
the following combination of characters: an erect or
decumbent annual herb with glabrous stems and leaves,
pedunculate axillary or terminal inflorescences, sepals
not clawed, outer sepals with rounded apices, a glabrous
adaxial sepal surface, densely woolly hairs on the base of
the abaxial sepal surface, 5 fertile stamens, staminal cup

99

T.A. Hammer

Swainsona 31 (2018)

appendages absent, a central style centrally placed on
the ovary apex, and a glabrous ovary.

Phenology. Ptilotus gomphrenoides flowers from April to
September, with a few outlying specimens having been
collected from October to January.

Distribution and habitat. Ptilotus gomphrenoides
occurs in the Carnarvon, Gascoyne, Pilbara and
northern Murchison IBRA regions of Western Australia
(Fig. 3B). The typical habitat is flat seasonally-
inundated floodplains, riverbanks or creek lines with
clayey red or brown soils.

Conservation status. Ptilotus gomphrenoides is not
listed as of conservation concern in Western Australia.

Selected specimens examined

WESTERN AUSTRALIA: Callina Creek c. 40 km WSW
of Marble Bar, 30 Apr. 2006, A.Bean 25214 (PERTH);
Chichester Ranges, 20 July 2011, S.Chalwell 379 (PERTH);
Turn off to Python Pool near Tom Price railway road,
16 July 2013, R.Davis 12261 (PERTH); 1.1 km S along
fenceline from Mount Brockman road and 22 km W of
Hamersley Station homestead, 24 Sep. 2006, D.Halford
Q9265 (PERTH); Nammuldi/Silvergrass lease area near
Mt Brockman, WNW of Tom Price, 28 Aug.-7 Sep. 1998,
M.Maier s.n. (PERTH 05986389); N of Catho Well mesa,
Mount Stuart Station, West Pilbara Iron project area, 7
Aug. 2008, KMcMaster LCH 25882 (PERTH); 14 miles S
of Nullagine, Sep. 1971, RMirrington 710919 (PERTH);
Gascoyne River crossing at Yinnetharra, Western Australia, 14
Aug. 2002, S.Patrick 4291 (PERTH); Karijini National Park,
900 m NNW of the summit of Mt Hyogo and 13.3 km SW
of Mt Bruce and 31.1 km NNE of Mt Bennett, Hamersley
Range, 24 Aug. 1995, S.van Leeuwen 2053 (PERTH); E side
of track, 5.3 km N of Ripon Hill Road on track of abandoned
Braeside Station Homestead, 10.5 km W ofPulgorah Cone,
119.9 km ESE of Marble Bar, Warrawagine Station, Pilbara,
22 Apr. 2004, S.van Leeuwen etal. PBS 7047 (PERTH).

Acknowledgements

The author would like to thank the staff and directors
of the cited herbaria for access to their collections
and loaning specimens. Russell Barrett is thanked for
his input and encouragement, and Robert Davis and
Kevin Thiele are thanked for providing substantial
assistance and giving permission to use photographs.
The author acknowledges the support of a Forrest
Research Foundation PhD scholarship and a University
Postgraduate Award (UWA).

References

Bean, A.R. (2008). A synopsis of Ptilotus (Amaranthaceae) in
eastern Australia. Telopea 12: 227-250.

Beni, G. (1962). Beitrag zu einer Revision der Gattung Ptilotus
R.Br. (Amaranthaceae) 4. Teil. Mitteilungen der Botanischen
Staatssammlung Munchen 4: 277-284.

Beni, G. (1971). Ein Bestimmungsschlussel fur die Gattung
Ptilotus R.Br. (Amaranthaceae). Mitteilungen der Botanischen
Staatssammlung Munch en 9: 135-176.

Beni, G. (1979). Erganzende Bemerkungen zu bisher wenig
bekannten Ptilotus-S ippen (Amaranthaceae) nebst
einigen Neubeschreibungen. Mitteilungen der Botanischen
Staatssammlung Munch en 15: 161-174.

Beni, G. (unpubl.). Ptilotus. Manuscript for: Flora of Australia,
Vol. 5; dated 1988. (Australian Biological Resources Study:
Canberra).

Bentham, G. (1870). Flora Australiensis, Vol. 5. (Reeve:
London).

Black, J.M. (1923). Additions to the flora of South Australia,
No. 21. Transactions and Proceedings of the Royal Society of
South Australia 47: 367-370.

Burbidge, N.T. (unpubl.). Key to species of Ptilotus. Manuscript.
(Australian National Herbarium: Canberra).

Department of the Environment (2018). Australia’s bioregions
(IBRA), IBRA7, Commonwealth of Australia. https://www.
environment.gov.au/land/nrs/science/ibra#ibra [accessed: 8
June 2018].

Farmar, L. (1905). Contributions to our knowledge of Australian
Amaranthaceae. Bulletin de I’Herbier Boissier ser. 2, 5(11):
1085-1091.

Hammer, T.A., Davis, R.W. & Thiele, K.R. (2015). A molecular
framework phylogeny for Ptilotus (Amaranthaceae):
Evidence for the rapid diversification of an arid Australian
genus. Taxon 64(2): 272-285.

Hammer, T.A. & Davis, R.W. (2017). Synonymisation of three
Western Australian taxa within Ptilotus (Amaranthaceae).
Nuytsia 28: 133-134.

Mueller, F.J.H. von (1863). Fragmenta Phytographiae Australiae,
Vol. 3. (Government Printer: Melbourne).

Ronse De Craene, L.P. (2013). Reevaluation of the perianth
and androecium in Caryophyllales: implications for flower
evolution. Plant Systematics and Evolution 299: 1599-1636.

Smith, M. & Jones, A. (2018). Threatened and Priority Flora
list for Western Australia. (Department of Biodiversity,
Conservation and Attractions: Kensington, Western
Australia.)

Vrijdaghs, A., Flores Olvera, H. & Smets, E. (2014). Enigmatic
floral structures in Alternanthera, Iresine, and Tidestromia
(Gomphrenoideae, Amaranthaceae): A developmental
homology assessment. Plant Ecology and Evolution 147:
49-66.

With the exception of images and other material protected by a trademark and subject to review by the Government of
South Australia at all times, the content of this publications is licensed under the Creative Commons Attribution 4.0 Licence
(https://creativecommons.Org/licenses/by/4.0/). All other rights are reserved.

© 2018 Board of the Botanic Gardens and State Herbarium (Adelaide, South Australia)

100

Swainsona 31:101 (2019)

© 2019 Board of the Botanic Gardens & State Herbarium (Adelaide, South Australia) ©0

CORRIGENDUM to: A new species of small black disc fungi, Smardaea
australis (Pezizales, Pyronemataceae), is described from Australia

Pamela 5. Catcheside a - b & David E.A. Catcheside b

3 State Herbarium of South Australia, GPO Box 1047, Adelaide, South Australia 5001
Email: Pam.Catcheside@sa.gov.au

b School of Biological Sciences, Flinders University, PO Box 2100, Adelaide, South Australia 5001
Email: David.Catcheside@flinders.edu.au

The description of the new species Smardaea australis
P.S.Catches. & D.E.A.Catches. (Catcheside et al.
2017) contained one error. Inadvertently, the asci were
described as “amyloid”. This is not correct, the species
has inamyloid asci, as do other species of the genus.

We thank Dr Mario Filippa for alerting us to the
mistake.

References

Catcheside, P.S., Qaraghuli, S. & Catcheside, D.E.A. (2017).
A new species of small black disc fungi, Smardaea australis
(Pezizales, Pyronemataceae), is described from Australia.
Swainsona 31: 17-26.

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Swainsona Volume 31—2017/18

Contents

Leptecophylla in Tasmania: a reassessment of four species

S. J. Jorman & G. Kantvilas .1

A new species of small black disc fungi, Smardaea australis (Pezizales, Pyronemataceae),
is described from Australia

P.S. Catcheside, S. Qaraghuli & D.EA. Catcheside .17

Tephromela baudiniana sp. nov. (lichenised Ascomycetes) from Kangaroo Island

G. Kantvilas &J.A. Elix .27

Two species of Bacidia De Not. with pruinose apothecia from Kangaroo Island

G. Kantvilas . 31

Goodeniaasteriscus (Goodeniaceae), a new arid zone species from north-western South
Australia and eastern Western Australia

PJ. Lang & RJ.-P. Davies . 37

Pertusaria crassilabra Mull. Arg. - a reinstated name for an Australasian lichen

G. Kantvilas ..45

The type of Sturt pea found

AS. George .49

Micarea kartana sp. nov. (lichenised Ascomycetes) from Kangaroo Island, South Australia

G. Kantvilas .55

Notes on the taxonomy of Australian Lindernia subg. Didymadenia (Linderniaceae)

W.R. Barker .59

Antrelloides atroceracea, a new genus and species in the Pezizaceae (Pezizales) from Australia

P.S. Catcheside & D.E.A. Catcheside .81

The Ptilotus murrayi species group: synonymisation of P. petiolatus under P. murrayi and
description of the new Western Australian species P. unguiculatus (Amaranthaceae)

T. A. Hammer .93

CORRIGENDUM to: A new species of small black disc fungi, Smardaea australis (Pezizales,
Pyronemataceae), is described from Australia

PS. Catcheside & D.E.A. Catcheside .101

Front cover: A new species described in this journal issue — Smardaea australis P.S.Catches. & D.E.A.Catches. (p. 19).