Telopea
Journal of Plant Systematics
Vo I 11(1) • 2005
Botanic Gardens Trust
SYDNEY
National Herbarium of New South Wales
Telopea is published by the National Herbarium of New South Wales, Royal
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Website: http://plantnet.rbgsyd.nsw.gov.au/Telopea
Cover Artwork
Telopea speciosh
David Mackay.
Cover Artwork
Telopea speciosissima (Sm.) R. Br., adapted by Helen Stevenson from the original by
Telopea
Journal of Plant Systematics
1 1 ( 1 ): 1-98 * 2005
CONTENTS
Paenula sloryi, a new genus and species related to Ixodia and Haeckeria
(Asteraceae: Gnaphalieae) A.E. Orchard
Morphological and ontogenetic studies on the gynostemium of some
Australian members of Diurideae and Cranichideae (Orchidaceae)
H. Kurzweil, P.H. Weston and A. J. Perkins
A new species of Goodetiia (Goodeniaceae) from Nocoleche Nature
Reserve, Far Western Plains, New South Wales
Belinda J. Pellow and John L. Porter
Reassessment of Indigofera pratensis var. coriacea Domin and var.
angustifoliola Domin (Fabaceae: Faboideae) with the recognition of a
new species Aniuska A. Kazandjian and Peter G. Wilson
Lectotypification of Schoenodum tenax (Restionaceae) and a note on
the type of Lygitiia itnberbis (Anarthriaceae) Barbara G. Briggs
New combinations and synonymies in the Australian Graphidaceae
A.W. Archer
A new species of Pomaderris (Rhamnaceae) from the Central Tablelands
of New South Wales J.C. Millott and K.L. McDougall
A colligate Spirogyra (Zygnemataceae, Zygnematophyceae) in Australia
Stephen Skinner and Timothy J. Entwisle
A new Zygnemopsis species (Zygnemataceae, Zygnematophyceae), with
mature zygospores, from Australia
Stephen Skinner, Hannah McPherson and Gillian M. Towler
Justice for Justice Barron Field Helen Hewson
1-9
11-33
35-41
43-51
53-58
59-78
79-86
87-89
91-93
95-98
Telopea 11 ( 1 ): 1-98 *2005
Scientific Editor
Joy Everett
Editor of Publications
Gary Bridle
Typesetting and Production Assistance
Julia Sideris and Debby McGerty
Other members of Editorial Committee
Elizabeth Brown, Brett Summerell, Peter Wilson
Website: http://plantnet.rbgsyd.nsw.gov.au/Telopea
ISSN 0312-9764
Telopea 10(4) was distributed on 14 December 2004.
National Herbarium of New South Wales
Royal Botanic Gardens Sydney
Mrs Macquaries Rd
Sydney NSW Australia 2000
Telopea 11(1) 1-9
Paenula storyi, a new genus and species
related to Ixodia and Haeckeria
(Asteraceae: Gnaphalieae)
A.E. Orchard
Centre for Plant Biodiversity Research, CSIRO Plant Industry, GPO Box 1600, Canberra
ACT 2601, Australia
Abstract
A plant collected incidentally 40 years ago SSE of Wollar in the Central Western Slopes district of
New South Wales is shown to represent a new genus and species, combining the paleae
characteristics of Ixodia with the leaf characteristics of Haeckeria s.str. and the phyllary
characteristics of Cassinia and Haeckeria. As in Haeckeria, and some species of Odixia and
Cassinia, it lacks a pappus. The genus Paenula and species Paenula storyi are described and
illustrated, the few available details of ecology summarised, and inter-relationships discussed.
Introduction
In an earlier paper (Orchard 1981) 1 discussed character distribution in a range of
genera related to Ixodia ( Odixia , Haeckeria, Ammobium, Cassinia and Ozothamttus
(as Helichrysum)). It was shown that this group of shrubby Asteraceae showed a
braided series of inter-relationships, where characters usually thought to be diagnostic
(presence and absence of pappus, presence and absence of paleae, production of a
showy ‘petal-like’ reflexed tip to the inner phyllaries, etc) seemed to be segregating
independently of each other. That paper resulted in the description of a new genus
Odixia, to accommodate two Tasmanian species formerly placed in Ixodia. Odixia is
separated from Ixodia in having only 5-6 florets per capitulum, membranous
phyllaries, of which the innermost are ±linear with a narrow white tip, and either
lacking paleae between the florets, or paleae flat, linear or filiform. Ixodia has 20-30
florets per capitulum, green ±fleshy phyllaries, of which the innermost arc clawed with
a broad white hooded tip, and the paleae wrap around and envelop the florets. There
were other minor character differences: for example, in Ixodia the abscissing fruits
leave peg-like scars on the receptacle, while in Odixia the scars are rough but not
peg-like. It was suggested that Odixia was probably more closely related to Cassinia
and Haeckeria than to Ixodia. Odixia, Cassinia and Haeckeria share a ±woody shrubby
habit, membranous phyllaries which are not noticeably expanded at the tips into
petaloid appendages, and paleae which, when present, are planar or at most,
longitudinally fluted. Ixodia is a short lived shrub, has green fleshy phyllaries, well
© 2005 Royal Botanic Gardens and Domain Trust
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2
Telopea 11(1): 2005
Orchard
developed showy petaloid terminal appendages on the inner phyllaries, and paleae
which wrap around the achene and base of the floret. Ixodia was last revised by Copley
(1982).
In a separate paper (Orchard 2004) I discuss the variation and limits of Haeckeria,
reducing that genus to just two species, H. punctulata and H. cassiniiformis. Thus
reduced, Haeckeria is defined as comprising short-lived woody shrubs bearing fleshy
trigonous leaves with embedded dark glands, capitula of 1,2 or 3 bisexual florets with
no paleae, florets epappose, and phyllaries membranous or cartilaginous, each
narrowly ovate, slightly hooded, incurved at tip and lacking showy petaloid
appendages.
In the course of preparing the above paper and another on Cassinia, I came across a
specimen from New South Wales which caused me to revisit the Ixodia complex. This
specimen was collected almost accidentally in 1961 by R. Story. He had collected
specimens of Cassinia quinquefaria from south-south-east of Wollar, from an
unnamed ‘nephaline hill*. In addition to flowering material of the Cassinia he collected
some other material that he thought was fruiting material of the same species.
However, during processing at the National Herbarium of New South Wales it was
noted that the fruiting material did not belong to C. quinquefaria , and it was annotated
‘Cassinia sp. nov.’ As the material was rather scrappy, it lay in limbo for the next 40
years.
Taxonomy
Examination of the above material in the course of my Cassinia revision revealed that
it was not part of that genus. The leaves are terete-trigonous, rugose, with an obscure
midrib, and the leaves have, on all three surfaces, deeply sunken dark pit-glands. The
capitula are CYissi/nn-like, with white phyllaries all slightly hooded, incurved at the tip,
spirally arranged, and lacking a showy reflexcd petaloid tip. The florets are all bisexual,
epappose, few in number (6 per capitulum) and each is enclosed in a stiff hyaline
sheath formed by its subtending palea.
In its leaf characters (trigonous with dark sunken glands) this plant most closely
resembles Haeckeria s. str. Some plants of Ixodia can also exhibit dark sunken pit-
glands, but the leaves in that genus are usually flattened, never trigonous in section. In
its capitulum structure the new species mimics Cassinia, and it was this allied with its
± terete leaves which undoubtedly led to its temporary identification as Cassinia sp. nov.
However, similar inflorescence structures and capitulum structures are also found in
Ozothannms and in Haeckeria s.str., and the leaf anatomy of Paenula is not reflected in
any species of Cassinia, where the leaves arc uniformly dorsiventral and never exhibit
dark sunken pit-glands. The interim close identification of this taxon with Cassinia is
thus to be disregarded. In lacking a pappus it also resembles Haeckeria, Odixia and
some anomalous Cassinia species (see Orchard, in prep.), but in its peculiar sheathing
paleae it is obviously linked to Ixodia.
The Story material clearly represents a new species, and it does not fit well in any of the
existing genera. Expansion of the limits of any of them to accommodate it would
introduce unsatisfactory anomalies elsewhere. The most satisfactory solution seems to
be to recognise it as a distinct genus, sister to Ixodia (or perhaps Haeckeria), and
distinguished from its near relatives as follows:
Paenula storyi, a new genus and species
Telopea 11(1): 2005
3
1. Paleae forming a stiff hyaline sheath around the achenes and base of the florets; leaves often
with dark sunken pit-glands.
2. Capitula of 20-30 florets; phyllaries green and subfleshy, leaves decurrent, terete or
flattened but not trigonous. Ixodia
2: Capitula of c. 6 florets; phyllaries white, membranous to cartilaginous; leaves sessile, not
decurrent, trigonous. Paenula
1: Paleae (if present) flat or at most longitudinally fluted; leaves glabrous, pubescent or with
dark sunken pit-glands; phyllaries membranous to cartilaginous
3. Pappus absent
4. Inner phyllaries with spreading petaloid tips; leaves dorsiventral with well
defined midrib, glabrous or pubescent but never with dark sunken pit-glands;
paleae present or absent . Odixia
4: Inner phyllaries lacking spreading petaloid tips
5. Leaves trigonous, lacking prominent midrib below, with dark sunken
pit-glands; paleae absent. Haeckeria
5: Leaves dorsiventral with prominent midrib below, glabrous, cottony
pubescent or scabrous but never with dark sunken pit-glands; paleae
present . Cassinia
3: Pappus of shortly toothed bristles
6. Paleae present (or if absent, florets 1-3 per capitulum); often aromatic shrubs
with a resinous or‘curry’ smell; phyllaries all with incurved tips . Cassinia
6: Paleae absent (florets rarely as few as 3, usually 8-10 or more); usually not
aromatic; phyllaries all with incurved tips or innermost spathulate with spreading
petaloid tips . Ozothamnus
Generic relationships within Asteraceae-Gnaphalieae
The two major papers relevant to how this new genus fits in the Gnaphalieae are
Anderberg (1991) and Bayer et al. (2002).
Anderberg (1991) published a number of trees where of all of the above genera
(obviously excluding Paenula) were in the Cassiniinae. Unfortunately Anderberg
included within his concept of Haeckeria species that are now considered to belong in
Ozothamnus (H. pholidota ), and Cassinia (H. ozothamnoides). This was reflected in his
prophetic statement (p. 87) under Haeckeria that “the genus appears to be
polyphyletic”. Despite this, it is interesting to note that his strict consensus tree for
Cassiniinae (Fig. 12, p. 32) shows Haeckeria + Apalochlamys as sister taxa to Ixodia +
Odixia, and this group of four genera sister to Ozothamnus, and that combined clade
in turn sister to Cassinia. Other analyses omitted Haeckeria and are thus
uninformative. On morphological grounds, the inclusion of Apalochlamys in
Anderberg’s subclade of Apalochlamys, Haeckeria, Ixodia and Odixia seems to me
problematical, but the grouping of the other three genera is credible morphologically.
I would place Paenula in this subclade.
Bayer et al. (2002) provided a different set of trees based on molecular sequence data.
These trees appeared to show that Ixodia and Haeckeria were situated on widely
different branches (branches E and G, Fig. 1, p. 807). However, the Bayer et al. analysis
sequenced only one individual from ‘Haeckeria’, and unfortunately this individual was
4 Telopea 11(1): 2005
Orchard
'g . Paenula storyt, a, habit; b, leaf; c, leaf tip; d, diagrammatic TS of leaf; e, capitulum;
i, phyllanes from outer to inner, dorsal views; j, phyllary, lateral view of (h); k, corolla; 1, style
and stylopodium; m, achene enclosed in palea, axial view; n, same, dorsal view; o, same, lateral
view; p, achene unusually only partly sheathed by palea; q, achene, lateral view; r, achene oblique
apica view, ca es - 1 cm (a) or 1 mm (b—r). All drawings by author, based on Story 7587.
Paenula storyi, a new genus and species
Telopea 11(1): 2005
5
from a taxon that is in fact an undescribed species of Cassinin (Orchard 2004). Thus
the analysis provides molecular support for my segregation of this taxon (on
morphological criteria) as a Cassinia (the Bayer et al. analysis placed it as sister to
Cassinia, represented by a collection of C. longifolia), but does not provide any
indication of the real relationships of Haeckeria to Cassinia or Ixodia, or any indication
of where Paenula might fit in this hypothetical phylogeny.
Paenula Orchard, gen. nov.
Frutices; foliis alternis, teretibus, plusminusve trigonis in sectionis, glandibus inclusis
fuscatis depressis; inflorescentiis cymosis, umbelliformibus; capitulis pedunculatis,
albis, ovoideis; phyllariis membranaceis vel cartilagineis, parum cucullatis, apicibus
petaloideis expansis destitutis; flosculis omnibus bisexualibus, tubularibus; paleis
cartilagineis, hyalinis, tubularibus achenia et basis corollae vaginantibus; pappibus
destitutis; acheniis exutis in paleam vaginantem.
Shrubs; leaves alternate, terete, itrigonous in section, with embedded dark sunken
glands. Inflorescence cymose, umbelliform, with pedunculate capitula. Capitula white,
ovoid; phyllaries membranous to cartilaginous, slightly hooded, lacking petaloid
spreading tips, with undivided stereome; florets all bisexual, tubular. Paleae tough,
hyaline, wrapping around and enclosing achene and base of corolla. Pappus absent.
Carpopodium well-developed. Achenes shed still enclosed in paleae.
Type: P storyi Orchard
Paenula storyi Orchard, sp. nov.
Caules glabri. Folia 20 mm longa, 1 mm lata, rugosa glandibus fuscatis depressis in
omnis paginis, glabra, sessilia; apice acuto, reflexo, non mucronato. Capitula in
pedunculis 2 mm longis. Capitula 3.0-3.5 mm longa, 1.5-1.7 mm lata, ovoidea;
phyllariis spiratim dispositis, cucullatis, glabris. Phyllaria externa ovata, 1.2 mm longa,
0.6 mm lata. Phyllaria interna oblonga, 2.4-2.8 mm longa, 0.7-0.8 mm lata. Flosculi c.
6 per capitulum. Corolla cylindrica, tubo 2.2 mm longo, 0.4 mm lato, vix tumido ad
basim. Achenium purpureo-brunneum, fusiforme, 1.1 mm longum, 0.3 mm latum,
trichomatibus gemellis sparsis appressis tectum.
Type: New South Wales: Central Western Slopes: 18 miles [c. 29 km] north by east of
Rylstone (about 12 miles [c. 19 km] S.S.E. of Wollar) R. Story 7587, 18 Mar 1961,
holotype NSW232395.
Stems glabrous, weakly longitudinally ribbed. Leaves 20 mm long, 1.0 mm wide,
rugose with dark sunken glands on all surfaces, glabrous, sessile, not stem clasping; tip
acute, reflexed, not mucronate. Inflorescence slightly domed. Capitula on peduncles
2.0 mm long; peduncles and inflorescence branches subtended by keeled deltoid bracts
1.0 mm long. Capitula 3.0-3.5 mm long, 1.5-1.7 mm diam., ovoid; phyllaries spirally
arranged, all somewhat hooded, incurved at tip. Outer phyllaries ovate, 1.2 mm long,
0.6 mm wide, glabrous; stereome green, drying brown in lower two thirds; opaque
white in upper one third and on wings. Inner phyllaries oblong, 2.4-2.8 mm long,
0.7-0.8 mm wide, glabrous, with poorly defined green-brown stereome in lower half
to two thirds; upper part and wings white opaque. Paleae cartilaginous; occasional
outer paleae oblong, hyaline, 2.1 mm long, 0.6-0.7 mm wide, longitudinally fluted,
with truncate erose tip; most paleae enveloping base of florets and achenes in a stiff
2-lipped tunic. Florets c. 6 per capitulum, all bisexual. Corolla cylindrical; tube 2.2 mm
6
Telopea 11(1): 2005
Orchard
long, 0.4 mm wide, scarcely swollen at base; corolla lobes deltoid, 0.4 mm long,
reflexed, glabrous. Anthers 1.25 mm long inch 0.13 mm deltoid appendage; anther tails
filamentous, 0.25 mm long. Style 2.0 mm overall, divided into 2 branches in upper 0.9 mm;
stylopodium transparent, conical, 0.25 mm long, very minutely rugulose. Style arms
truncate, with well developed brush of clavate hairs. Achene purple-brown, fusiform,
1.1 mm long, 0.3 mm diam., with sparse appressed twin hairs for entire length, and a
denser ring of hairs apically.
Distribution: New South Wales, Central Western Slopes. Known only from the type
specimen, collected SSE of Wollar.
Ecology: Found on a nephaline hill, with Cassinia quinquefaria (Story 7577) and
Eucalyptus albens dominant. Other material collected at the same time and place
comprised Eucalyptus goniocalyx (Story 7574), Eucalyptus laevopinea (Story 7575),
Grevillea ramosissima subsp. ramosissima (Story 7576), Eucalyptus blakelyi (Story 757S)
Eucalyptus dawsonii (Story 7579) and Muelleriua bidwillii (Story 7580). Fruits were
present in mid-March. At or immediately after seed shed, capitula abscise below the
phyllaries, leaving the small subtending bracts and peduncles still attached to the
twiggy remains of the inflorescence.
Etymology: The generic name paenula is a word used by Cicero to describe a coat
without sleeves and close to the body, worn on a journey, i.e. a kind of close-fitting
cloak. It alludes to the distinctive tightly cloaked fruits of this taxon. The species
epithet commemorates the original discoverer, Robert Story, ecologist (1913-1999).
Notes
It may be argued that this plant really represents an Haeckeria which has retained its
paleae. After all, in Haeckeria punctulata and H. cassiniiformis, the character of the
missing paleae can not be determined, except by analogy. Wakefield (1951) and
Orchard (2004) have noted that in Cassinia, which differs from Haeckeria in usually
having a pappus and paleae, the paleae are apparently absent in capitula with only 1, 2
or 3 florets. In these inflorescences it is argued that the paleae (situated on the abaxial
side of the florets) are adjacent to and indistinguishable from the inner phyllaries. Only
in capitula with more than 3 florets will paleae be interposed between adjacent florets.
By analogy, if the few-flowered capitula of Haeckeria have paleae, then they must be
similarly indistinguishable from the inner phyllaries and thus ±flat. Therefore in
Haeckeria the paleae are either completely absent (as most believe), or ±flat and
resembling the inner phyllaries. In either case, the situation differs from that in Paenula
where the paleae are sheathing.
Paenula storyi was searched for unsuccessfully in February 2004 near its original place
of discovery, in the vicinity of Growee Gulph, and again, unsuccessfully, in early March
2004 on Barigan Road, SSE of Wollar. The latter seems the most likely source of Story’s
original collection the respective reference distances from Rylstone and Wollar turn
out to miss each other by about 10 kilometres. At about the correct distance (12 miles)
rom Wollar there is in fact a small hill composed largely of basaltic boulders, on the
property of Mr John Jakes, “Derowen”. This hill bears a remnant scrub of Eucalyptus
albens, Cassinia quinquefaria and Bursaria spinosa on its summit, with occasional
egumes and herbaceous Asteraceae, but there was no sign of Paenula. A few kilometres
Paenula storyi, a new genus and species
Telopea 11(1): 2005
7
Fig. 2. Distribution of Paenula storyi, New South Wales.
8
Telopea 11(1): 2005
Orchard
further along the same valley there is a volcanic plug, which now has only a grassy
covering. All other hills in the near vicinity seemed to be mainly sandstone. The upper
slopes of these (generally larger) hills are covered with a dense forest of Eucalyptus spp.,
Callitris,Xanthorrhoea and Cassinia quinquefaria. No material resembling Paenula was
discovered in a vegetation survey of the adjacent Merriwa area by McRae & Cooper (1985).
This lack of success in rediscovering Paenula may signify nothing more than that the
plant is short-lived. Reference has already been made to the morphological similarity
between Paenula and the two Haeckeria species. Several collections of Haeckeria bear
notes on their ephemeral nature. Haeckeria species may appear in abundance after
major disturbance such as fire or ‘chaining’ of the mallee scrub in which they grow.
However they are usually senescent in 4-5 years and disappear completely within 10
years, awaiting the next disturbance. It is very possible that Paenula is behaving
similarly and will be re-discovered in the Wollar/Rylstone area after a future
disturbance.
It has been suggested by readers of earlier drafts of this paper that Paenula storyi might
also be explained as a short-lived hybrid or polyploid, but those making this suggestion
offer no suggestions as to parent taxa or (diploid) ancestor. However, the possibility is
worth exploring. My morphological analysis suggests that the two most diagnostic
characteristics of this taxon are its cloaking paleae (in the Cassiniinae shared only with
Ixodia, the nearest representatives of which are currently found 1000 km to the west of
Paenula), and the trigonous leaves with sunken pit-glands (in the Cassiniinae shared
only with Haeckeria s. str., the nearest representatives of which are currently found 900
km south-west of Paenula). The only Cassiniinae found near the location of Paenula
are Ozothamnus and Cassinia species, none of which exhibit either of these character
states. While this does not exclude the possibility of Paenula having arisen through a
hybridisation or polyploidy event in the distant geological past, when Haeckeria and
Ixodia may have been present in NSW, it begs the question of how long a self-
perpetuating taxon needs to exist before it can be recognised as a genus in its own
right? I believe that a recent hybrid or polyploidy origin for Paenula is extremely unlikely.
Acknowledgments
The author is grateful to his wife Theresa, patient and observant field assistant on
numerous field trips in search of Cassinia and related taxa, and who searched the AVH
database for other Story collections from the type locality. Laurie Adams supplied
background information on Robert Story. The Centre for Plant Biodiversity Research,
CSIRO, Canberra, kindly provided excellent institutional support for this study and
related fieldwork. Brendan Lepschi patiently joined me in the search for Paenula SSE
of Wollar. The author is particularly grateful to John Jakes, owner of‘Derowen’, for
hospitality and advice during the unsuccessful search on his property. 1 am also
grateful to Barr)' Conn, Karen Wilson and John Benson of the National Herbarium of
NSW, for background information on Story’s collections and survey of 1961. Finally,
1 am very grateful to the National Herbarium of NSW for including this interesting
material in my loan of Cassinia.
Paenula storyi, a new genus and species
Telopea 11(1): 2005
9
References
Anderberg AA (1991) Taxonomy and phylogeny of the tribe Gnaphalieae (Asteraceae), Opera
Botanica 104: 5-195.
Bayer R), Greber DG & Bagnall NH (2002) Phylogeny of Australian Gnaphalieae (Asteraceae)
based on chloroplast and nuclear sequences, the trnL Intron, trnL/trnF Intergenic Spacer, matK,
and ETS. Systematic Botany 27(4): 801-814.
Copley PB (1982) A taxonomic revision of the genus Ixodia (Asteraceae). Journal of the Adelaide
Botanic Gardens 6:41-54.
McRae RHD & Cooper MG (1985) Vegetation of the Merriwa area, New South Wales.
Cunninghatnia 1(3): 351-369.
Orchard AE (1981) The generic limits of Ixodia R.Br. ex Ait. (Compositae - Inuleae). Brunonia
4: 185-197.
Orchard AE (2004) A reassessment of the genus Haeckeria (Asteraceae - Gnaphalieae), with
definition of a new species in Cassinia. Australian Systematic Botany 17: 447-467.
Wakefield NA (1951) Some notes on Cassinia with description of a new species. Victorian
Naturalist 68: 69-70.
Manuscript received 19 May 2004, accepted 11 July 2005
Telopea 11(1): 11-33
Morphological and ontogenetic studies on the
gynostemium of some Australian members of
Diurideae and Cranichideae (Orchidaceae)
H. Kurzweil 1 , P.H. Weston 2 and A.J. Perkins 2
'Compton Herbarium, South African National Biodiversity Institute, 7735, Claremont,
South Africa. To whom correspondence regarding reprints should be addressed.
2 National Herbarium of New South Wales, Royal Botanic Gardens Sydney, Mrs. Macquarie’s
Road, Sydney, NSW 2000, Australia.
Abstract
Gynostemium development of Genoplesium fimbriatum (Orchidaceae—Diurideae) is described
and documented by means of SEM micrographs. Other Australian representatives of tribes
Diurideae and Cranichideae are briefly treated. Particular attention is paid to the early stages which
are essential for the correct interpretation of the gynostemium structure. It was found that
gynostemium development largely conforms to that of the other Orchidaceae that have been
examined in this respect. Several of the species have auricles (filament appendages) next to the
anther and a shallowly or prominently three-lobed median stigma lobe in middle ontogeny which
are both interesting similarities shared with tribes Orchideae and Diseae. As in the latter two tribes,
the mature lateral gynostemium appendages (column-wings’) of some of the study species are
of dual origin, comprising both a staminodial and an auricular element.
Introduction
The available literature on the Australian terrestrial orchids consists mainly of floristic,
taxonomic and phylogenetic studies. Although a limited amount of information on the
flower structure of many species can be obtained in the existing literature, little
detailed work on floral morphology has been carried out on a broad scale. A notable
exception is the careful study of Rasmussen (1982) who described gynostemium
structure and development of a few Australian terrestrial orchids in great detail. Very
recently detailed information on the flower morphology and ontogeny of Calochilus
has been made available (Perkins 2001).
In the present paper gynostemium development in Genoplesium fimbriatum
(= Prasophyllum fimbriatum) is described in detail and that of several other Australian
terrestrial orchids is more briefly characterised. Ontogenetic information on a few
more superficially studied species is summarised in a table. Development was found to
be basically similar to that described in other orchids where it has been more fully
described (Kurzweil 1987a—b). The species examined here represent most subtribes of
© 2005 Royal Botanic Gardens and Domain Trust
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Telopea 11(1): 2005
Kurzweil, Weston and Perkins
tribe Diurideae and subtribe Pterostylidinae of tribe Cranichideae (all taxa sensu
Pridgeon et al. 2001,2003; see Table 1). Special attention is paid to the early and middle
development of the gynostemium — here defined as comprising the ontogeny before
the stage where the column-part (if present) starts elongating, or up to the stage where
the stigma starts becoming papillose. An understanding of this particular phase is
essential for a correct interpretation of the adult gynostemium architecture. While the
examination of a few selected species is obviously insufficient as a character survey in
large groups, the present paper is intended as a preliminary analysis and is furthermore
aimed at stimulating future research. It is hoped that comparative investigations of a
large number of species will be undertaken one day, with the aim of improving our
understanding of the fascinating flowers of these orchids and thereby contributing
characters for phylogenetic analysis. The results presented here reveal similarities
shared by at least some members of all tribes of Orchidoideae that have been examined
for floral development. Interestingly, these have a similar phylogenetic distribution to
some other morphological features (e.g. root tubers; Dressier 1981, 1993). Recent
molecular studies (e.g. Cameron et al. 1999; Kores et al. 2000, 2001; Clements et al.
2002) provide a robust phylogenetic framework within which the evolution of these
features can be interpreted.
Ontogeny is probably the best criterion for recognising primary homology (de Pinna
1991) and frequently a reasonable homologisation of organs is not possible without a
sound knowledge of their development. Ontogenetic studies have also proven useful in
botany and zoology in sometimes allowing the relative generality of homologous
character states to be observed directly (Weston 1988, 1994). Furthermore, ontogeny
can also contribute valuable taxonomically significant characters. We expect that our
knowledge of the phytogeny and morphological evolution of the Australian terrestrial
orchids will benefit greatly from a better understanding of its floral and especially
gynostemium structure and development.
Material and methods
Most of the plant material was collected directly in the field in New South Wales
(Australia) by the second author. It was preserved in FAA (ethanol 70% : glacial acetic
acid : formaldehyde = 18:1:1) and subsequently transferred to 70% ethanol.
The exact sources of the material are given in the appendix.
For the present SEM investigations the material was prepared in ethanol 70% under a
dissecting microscope. Samples were chemically dehydrated in FDA (= formaldehyde-
dimethylacetal) and subsequently critical-point-dried directly from FDA without the
use of an intermedium, using CO2 as the carrier gas (technique after Gerstberger
& Leins 1978). The dry samples were coated with AuPd in a ‘BALZERS’ sputter-coater
and viewed and photographed in a CAMBRIDGE STEREOSCAN S200 scanning
electron microscope at 10 kV.
The terminology used here follows that published in earlier papers of the first author
(particularly in Kurzweil 1987a—b).
Australian members of Diurideae and Cranichideae
Telopea 11(1): 2005
13
Table 1. Systematic distribution of the species studied among the Australian representatives of
Diurideae and Cranichideae, and the extent of our investigation.
The classification follows Pridgeon et al. (2001, 2003): early ontogeny ... up to the three-carpel-apex
stage; middle ontogeny ...up to the start of the elongation of the column-part (if present) or up to the
stage where the stigma starts becoming papillose; late ontogeny ... up to anthesis. The figure gives the
number of different stages observed.
Taxon
TRIBE DIURIDEAE
Subtribe Acianthinae
Total
Ontogeny
early
middle lat
Acanthus fornicatus
12
3
3 6
Corybas fimbriatus
Subtribe Caladeniinae
8
-
1 7
Caladenia carnea
1
-
1
C. catenata
1
-
1
Eriochilus autumnalis
4
-
4
Glossodia minor
Subtribe Cryptostylidinae
4
-
4
Cryptostylis erecta
Subtribe Diuridinae
20
7
9 4
Diuris longifolia
12
1
7 4
Orthoceras strictum
Subtribe Drakaeinae
22
■
19 3
Caleana major
16
6
1 9
Chiloglottis sp.
Subtribe Megastylidinae
4
"
4
Lyperanthus suaveolens
2
-
2
Rimacola elliptica
Subtribe Prasophyllinae
13
5
5 3
Microtis parviflora
8
1
4 3
Genoplesium fimbriatum
Subtribe Rhizanthellinae
Subtribe Thelymitrinae
31
20 11
Calochilus campestris
14
6
4 4
C, roberstsonii
4
-
4
Thelymitra carnea
TRIBE CRANICHIDEAE
Subtribe Goodyerinae
Subtribe Pterostylidinae
13
4 9
Pterostylis concinna
15
6 9
Subtribe Spiranthinae
14
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Kurzweil, Weston and Perkins
Observations
Table 2. Various features observed in the species studied:
Ic ... lateral carpel apices/stigma lobes; me ... median carpel apex/stigma lobe.
The ontogenetic stages correspond to those defined in Table 1. x = character present, (x) = character
obscure/weakly developed, - = character not present, ? = not clear, 0 = stages not observed, sep =
separate, con = connate to a ridge, emarg = connate to an emarginate ridge.
Taxon
(in brackets numbers of
illustrations in the present paper)
Acanthus fornicatus (Fig. 7A-D)
Caladenia carnea
C. catenata (Fig. 5D)
Caleana major (Fig. 3D-F)
Calochilus campestris (Fig. 6)
C. robertsonii
Chiloglottis sp. (Fig. 5E)
Corybas fimbriatus (Fig. 5F-H)
Cryptostyiis erecta (Fig. 4A-B)
Diuris longifolia (Fig. 4C-E)
Eriochilus autumnalis
Genopiesium fimbriatum
(Fig. 1A-H, 2A-C)
Glossodia minor
Lyperanthus suaveolens (Fig. 51)
Microtis parviflora
Orthoceras strictum (Fig. 4F-I)
Pterostylis condnna (Fig. 7E-I)
Rimacola elliptica (Fig. 3A-C)
Thelymitra carnea (Fig. 5A-C)
Staminodes
Auricles
Ic
me
(middle stage)
early/
middle late
middle late
middle late
X
X
-
-
con
con
clearly 3-lobed
0
X
0
0
0
emarg
0
0
X
0
0
0
emarg
0
X
X
-
X?
0
emarg
obscurely 3-lobed
X
X
(X)
X
con
con
clearly 3-lobed
0
0
0
X
0
0
0
0
X
-
-
0
0
0
0
X
-
-
0
emarg
clearly 3-lobed
X
X
-
-
sep
emarg
unlobed
X
X
-
?
con
con
unlobed
0
?
0
-
0
emarg
0
X
X
X
X
sep
emarg
obscurely 3-lobed
0
X
0
-
0
0
0
0
X
0
X
0
emarg
0
X
X
-
-
0
con
obscurely 3-lobed
X
X
(X)
?
sep
emarg
obscurely 3-lobed
X
X
(X)
(X)
sep
sep
obscurely 3-lobed
X
X
(X)
X
emarg
emarg
unlobed
X
X
(X)
0
emarg
0
unlobed
Genopiesium fimbriatum (R. Br.) D.L. Jones & M.A. Clem.
Commonly called ‘Midge Orchid’, the genus Genopiesium (tribe Diurideae) comprises
about forty species in open situations in sclerophyllous forest and heathland in
Australia (particularly in the south-eastern areas). New Zealand and New Caledonia.
Vegetative as well as floral morphological features mark affinities to Prasophyllum in
which genus Genopiesium was included in the past. All species are sympodial herbs
with root tubers and have a single terete and hollow leaf (Jones 2001). The
inflorescence bursts through the leaf at a predetermined point near its tip. Flowers are
non-resupinate and have a mobile lip. Pollination is mainly carried out by small flies
of the superfamily Chloropoidea, which may be attracted by nectar, fruity perfumes or
mobile lips (Bower 2001). A few species are self-pollinating. Molecular phylogenetic
analyses of small samples of species (Kores et al. 2001; Clements et al. 2002) suggest
that Genopiesium s.l. is paraphyletic to Prasophyllum s.s.. Jones et al. (2002) transferred
G. fimbriatum and 45 other species to the genus Corunastylis on the basis of these
results. However, we prefer to retain the existing taxonomy until more species in the
Prasophyllinae have been phylogenetically analysed and relationships are more
clearly resolved.
Australian members of Diurideae and Cranichideae
Telopea 11(1): 2005
15
Fig. 1. Genoplesium fimbriatum ; gynostemium development. A, D-F, Front views of the
gynostemium in different stages. B-C, Base of gynostemium and whole gynostemium in side view.
G, Stigma in a late stage seen from the front. H, Adult anther-stigma-complex, lateral gynostemium
appendages (column-wings) largely removed. — SEM micrographs. Bars: 0.1 mm. Abbreviations:
A = anther, a = staminodes, c = carpel apices, h = hamulus, la = lateral gynostemium appendages,
me -- median carpel apex, stg = stigma. The arrowheads mark suspected auricles (= filament
appendages; see Kurzweil 1987b). Source: Weston 1277.
16
Telopea 11(1): 2005
Kurzweil, Weston and Perkins
Genoplesium fimbriatum, the ‘Fringed Midge Orchid’, is endemic to south-eastern
Queensland and the coastal areas of New South Wales where it is rather common in
suitable habitats. Its flowers measure about 9-11 mm across and are prominently hairy.
Sepals and petals are yellowish green with red stripes and the lip is generally pale red.
The ovate median sepal is hooded and has long marginal cilia, and the divergent or
spreading lateral sepals are linear to lanceolate in shape and have entire margins. Cilia
are also present on the narrowly ovate or ovate and pointed petals. A characteristic
feature of the species is the linear lip which is hinged on a short strap and has many
long pink or red marginal cilia which tremble in the wind. It is conspicuously recurved
at its apex and has a long median callus consisting of two narrow ridges. The stout
gynostemium (Fig. 1H) lacks a long column-part and has a pronounced basal column-
foot. Its most prominent organs are the erect column-wings which are approximately
as tall as the anther. They are deeply two-lobed and basally fused to the anther-stigma-
complex. The erect anther is comparatively narrow, measuring about one third of the
gynostemium. An elongate, apical connective process is visible on top of the anther.
The entire stigma is papillose and funnel-shaped.
In the earliest stage available (Fig. 1A) the anther is erect and its division into two
thecae is made visible by a prominent gap between them which is especially
pronounced in its lower part. All three carpel apices have already emerged with the
median one being the largest. The latter is an erect and entire lobe in front of the
anther. The two lateral carpel apices are separate structures in front of the median
carpel apex. Prominent bulges are visible at the inner base of the petals, i.e. in a
position superposed to them (a in Fig. 1A). On account of their position and their
early initiation they are here interpreted as vestiges of the staminodes aj and 33 which
corresponds well with the situation in other groups of monandrous Orchidaceae
(Kurzweil 1985, 1987a). They will be simply referred to as ‘staminodes’ below.
Fig. 2. Genoplesium fimbriatum ; gynostemium in different developmental stages with the anther
removed, showing the median carpel apex, the staminodes and the suspected auricles. A-B, side
views; C, dorsal view. — SEM micrographs. Bars: 0.1 mm. Abbreviations: a = staminodes,
h = hamulus, me = median carpel apex. The arrowheads mark suspected auricles (= filament
appendages; see Kurzweil 1987b). Source: Weston 1277.
Australian members of Diurideae and Cranichideae
Telopea 11(1): 2005
17
These staminodes later grow into the anterior lobes of the prominent two-lobed wings
next to the anther (Fig. 1D-H). In slightly later stages additional structures appear as
small but clearly visible bulges, emerging in a dorsolateral position on the anther
(arrowheads in Fig. 1B-C). They obviously do not occupy a position superposed to the
petals and are thus distinct from the staminodes. They are also not simply posterior
outgrowths of the staminodes which one would expect to develop late in ontogeny. It
is here suggested that the dorsolateral structures in question are appendages of the
filament of the anther, constituting auricles comparable to those in Orchideae
(Kurzweil 1987b). The further development of these auricles is also visible in
gynostemia where the anther has been removed (arrowheads in Fig. 2A-C). Later in
ontogeny they become basally fused with the staminodes and grow up together with
them while their common base elongates. The prominent lateral gynostemium
appendages (‘column-wings’ in taxonomic and floristic treatments) are thus of dual
origin as in some other Orchidaceae (see discussion), being derived from both lateral
staminodes of the inner staminal whorl and filament appendages of the fertile anther
(termed auricles). Rather early, the median carpel apex develops into an obscurely
three-lobed organ (Fig. 1D-G). The lateral carpel apices become fused at their base
and the resulting ridge-like structure remains significantly emarginate. Its derivation
from two separate structures is thus clearly visible.
In middle and late ontogenetic stages the staminodial (= anterior) portions of the
lateral gynostemium appendages become pointed and minutely lacerate (Fig. IF, 2C).
The posterior auricular portions remain rounded and oblong lobes and their surface
remains smooth throughout ontogeny. In middle and late stages they are slightly
shorter than the staminodial portions (Fig. 2C). The three carpel apices form an oval
pad with a flat front face in late development (Fig. 1G), and the median carpel apex is
by far the largest of the three carpel apices. It is a long erect tongue-like lobe. As in
other monandrous orchids the rostellum of the mature gynostemium is entirely
derived from the upper part of this organ. Its apical portion develops into a prominent
hamulus-type stipe (,sensu Rasmussen 1982) (Fig. 1H, 2A-C). The lower portion of the
median carpel apex and the two lateral carpel apices become conspicuously papillose
(Fig. 1G). They eventually form the receptive stigma surface and are thus referred to as
the stigma lobes. In late stages the lower portion of the anterior part of the funnel-rim
which is derived from the lateral stigma lobes grows forward (Fig. 1H). The prominent
elongate apical connective process of the anther develops in late ontogenetic stages.
Notes on other species
Rimacoln elliptica (R. Br.) Rupp
Rimacola elliptica, the sole species of its genus, is endemic to the Sydney region,
including the Blue Mountains (New South Wales). It is most closely related to the
monotypic New Zealand genus Waireia (Clements et al. 2002), which, together, were
originally included in the rather more distantly related genus Lyperanthus (Kores et al.
2001). The arching inflorescences bear a few resupinate flowers with narrowly
lanceolate sepals, falcate petals, and an unlobed or obscurely three-lobed lip with a
basal callus. The mature gynostemium has a long column-part and prominent
column-wings are present. The apex of the anther is broadly rounded in the early and
middle developmental stages but exhibits a short subacute connective process in late
stages (Fig. 3C). Prominent staminode primordia are visible in early stages
18
Telopea 11(1): 2005
Kurzweil, Weston and Perkins
(a in Fig. 3A) and later develop into the obscure column-wings (a in Fig. 3C). In
addition, obscure bulges probably homologous with auricles are developed in middle
stages (arrowhead in Fig. 3A) and grow into the posterior portion of the shallowly two-
lobed lateral gynostemium appendages that are visible in the late stages (arrowhead in
Fig. 3B). In the mature flower the suspected auricles are visible as small dorsolateral
teeth next to the anther (arrowhead in Fig. 3C). The median carpel apex is unlobed in
all stages observed (not shown). The initiation of the lateral carpel apices was not
observed but a slightly later stage reveals that they are partly fused to an emarginate
ridge (not shown).
Caleana major R. Br.
Caleana is a small genus in eastern Australia and New Zealand. C. major has erect
inflorescences with a few non-resupinate flowers. The median sepal is hood-like while
the lateral sepals are linear and reflexed. A most conspicuous part of the flower is the
duck’s head-shaped lip. The gynostemium is very prominent with its wide column-
Fig. 3. Gynostemium development of various species. A-C, Rimacola elliptica, sidc/dorsal views;
D-F, Caleana major, front views. — SEM micrographs. Bars: 0.1 mm in A-B, D-E; 1 mm in C, F.
Abbreviations: A = anther, a = staminodes, me = median carpel apex, stg = stigma. The arrowheads
mark suspected auricles (= filament appendages; see Kurzweil 1987b). Sources: a-b: Weston 1586;
c: Bishop J67/31-37; d-f: Weston 1229.
Australian members of Diurideae and Cranichideae
Telopea 11(1): 2005
19
wings," ... so broad that they form an inverted cup-like basket” (Bernhardt 1993: 194).
In the mature flower these column-wings extend over the full length of the column-
part while in other orchids marked column-wings are confined to the upper portion
of the column-part (for example in Ritnacola elliptica). In early ontogeny staminode
primordia are clearly visible (a in Fig. 3D), and later develop into the prominent
column-wings of the adult gynostemium (Fig. 3F). The anther is somewhat narrow
and pointed in e-arly and middle ontogeny (Fig. 3E). No prominent additional lateral
or dorsolateral gynostemium appendages are visible in early or middle ontogenetic
stages. However, an obscure tooth on the side of the gynostemium is visible in a late
stage (illustrated in Kurzweil 1998, Fig. 11D). The present investigations are
inconclusive with regard to the ontogenetic derivation of the tooth, but the presence
of staminodial column-wings and dorsolateral gynostemium teeth may suggest a dual
origin of the lateral gynostemium appendages from staminodes and auricles like in
many other orchids. The median carpel apex is obscurely three-lobed in middle and
late stages (not shown).
Cryptostylis erecta R. Br.
Cryptostylis is a primarily Australasian genus of about twenty species. The erect
inflorescences bear a few non-resupinate flowers with linear sepals and petals and a
large ovate-oblong lip. The gynostemium development of C. erecta (endemic to
Queensland, New South Wales and Victoria) has been briefly examined. In middle
stages prominent staminode primordia occur (Fig. 4A). Structures likely to be auricles
were not observed in any stage. In late stages the lateral appendages are several-toothed
lobes (Fig. 4B), and their lobed appearance may suggest that they incorporate auricular
tissue as in many other orchids. The median carpel apex is a large erect, entire lobe and
is strongly bulging to the front (Fig. 4A). The stigma is an erect pad with the papillose
area derived from all three stigma lobes. Lateral carpel apices are separate structures at
their time of initiation (not shown).
Diuris lottgifolia R. Br.
The genus Diuris (± 55 species, Jones 2001) is amongst the most attractive of the
Australian terrestrial orchids with their fairly large resupinate flowers. The slender
lateral sepals are mostly reflexed while the median sepal is developed as a shallow hood.
Most prominent are the ear-like clawed petals. The gynostemium lacks a basal column-
part as anther, style and column-wings are free-standing structures and are only fused
at their base. Only a few developmental stages of the Western Australian D. longifolia
were available for study. The large erect lateral appendages of the mature gynostemium
arise from prominent staminode primordia that are visible in early ontogeny (Fig. 4C).
Auricles are not visible in any early or middle stage. However, the lateral appendages of
the gynostemium are shallowly two-lobed in late stages which may point to a dual
origin of the lateral appendages from staminodes and auricles. The median carpel apex
is unlobed in all stages (Fig. 4C-E). In middle stages it is somewhat infolded in its
central portion which grows into the gap between the two thecae (Fig. 4D), and in late
ontogeny this central portion forms the viscidium. The lateral carpel apices emerge
connate as an undivided edge (Fig. 4C).
20
Telopea 11(1): 2005
Kurzweil, Weston and Perkins
Fig. 4. Gynostemium development of various species. A-B, Cryptostylis erecta, side views;
C-E, Diuris longifolia, front views; F-I, Orthoceras strictum. F-H, front views; H, three-quarter
view; 1, adult gynostemium in side view. — SEM micrographs. Bars: 0.1 mm in A-D, F-H; 1 mm
in E, 1. Abbreviations: A = anther, a = staminodes, c = carpel apices, me = median carpel apex, stg
= stigma. The arrowheads mark suspected auricles (= filament appendages; see Kurzweil 1987b).
Sources: A-B: Weston 2473; C-E: sine collector; F, H: Weston 1584; G: Weston 1271; I: Weston 1279.
Australian members of Diurideae and Cranichideae
Telopea 11(1): 2005
21
Fig. 5. Gynostemium development of various species. For reference see Table 2. A-C, Thelymitm
camea. A, front view; B, dorsal view; C, side view; D, Caladenia catenateu front view; E, Chiloglottis
sp„ front view; F-H, Corybas fimbriatus. F-G. side views; H, front view; — i, Lyperanthus
suaveolens, dorsal view. — SEM micrographs. Bars: 0.1 mm in A-C, E-H; 1 mm in D, 1.
Abbreviations: A = anther, a = staminodes, lc = lateral carpel apices, me = median carpel apex,
stg = stigma. The arrowheads mark suspected auricles (- filament appendages; see Kurzweil
1987b). Sources: A: Weston 1227 ; B-C: Weston 1232; D: Abell 67; E: Weston 1278; F-H: Weston 1392;
I: Weston 1224.
22
Telopea 11(1): 2005
Kurzweil, Weston and Perkins
Orthoceras strictum R. Br.
The small genus Orthoceras shares many floral features with Diuris and is therefore
placed in subtribe Diuridinae (Dressier 1993), differing from the latter genus most
obviously in its horn-like and spreading lateral sepals and minute petals. O. strictum,
from Australia, New Zealand and New Caledonia has erect inflorescences with a few
small resupinate flowers. Its gynostemium ontogeny has been examined. Staminode
primordia can be seen in early and middle stages (Fig. 4F). Staminodes are prominent
throughout ontogeny and develop directly into the large tooth-like lateral
gynostemium appendages which are visible in late stages and in the mature
gynostemium (Fig. 4FI-I). An obscure dorsolateral tooth reminiscent of an auricle was
observed in one gynostemium in a middle stage (illustrated in Kurzweil 1998,
Fig. 1 IF). Flowever, this bulge does not appear to be a constant feature as many
gynostemia without it were also observed. An obscure dorsal process is also visible in
the adult gynostemium at the base of the large staminodes (arrowhead in Fig. 41), and
it is possible that it is derived from such an auricle. In addition, there is also a ventral
bulge at the base of the staminodes in late stages and in the mature flower (? in Fig. 41).
Due to its very late ontogenetic origin it is most probably only a secondary outgrowth
of the staminodes. The median carpel apex is obscurely three-lobed in middle and late
ontogeny (Fig. 4G-H). Lateral carpel apices emerge as separate structures but soon
become connate into an emarginate ridge.
Calochilus campestris R. Br.
Calochilus is a genus of approximately 18 species (Jones 1988; Jones & Lavarack 1989;
Jones & Gray 2002; Jones & Clements 2004). All species have a single fleshy leaf up to
40 cm long. Fertile plants produce a raceme 20 to 100 cm tall bearing two to sixteen
resupinate flowers. Most species have multicellular hair-like processes that cover the
surface and margins of the obscurely three-lobed labellum; these are usually green, red,
crimson or purple in colour. These floral features give rise to their common name
‘Beardies’ or ‘Bearded Orchids’, and their scientific name 'Calos’ meaning beautiful,
and ‘cheilos meaning lip (Jones, 1988).
In C. campestris, the mature gynostemium is short and arched forward. Column-wings
are present but obscure and fused to the anther filament to form a mitra or hood,
which dorsally covers the gynostemium. The apex of the anther is narrowly elongate
with a terminal, subacute connective process visible in the early and middle
developmental stages (Fig. 6A), becoming obtuse and densely papillate in late
developmental stages (Fig. 6B). Staminode primordia are clearly visible in early stages
and are already connate, forming a mitra (Mi in Fig. 6A, C). The mitra later grows up
to the base of the anther (Mi in Fig. 6C, D, F). Mature apices of the mitra are covered
with tuberculate appendages and are likely to be homologous with auricles (arrowhead
in Fig. 6C-D). During the middle stages of development the auricles become heavily
sculptured (arrowhead in Fig. 6E-F). During late developmental stages, two orbicular
bulges form on the ventral surface of the mitra, on both sides of the base of the stigma
(g in Fig. 6F). Their anatomical homology and function are unknown but they form
the prominent gland-like eyes’ of the mature flower. The median carpel apex is
strongly elongate (me in Fig. 6A, F) and connate to the lateral carpel apices, appearing
three-lobed in all stages observed. In mature flowers, a viscidium forms on the ventral
surface of the median carpel apex (me in Fig. 6F).
Australian members of Diurideae and Cranichideae
Telopea 11(1): 2005
23
Fig. 6. Gynostemium of Calochilus campestris in different developmental stages. A, during early
development in front view; B, papillae on connective apex during late development; C, during early
development in dorsal view; D, during middle development in dorsal view; E, auricles during late
development in front view; F, during anthesis in front view. — SEM micrographs. Bars: 1 mm in
A, C-F; 0.1 mm in B. Abbreviations: A = anther, Mi = mitra, me = median carpel apex, g = gland¬
like “eyes” on mitra, stg = stigma. The arrowheads mark suspected auricles (= filament appendages;
see Kurzweil 1987b). Sources: NSW446115.
24
Telopea 11(1): 2005
Kurzweil, Weston and Perkins
Acianthus fortiicatus R. Br.
The genus Acianthus, as re-circumscribed by Jones et al. (2002), includes eight species
in Australia and New Zealand. A. fornicatus from New South Wales and Queensland
has erect inflorescences with a few small resupinate flowers. The sepals and the petals
are free, unlobed and spreading. The ovate lip is unstalked and bears a median callus.
A long column-part is present in the knee-like bent gynostemium. Lateral gynostemium
appendages (column-wings) are rather small. Small staminode primordia are visible in
early ontogeny (Fig. 7A) and later develop into the small lateral appendages of the
mature gynostemium as seen in Fig. 7D. Structures likely to be auricles are not visible
in any stage. A prominent feature of the ontogeny is the median carpel apex which is
conspicuously three-lobed in middle stages with its lobes roughly equally long (Fig.
7B). Its central lobe loses its prominent appearance later on and grows into the gap
between the two thecae which is reminiscent of the intrathecal rostellum portion of
tribe Orchideae (Fig. 7C-D). The lateral carpel apices emerge connate to an undivided
edge and also remain undivided later on (not shown). The anther is apically
emarginate throughout ontogeny.
Pterostylis concinna R. Br.
A single species of the large genus Pterostylis (subtribe Pterostylidinae sensu Jones &
Clements 2002 - about 120 species in Australia, New Zealand, New Guinea and New
Caledonia) was available for study, and is of considerable interest as it is the only study
species outside tribe Diuridcae, having been transferred to tribe Cranichideae recently
(Pridgeon et al. 2003). The terminal inflorescences bear a single or a few resupinate
flowers with a prominent galea made up of median sepal and petals, two basally fused
lateral sepals and a mobile lip. In P. concinna the slender and curved gynostemium has
a long column-part. Large lateral gynostemium appendages (column-wings) are
present in the apical portion and are expanded next to the anther so that they almost
obscure it. They consist of rather wide and obtuse lobes pointing down, and narrow
and acute lobes pointing up. The stigma of the mature gynostemium is an entire pad
at the base of the column-part. Early stages of the gynostemium ontogeny of this
species were not available for study. In a middle stage (‘three-carpel-apex stage’) the
large anther is erect and the three carpel apices are visible with the median one being
as usually the largest (Fig. 7E). Prominent staminode primordia are clearly visible in
this stage. As could be expected they develop directly into the large apical wings of the
late stages and the mature gynostemium (Fig. 7E-I). From the ontogeny it is evident
that both the acute upper and the obtuse lower part of the mature lateral gynostemium
appendages are derived from these staminodes. In middle ontogenetic stages there is
also an insignificant suspected auricle visible (arrowhead in Fig. 7H), and in the
mature gynostemium the structure takes the shape of an obscure small tooth next to
the anther (arrowhead in Fig. 71). The median carpel apex is an obscurely three-lobed
organ in middle ontogeny (Fig. 7F-G). Lateral carpel apices are separate structures
soon after their initiation and remain separate until the column-part starts elongating.
Discussion
I he ontogeny of the anther and the three carpel apices, as well as the late formation of
the gynostemium of the species examined here, conform basically with that of all other
Australian members of Diurideae and Cranichideae
Telopea 11(1): 2005
25
Fig. 7. Gynostemium development of various species. A-D. Acianthusfomicatus. Gynostemium/base
of gynostemium in front view. E-I. Pterostylis concmna. E-G. Font views. H. Side view. I. Adult
gynostemium in side view with the left lateral gynostemium appendage removed. — SEM
micrographs. Bars: 0.1 mm in A-H; 1 mm in 1. Abbreviations: A = anther, a = staminodes,
c = carpel apices, me = median carpel apex
26
Telopea 11(1): 2005
Kurzweil, Weston and Perkins
monandrous orchids studied so far (e.g. Irmisch 1842; Payer 1859; Wolf 1865; Pfitzer
1888; Capeder 1898; Heusser 1915; Hirmer 1920; Jeyanayaghy & Rao 1966; Rao 1967;
Sattler 1973; Yang 1982; Rasmussen 1982, 1985b; Kurzweil, several papers; Leins et al.
1988; Schill et al. 1992; Linder & Kurzweil 1996; Luo & Chen 2000).
The anther is the first organ of the gynostemium to become visible. Like the six
perianth organs, it develops from the marginal parts of a transversally elliptic floral
primordium. The anther is erect in early stages and the gap between its two thecae is
of very early ontogenetic origin. As in other orchids it is the largest organ of the
gynostemium throughout early development, but in many species its growth later
decreases; the anther is therefore often comparatively small in the mature
gynostemium.
The three carpel apices emerge in front of the anther in early ontogeny and later
develop directly into the stigma lobes of the mature flower. The median carpel apex is
the first carpel apex to be initiated. It is erect and unlobed in early stages, and measures
approximately 3 A of the width of the anther soon after its initiation. It remains the
largest carpel apex in the early and middle ontogeny, but like the anther it often loses
its prominent appearance later on and is fairly small in the mature flower. Its apical
portion becomes structurally modified in late ontogeny and develops into the
rostellum. The derivation of the non-receptive rostellum from the median carpel apex
is also found in all other orchids examined so far. The two lateral carpel apices emerge
just after the median one in a position directly in front of it. Just after their initiation
they are either separate ( Genoplesium fimbriatum, Pterostylis concinna, Cryptostylis
erecta, Orthoceras strictum ) or connate into an entire or emarginate ridge ( Rimacola
elliptica, Acianthus fornicatus, Diuris longifolia, Thelymitra carnea, Calochilus
campestris). Both these character states are also found in the other orchids (Kurzweil
1998; Kurzweil & Kocyan 2002). The lateral carpel apices are initially about one third
to half as wide as the median carpel apex which is clearly visible where the lateral
carpel apices are separate or connate into an emarginate ridge. On the basis of
outgroup comparisons with other basally diverging families of Asparagales, the most
recent common ancestor of the orchids probably had three spreading stigma lobes, and
therefore the separate emergence of the lateral carpel apices is most likely the more
primitive condition for the family as a whole. In late ontogeny the two lateral stigma
lobes are generally basally united and also marginally fused with the median stigma
lobe to form a funnel-like structure. Finally, the lateral stigma lobes develop into the
receptive stigma and also a portion of varying size is contributed by the median stigma
lobe. The surface of the stigma is papillose in the late stages.
In some species the organ-complex of the three stigma lobes is elevated together with
the anther and the column-wings in that the common base of all of these organs
elongates, and thus the column-part of the gynostemium is formed. The formation of
this column-part takes place in the late stages. The resulting gynostemium can be very
long (e.g. Rimacola elliptica, Caleana major, Pterostylis concinna, Acianthus fornicatus).
The fusion may also be absent or very weak and is then confined to the base of the
organs of the gynostemium ( Genoplesium fimbriatum, Diuris longifolia, Orthoceras
strictum). The weak fusion of the gynostemium in Diuris, Orthoceras and Genoplesium
has been thought to be a primitive condition (see Dressier 1986: 10). However, recent
molecular phylogenetic analyses of the Diurideae and outgroups (Kores et al. 2000,
2001, Clements et al. 2002) strongly imply that weak fusion has been secondarily
derived in these taxa. Such a reversal is readily explicable as the result of neotenic loss
Australian members of Diurideae and Cranichideae
Telopea 11(1): 2005
27
of gynostemium fusion and elongation late in floral organogeny. In the other
monandrous orchids the parts of the gynostemium are completely fused, and very
elongate column-parts occur in Epidendroideae and two genera in the the
Orchidoideae-Diseae: Satyrium and Pachites.
Lateral gynostemium appendages
Of particular interest are the lateral gynostemium appendages which are often referred
to as column-wings in floristic and taxonomic works. It was found that they are
ontogenetically directly derived from massive bulges superposed to the petals which
can be observed in early and middle ontogeny. On account of their early initiation and
the position in front of the petals they are here interpreted as vestiges of the lateral
stamens of the inner staminal whorl (i.e. staminodes in a position aj and ^ which
corresponds well with most other Orchidaceae (e.g., Kurzweil 1987a, 1988). Previously,
similar prominent staminodes were also observed and illustrated in middle stages of
the gynostemium development of Gavilca glandulifcra, a member of the tribe
Chloraeeae, and the diurids Ditiris punctata, Prasophyllum concinnutn and Microtis
parviflora (Rasmussen 1982). Since orchids have most probably evolved from ancestors
with 3+ 3 stamens (see also Rasmussen 1982) the presence of pronounced staminodes
must be interpreted as an ancestral feature in each of the two clades of monandrous
orchids, the Vanilloideae and Orchidoideae-Epidendroideae. The primitive feature of
massive staminodes has apparently constantly been retained in the orchid subfamily
Epidendroideae (Kurzweil 1998; Kurzweil & Kocyan 2002). Interestingly, in
Epidendroideae the staminodes sometimes disappear in late ontogeny as they become
incorporated into the gynostemium. In most species of tribes Cranichideae, Diseac
and Orchideae (scnsu Pridgeon et al. 2001, 2003) the staminodes a] and a2 are small
or obscure in early ontogeny (Kurzweil 1987b, 1988). Given the sister group
relationship between Diurideae and Cranichideae-Chloraeeae on the one hand and
Diseae and Orchideae on the other (Kores et al. 2001; Chase et al. 2001) staminode
reduction has probably evolved independently in Cranichideae and OrchideaeDiseae.
In a few species examined here there are also additional lateral gynostemium
appendages which become visible in middle developmental stages. They are initiated
as small or obscure but clearly visible bulges in a dorsolateral position on the anther.
Their time of initiation and place of origin corresponds well with that of the filament
appendages termed auricles of tribe Orchideae (Kurzweil 198/b; Luo & Chen 2000),
and the structures are here also interpreted as such. The homology of the auricles of
Orchideae with filament appendages was originally suggested by Vermeulen (1966)
whose investigations were based on the study of adult and especially teratological
flowers. In Orthocerasstrictum such auricles are not constant in their appearance. This
points to the possibility that the auricles may be genetically fixed even if they are
externally not always expressed. In superficial appearance the auricles observed here
are mostly unsculptured, differing from those of Orchideae which are usually strongly
sculptured. In the present study heavily sculptured auricles were only observed in
Lyperanthus suaveolens and Calochilus campestris, and previously a heavily sculptured
auricle was illustrated in an unidentified Calochilus species (Kurzweil 1998, Fig. 11L).
No sign of any auricles was here found in Acianthus fornicatus, Cryptostylis erecta,
Chiloglottis sp., Corybas fimbriatus and Microtis parviflora. Auricles in Diurideae have
been reported before by Dressier (1986: cladogram on p. 13) although it is not clear
whether the term was used in the sense of filament appendages (however, Dressier
28
Telopea 11(1): 2005
Kurzweil, Weston and Perkins
included Chloraeinae in his tribe Diurideae which are now recognised as a distinct
tribe Chloraeeae; Pridgeon et al. 2003). In other orchids, auricles have so far only been
reported in tribes Orchideae and Diseae. While they are usually prominent in
Orchideae they are mostly small and reduced in Diseae. The only exception known as
yet is the genus Bartholina (Orchideae) where the auricles are absent (Kurzweil &
Weber 1991), and this is probably the result of secondary loss in this genus. The shared
occurrence of auricles in Diurideae, Orchideae + Diseae, Chloraeeae and Pterostylis, a
basally diverging lineage of Cranichideae, is most parsimoniously interpreted as a
synapomorphy for these tribes that has been secondarily lost in most Cranichideae.
The appearance of the auricles in the mature flowers differs among the species
examined. In Cenoplesium fimbriatutn the staminodes and the auricles become fused
in the course of development while their common base elongates. These two organs
together develop into the prominent lateral gynostemium appendages (column-
wings) which are therefore of dual origin. The auricles of Rimacola elliptica develop
into small processes on top of the staminodial wings. In Calcutta major and Diuris
longifolia prominent auricles were not observed in early and middle stages but the
lateral appendages of the late-ontogenetic gynostemium have a small apical tooth or
are shallowly two-lobed, respectively. Compared with the other species examined here
this may again suggest an involvement of both staminodes and auricles, although there
is obviously no clear ontogenetic evidence for a dual origin. The auricles of Pterostylis
concinna arc small lobes next to the anther behind the column-wings. Previously, a
distinct two-lobing of the lateral gynostemium appendages was found in middle
developmental stages in Gavilea glandulifera and Prasophyllum concinnum but a
possible ontogenetic derivation from two different structures was not suspected
(Rasmussen 1982). In view of the present findings the two-lobing found in these two
species may be suggestive of the origin from both staminodes and auricles. It is also
noteworthy that the dorsal lobe of the lateral gynostemium appendages of
Prasophyllum concinnum is said to contain raphides (Rasmussen 1982) which is an
anatomical feature frequently associated with auricles. In other orchids, a dual origin
of the lateral gynostemium appendages is common in Orchideae where the auricles are
frequently heavily sculptured appendages on top of or at the posterior end of a
staminodial base (Kurzweil 1987b, 1990; Kurzweil & Weber 1991, 1992; Luo & Chen
2000 ).
Three-lobing of the median carpel apex
It was found that the median carpel apex of Acianthus fornicatus, Calochilus campestris
and Corybas fimbriatus is clearly three-lobed in middle developmental stages, and
Genoplesium fimbriatum, Caleana major, Pterostylis concinna, Microtis parviflora and
Orthoceras strictum have a shallowly three-lobed median carpel apex in this stage. This
is remarkable as the vast majority of Orchidaceae have an unlobed median carpel apex
throughout middle and late development, which is apparently the basic condition in
the family. The three-lobing observed here is reminiscent of tribes Orchideae and
Diseae where the median carpel apex is usually deeply three-lobed throughout
ontogeny (Vogel 1959; Vermeulen 1959, 1966; Dressier 1981, 1993; Kurzweil, several
papers) although it has been reduced to a two-lobed or unlobed structure in some
Diseae (Kurzweil 1991, 1996; Linder & Kurzweil 1996). This shared feature might be
another morphological synapomorphy for the Orchidoideae, albeit one that has been
secondarily lost in a number of different lineages.
Australian members of Diurideae and Cranichideae
Telopea 11(1): 2005
29
Alternatively, the three-lobing of the median carpel apex observed here may also
merely be a precondition for the formation of an apical viscidium in late ontogeny, and
is therefore not necessarily evidence of phylogenetic relationship. However, this
explanation is unlikely as one would not expect this feature to already be expressed
early in ontogeny.
Phylogenetic considerations
The phylogenetic relationships and the delimitation of diurid orchids have been rather
disputed among taxonomists.
In the past the diurid orchids were often treated in a subgroup of Orchidaceae that
contains terrestrial orchids with a large number of primitive characters
(e.g. ‘Acrotonae-Polychondreae’, Schlechter 1926; ‘Neottioideae’, Garay 1960, 1972;
‘Epidendroideae-Contribc Neottianthae’, Vermeulen 1966;‘Neottioideae’, Brieger et al.
1970-2000). After realising that this group is an artificial aggregation that contains
several basally diverging lineages of both Orchidoideae and Epidendroideae
(e.g. Rasmussen 1982; Dressier 1993; Cameron et al. 1999), its tribes and subtribes
were either transferred to other subfamilies or raised to the rank of separate new
subdivisions (e.g. subfamily Spiranthoideae; Dressier 1979, 1981, 1993). Although the
diurid orchids, as circumscribed by Dressier (1981, 1993), were in the past treated as a
natural group, it is now evident that they are not monophyletic (Kores et al. 1997,2000;
Cameron et al. 1999; Molvray et al. 2000; Pridgeon et al. 2001). Groups which do not
belong to the core clade of diurid orchids are the Pterostylis and Chlomea groups.
A relationship of the diurid orchids with the Orchideae and Diseae (subfamily
Orchidoideae) is now widely accepted on the basis of morphological and anatomical
data (Dressier 1979, 1981, 1993; Rasmussen 1985a; Pridgeon et al. 2001) and was also
confirmed by molecular investigations (Kores et al. 1997, 2000; Cameron et al. 1999;
Molvray et al. 2000). The most frequently cited morphological feature shared by diurid
orchids on the one hand and Orchideae and Diseae on the other are root tubers. Some
authors have expressed doubt about the homology of these structures because of their
anatomical diversity (see Pridgeon & Chase 1995 and references therein). However, the
unusual structural similarities that the root tubers of all of these groups share
prompted Dressier (1993) and Pridgeon & Chase (1995) to postulate their homology.
An implication of this idea in the light of more recently published molecular
phylogenetic analyses (Kores et al. 1997, 2000; Cameron et al. 1999; Molvray et al.
2000) is that root tubers are a synapomorphy for the subfamily Orchidoideae but that
they have been secondarily lost in several different lineages.
Conclusions
Two interesting similarities shared by Diurideae and Pterostylis (Cranichideae) with
tribes Orchideae and Diseae were found in the present study, namely the auricles and
the early three-lobing of the median carpel apex, which add further support to the
theory of a close relationship of the groups.
A) Definite auricles or obscure structures suggestive of such auricles were here found
in eight of 11 species where the early or middle ontogeny was examined.
Auricles of diurid orchids have been observed before but due to the lack of ontogenetic
30
Telopea 11(1): 2005
Kurzweil, Weston and Perkins
studies the similarity in their ontogeny to the auricles of the Orchideae and Diseae was
not noted.
B) The second point of interest relates to the early three-lobing of the median carpel
apex in six of the 11 species where relevant stages were observed. The median carpel
apex is frequently only shallowly lobed but in three species the three-lobing is rather
pronounced.
An extensive discussion of the systematics of the diurid orchids which is based on
morphological-ontogenetic gynostemium characters would obviously require far
more complete sampling.
Acknowledgments
Collecting was made possible with a permit from the NSW National Parks and Wildlife
Service (license no. A2409) which is gratefully acknowledged. The first author would
also like to express his gratitude to Ruth Rudkin, Alan Dash, Wal and Gill Upton,
Jim Lycos, Graeme Bradburn, John Riley, Darryl Smedley and Andrew Harvie for their
assistance during a fieldtrip to the Sydney region. The second author would like to
thank the late Tony Bishop for company and assistance in the field while collecting
material for this study. We also thank the staff of the Electron Microscope Unit of the
University of Cape Town (Figures 1-5, 7) and the University of Sydney (Figure 6) for
providing SEM time and for technical support.
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Weston PH (1994) Methods for rooting cladistic trees. Pp. 125-155, in Siebert DJ, Scotland RW
8c Williams DM (eds) Models in phylogeny reconstruction (Oxford University Press: Oxford)
Woll T (1865) Beitrage zur Entwicklungsgeschichte der Orchideenbliite. Pringsheim Jahrbuch fiir
die wissenschaftliche Botanik 4: 261-384.
Yang S-H (1982) Studies on the development of flower in Gastrodia elata Bl. Acta Botanica Sinica
24: 21-27. (in Chinese language)
Manuscript received 13 July 2004, accepted 28 April 2005
Australian members of Diurideae and Cranichideae
Telopea 11(1): 2005
33
Appendix: List of material studied
The following list cites all specimens which have been studied in their floral ontogeny.
In most cases herbarium vouchers were deposited in the herbarium of the Royal
Botanic Gardens Sydney (NSW). The nomenclature follows mainly Jones (1988).
PHW is an abbreviation for the second author of the paper, RH. Weston; NSW
indicates a number in the National Herbarium of New South Wales.
Acianthus fornicatus R. Br. - PHW 1393
Caladenia earned R. Br. - Kurzweil 1928
C. catenata (Smith) Druce - Abell 67
Caleana major R. Br. - PHW 1229, Kurzweil s.n.
Calochilus campestris R. Br. - NSW446115
C. robertsonii Benth. - Abell 72
Chiloglottis sp. - PHW 1278
Corybas fimbriatus (R. Br.) Rchb. f. - PHW 1392
Cryptostylis erecta R. Br. — PHW 2473
Diuris longifolia R. Br. - sine collector
Eriochilus autumnalis R. Br. - PHW 1382
Genoplesiutn fimbriatum R. Br. - PHW 1277, PHW s.n.
Glossodia minor R. Br. — PHW 1235, PHW 1401
Lyperanthus suaveolens R. Br. - PHW 1224
Microtis parviflora R. Br. - PHW 1251
Orthoceras strictum R. Br. — PHW 1271, PHW 1279, PHW 1584
Pterostylis concinna R. Br. - PHW 1398
Rimacola elliptica (R. Br.) Rupp - Bishop J67/31-37, PHW 1586
Thelymitra carnea R. Br. - PHW 1227, PHW 1232, PHW 1242, PHW 1404
—
Telopea 11(1): 35-41
A new species of Goodenia (Goodeniaceae)
from Nocoleche Nature Reserve, Far Western
Plains, New South Wales
Belinda J. Pellow 1 and John L. Porter 2
' Janet Cosh Herbarium, School of Biological Sciences, University of Wollongong, Northfield’s
Avenue, Wollongong, NSW 2522, Australia, email: bpellow@ uow.edu.au
2 Waterbird and River Ecology Unit, NSW Department of Environment and Conservation,
P.O. Box 1967, Hurstville, NSW2220, Australia
Abstract
Goodenia nocoleche was cultivated in sediment collected from a freshwater temporary wetland in
Nocoleche Nature Reserve. Here it is newly described and illustrated with additional notes on its
ecology.
Introduction
A species of Goodenia was cultivated from sediments collected in Nocoleche Nature
Reserve during a study on seed banks of arid zone wetlands of the Paroo-Bulloo River
region of New South Wales (Porter 2002). It represents a new species. An ephemeral
herb, it has also been observed growing in its natural habitat at Pied Stilt Swamp and
a clay pan on the Wanaaring-Hungerford Rd. (29°28 00 S, 144°24 00 E).
Methods
Non-dormant seed banks of wetlands were sampled in June and September 1997
following the methods of Brock et al. (1994) to collect sediment cores. Sixty sediment
samples were collected randomly from each wetland, (depth = 2.5 cm, surface area =
0.012m 2 ). Sediment samples were placed in shallow plastic trays for drying, then
submerged in water (60 cm depth) to stimulate germination.
Examination of the cultivated specimens revealed a yellow corolla with ovules and
seeds numerous in two rows on either side of the septum, characters which placed
them in the section Porphyranthus of the genus Goodenia. However, the small flowers
with calyx lobes which are just short of or equalling the corolla lobes distinguish it
from others in this section.
© 2005 Royal Botanic Gardens and Domain Trust
ISSN0312-9764
36
Telopea 11(1): 2005
Pellow and Porter
Taxonomy
Goodenia nocoleche B. Pellow & J. L. Porter sp. nov.
Herba ephemera a speciebus duabus artissime cognatis (G. lamprospenna atque
G. paniculata) corolla parva 5-5.5 mm longa, lobis calycis 4-6 mm longis corollam
subaequantibus distinguenda.
Holotype: NEW SOUTH WALES: cultivated from seed collected 15 km S of
Wanaaring, Pied Stilt Swamp, Nocoleche Nature Reserve, /. Porter, 29 Mar 2000
(NSW 458574).
Ephemeral amphibious herb to 40 cm high, vegetative parts glabrous. Basal leaves with
floating lamina 20-40 mm long, 7—14 mm wide, thin, glossy green above, flat,
lanceolate, margins undulate and minutely toothed, apex acute with tooth. Petioles
elongated to 60 cm long, much longer than lamina, lengthening with water depth.
Cauline leaves present at base of flowering stem 10-40mm long, 2-3mm wide. Flowers
in racemes. Bracts longer towards base of inflorescence, linear and sometimes
resembling leaves, 10-40 mm long. Bracteoles linear, 8-10 mm long, 0.6-1 mm wide,
with minute simple and glandular hairs; bracteoles occasionally producing axillary
buds which extend into new inflorescence branches. Pedicels 2-5 mm long without
articulation. Floral tube 3-5 mm long with minute simple and glandular hairs. Calyx
lobes linear, 4-6 mm long, 0.8-1 mm wide, numbering 5 or sometimes 6, attaching in
top !/s of floral tube and covered with minute simple and glandular hairs. Corolla
yellow, 5-5.5 mm long, lobe tips often tinged with pink or purple; glandular and
simple hairs present externally, glabrous internally; no enations observed; anterior
pocket obscure; abaxial lobes 2 mm long with wings approximately 0.5 mm wide;
adaxial lobes 2.5 mm long with wings approximately 0.5 mm wide. Staminal filaments
1.8-2.0 mm long, anthers 0.5-0.8 mm long. Ovary with numerous ovules in 2 rows on
either side of the septum; septum almost as long as the ovary. Style 2-2.5 mm long
with scattered long simple hairs; indusium 1-1.5 mm long, 1-1.3 mm wide, purplish
in colour, hairs present on adaxial rim, abaxial rim without hairs. Fruit pale, obconical,
5-6.5 mm long, 2-3 mm wide, hispid with short simple and glandular hairs. Seed 1 mm
long, 2.5 mm wide, light brown, glossy, reticulation faint, winged. Figures 1 and 2.
, elected specimens examined: New South Wales: Far North Western Plains: Cultivated from
S'vfonc 0 CCted , fr ° n ! P ' ed Stilt Swam P’ Nocolcche Nature Reserve. 15km south of Wanaaring.
I . 2 18 S, 144°05'12"E. 100m alt. /.L. Porter ILP 255, 29 Mar 2000 (8107-WOLL, 8108-WOLI
8111-WOLL, 8113-WOLL). '
Distribution: This species has been recorded in the seed banks of five wetlands in the
1 aioo-Bulloo River region. It was common (mean density 1,511 m 2 ± 189 se) in the
seed bank of one freshwater temporary wetland, Pied Stilt Swamp and uncommon (mean
Swamp (29 ° 52 ' 45 " S 144°06'30"E), Waitchie Lake
(.0 loot) S 144 01 15 E), Lake Altibouka (29°49'00"S 142°45'00"E) and I akp
3). has also been ZdS f™, S
suvations growing in Pied Stilt Swamp and another unnamed wetland c. 70 km
no. h of Wanaanng. Three of these locations (Pied Stilt, Momba and Numalla) are
within conservation reserves on the Paroo river floodplain (Fig. 3).
7i‘ Ch ‘ m ° St ab “" dant in ('"’Ponry freshwater wetlands.
St.lt and Momba Swamps are small (22 and 61 ha), shallow (1.3 and 1.4 m)
Goodenia nocoleche (Goodeniaceae)
Telopea 1 1(1): 2005
Fig. 1. Goodenia nocoleche B. Pellow & J.L. Porter, a, habit; b, flower; c, fruit. Scale bar:
b = 6mm; c = 6 mm.
38
Telopea 11(1): 2005
Pellow and Porter
Fig. 2. Seeds of Goodenia nocoleche B. Pellow & J.L. Porter.
Fig. 3. Occurrence of Goodenia nocoleche (inset, dark square) in Australia’s arid zone (< 500 mm
annual average rainfall, stippled) and five wetland seed banks sampled: Pied Stilt Swamp (1),
Momba Swamp (2), Waitchie Lake (3), Lake Altibouka (4), Lake Numalla (5). Unnamed claypan
wetland where Goodenia nocoleche was observed growing (6). Conservation reserves on the Paroo
River are Currawinya National Park (CWY), Nocoleche Nature Reserve (NOC) and Paroo Darling
National Park (PDY). Town (dark square) is Wanaaring (WAN).
Goodenia nocoleche (Goodeniaceae)
Telopea 11(1): 2005
39
‘claypan’ basins with uneven substrates, caused partly by expansion and contraction of
grey or brown cracking clays. They fill predominantly from local rainfall and only
connect to the Paroo floodplain via Momba Creek after heavy local rain. From
1990-2000 the swamps held water for a total of 17 and 22 months respectively. Their
water is turbid after flooding, clearing gradually during the drying phase, especially in
Pied Stilt Swamp. Plant communities in these temporary wetlands undergo
considerable changes during erratic filling and drying cycles, as aquatic plants, sedges,
annual grasses, herbs and forbs become abundant during and after inundation, before
declining again. During prolonged dry periods ground cover may be virtually absent
and many species remain hidden below the surface as propagules in seed banks or
underground storage organs.
Vegetation around the swamp margins consists of a narrow band of trees, mostly
Eucalyptus largiflorens and E. populnea subsp. bimbil. Within the swamps trees are
absent, and perennial shrub understories are absent or sparse with occasional grasses.
After flooding sedges are common at the margins, and in open water floating-leaved
submergent or partially emergent species may occur. (Kingsford & Porter 1999).
Ecology: Goodenia nocoleche can germinate and grow in standing water up to 0.6 m
deep, with floating leaves on greatly extended petioles (Fig. 4), similar in appearance to
Potomogeton octandrus. As water recedes inflorescences emerge and grow rapidly. The
plants die back rapidly as sediments dry completely. Goodenia nocoleche has been
observed growing in Pied Stilt Swamp during several flood events (December 1998,
2000) and appears to be a summer annual needing inundation in shallow temporary
freshwater wetlands to stimulate a germination response from the persistent seed bank,
followed by partial drying to initiate flowering. These observations are supported by
the behaviour of the species in cultivation. Ability to initiate vegetative growth and
Fig. 4. Goodenia nocoleche B. Pellow & J.L. Porter in cultivation.
40
Telopea 11(1): 2005
Pellow and Porter
form floating leaves early in the drying cycle of temporary wetlands may confer some
advantage compared to other low growing amphibious species that also colonise the
exposed sediments of drying wetlands (e.g. Mimulus repens, Ranunculus sessiliflorus
var. pilulifer, Pratia darlingensis). In functional group terms, Goodenia nocoleche is an
‘amphibious fluctuation-responder’ because it alters it growth pattern or morphology
in response to the presence or absence of water (Brock & Casanova 1997). Amphibious
and aquatic plants in arid zone wetlands are generally poorly known, because many
species are short-lived and access to these remote areas is difficult when water is
present. Potential threats to this species include invasive weeds that are spread by water
such as Pistia stratiotes and Xanthium occidentale and alterations to flow regimes of
rivers and floodplains.
Etymology: the epithet is taken from the name of the Nature Reserve where this
species was found. It is derived from an aboriginal word meaning ‘place of many
waterholes’
Notes: In his description of two new species of Goodenia from the Northern Territory,
Albrecht (2002) defines bracteoles according to Briggs and Johnson (1979) calling the
structures opposite or sub-opposite bracteoles due to the presence of buds in their
axils. We have chosen to keep the term bracteole for these occurrences, according to
Carolin (1990, 1992). Although buds do occur within the axis of some bracteoles of the
inflorescence the majority of the bracteoles do not exhibit them.
A specimen of Goodenia lamprosperma (NSW 460174) collected in a semi-aquatic
environment exhibits similar leaf morphology to that of G. nocoleche. Other specimens
of G. lamprosperma not recorded as growing in semi-aquatic situations exhibited
different leaf morphology. The characters of extended leaf petiole, larger lamina and
reduced stem length may be related to the semi-aquatic habitat. Goodenia nocoleche has
been observed in the field with terrestrial leaves immediately prior to flowering in
situations where the water levels have receded but no specimens have been collected
for examination. Seedlings of G. nocoleche (8110-WOLL) show a number of delicate,
lanceolate leaves 12mm long and 2mm wide which are produced prior to the
development of the basal leaves with extended petioles.
Acknowledgments
We would like to thank the staff at the National Herbarium of New South Wales, in
particular Joy Everett for her comments on the manuscript. Peter Wilson provided the
Latin diagnosis and Catherine Wardrop prepared the botanical illustrations. Roger
Carolin provided advice and encouragement.
References
Albrecht DE (2002) New species and notes on central Australian Goodenia (Goodeniaceae).
Nuytsia 15: 1-9.
Briggs BCi & Johnson LAS (1979) Evolution in the Myrtaceae—evidence from inflorescence
structure. Proceedings of the Linnean Society of New South Wales 102: 157-256.
Brock MA & Casanova MT (1997) Plant life at edge of wetlands: ecological responses to wetting
and diying patterns. In N klomp & LI Klomp (eds) Frontiers in ecology: building the links.
(Elsevier Science: Oxford, UK)
Goodenia nocoleche (Goodeniaceae)
Telopea 11(1): 2005
41
Brock MA, Theodore K & O’Donnell L (1994) Seed-bank methods for Australian wedands. Aquatic
Botany 45: 483-493.
Carolin RC (1990) Nomenclatural notes and new taxa in the genus Goodenia (Goodeniaceae).
Telopea 3: 517-570.
Carolin RC (1992) Goodenia. Pp. 147-281 in AS George (ed.) Flora of Australia, vol. 35. (Australian
Government Publishing Service: Canberra)
Kingsford RT & Porter J (1999) Wetlands and waterbirds of the Paroo and Warrego Rivers.
Pp. 23-50 in RT Kingsford (ed.) A free-flowing River: The Ecology of the Paroo River
(NSW NPWS: Sydney Australia)
Porter J (2002) Effects of salinity, turbidity and water regime on arid zone wetland seed banks.
Verb. Inernat. Verein. Litnnol. 28: 1486-1471.
Manuscript received 1 September 2004, accepted 2 May 2005
.
Telopea 11(1): 43-51
Reassessment of Indigofera pratensis var.
coriacea Domin and var. angustifoliola Domin
(Fabaceae: Faboideae) with the recognition of
a new species
Aniuska A. Kazandjian 1 and Peter G. Wilson 2
‘Tropical Plant Sciences, School of Tropical Biology, James Cook University, Townsville QLD 4811,
Australia, and Fundacion Instituto Botanico de Venezuela, Caracas 1010-A, Venezuela
2 National Herbarium of New South Wales, Royal Botanic Gardens,
Sydney NSW 2000, Australia
Abstract
The two varieties of Indigofera pratensis described by Karel Domin are reassessed. Indigofera
pratensis var. coriacea is distinct from the widespread Indigofera pratensis sens. str. in a number of
reproductive and vegetative characters and is here raised to specific rank as Indigofera scabrella.
Indigofera pratensis var. angustifoliola is judged to be an environmental or rare geographic variant
of Indigofera pratensis sens. str. and not worthy of continued recognition. Lectotypes are designated
for both names.
Introduction
The Indigofera pratensis complex is widespread in open habitats in forest, woodland
and shrubland or in savanna. Plants are usually found as low to tall shrubs with
purplish-pink flowers that are usually more than 10 mm long and borne in dense,
showy axillary racemes. In eastern Queensland, its range extends from the northern
part of Cape York (c. 12°35') to just north of Brisbane (c. 27°15') but there are also
scattered populations in inland Queensland, particularly between Townsville and
Hughenden and in the vicinity of Mount Isa, with a few records from the north¬
eastern part of the Northern Territory. A survey of herbarium specimens of I. pratensis
was carried out as part of a review of this complex (Kazandjian 2002) and revealed
considerable variation such that some morphotypes in this complex are apparently
distinct at species or subspecies level. The inland populations of the complex were
largely unknown when Domin (1926) divided Indigofera pratensis into three varieties:
I. pratensis var. typica (- I. pratensis var. pratensis), I. pratensis var. coriacea and
I. pratensis var. angustifoliola. The two new varieties were described from limited
material and distinguished primarily on leaflet characters. In the present paper, we
reassess and lectotypify these infraspecific taxa.
© 2005 Royal Botanic Gardens and Domain Trust
ISSN0312-9764
44
Telopea 11(1): 2005
Kazandjian and Wilson
Indigofera pratensis var. coriacea and Indigofera pratensis var. pratensis
Domin (1926) distinguished variety coriacea as having fewer, smaller leaflets with a
more coriaceous texture, somewhat prominent lateral venation but obscure reticulate
venation. Indeed, he suggested that I. pratensis var. coriacea might possibly be treated
as a new species (‘vielleicht eine selbstiindige Art’).
Hairs
Hairs in these two taxa can be divided into two types, here designated T1 and T2. The
T-shaped hairs are the most commonly encountered type in Indigofera and have been
described as ‘uniseriate macroform biramous hairs’ often with ± equal arms
(Prabhakar et al. 1985, Schrire 1995). Tl hairs are ‘appressed’ following the definition
proposed by Hewson (1988), that is, the angle of elevation above the surface is in the
range 0°-15°. T2 hairs have their arms ascending at angles greater than this, to around
45°, with arms straight. Indigofera pratensis var. pratensis has both kinds, with the T2
type less frequent but tending to have longer arms, while var. coriacea has hairs of the
Tl type only (Fig. 1 c, f). Hair density on the leaflets varies somewhat between the two
taxa. In I. pratensis var. pratensis the leaflets have sparser hairs and are usually markedly
discolorous with the additional feature of rather conspicuous reticulate venation,
particularly on the lower surface (Fig. 2 a, b), whereas in var. coriacea, the leaflets have
moderately dense hairs on both surfaces and are more or less concolorous (Fig. 2 c, d).
Stipules
Stipules differ markedly in length between these two taxa (Fig. 1 b, e): they are
triangular-linear and 5-12 mm long in var. pratensis, but deflate and 1.0—1.5(—2) mm
long in var. coriacea. These length differences are parallelled in the inflorescences
(Fig 1 a, b) where the bracts match the stipules in shape and relative length, being 2-3 mm
and c. 0.5 mm long, respectively. There is a similar contrast in the size and shape of the
calyx teeth: in var. pratensis they are 0.9—1.5 mm long and acute, while in var. coriacea
they are 0.5-0.9 mm long and obtuse.
Inflorescence characters
Overall inflorescence characters also differ. The inflorescences of I. pratensis var.
pratensis appear to be denser than those of var. coriacea (Fig. 1 a, d). This is due to two
features, the spacing of flowers and the length of the pedicels. On the racemes, the
flowers are not always equally spaced and are often in loose groups of two or three with
a longer space between the groups. In this study, density of flowers was measured as
the distance between adjacent recently opened flowers (or groups of flowers) and was
found to be (4—) 5-8 mm in var. coriacea but only 2—3(—4) mm in var. pratensis. Pedicel
length is clearly distinct: only 1.5-2.5 mm in var. pratensis compared with 3-5(-7) mm
in var. coriacea.
Plants grown under standardised environmental conditions were also well separated
using the same morphological criteria (Kazandjian 2002), confirming that these
character variations are not environmentally induced. Furthermore, the measurable
morphological differences between the two taxa are maintained even though their
populations are (at least partially) sympatric. Indigofera pratensis var. coriacea has a
restricted distribution to the west of the Atherton Tableland, primarily around
CJiillagoe and the Walsh River but extending to the Mount Surprise area, and
Indigofera pratensis complex
Telopea 11(1): 2005
45
Fig. 1. Morphological characters. Indigofera pratensis var. pratensis. a, inflorescence showing long
bracts, short pedicels and crowded flowers; b, stipule, c, SEM of stem showing Tl- and T2-type
hairs. Indigofera pratensis var. coriacea d, inflorescence showing short bracts, long pedicels, short
sepals and well-spaced flowers; e, stipule; f, SEM of stem showing Tl -type hairs. (a,b from Puttock
UNSW 14239-, c from G. Wilson 5 (JCT); d,e from Kazandjian NSW 705606; f from ]ackes 9954
(JCT). Scale bar: a = 3mm, b = 3 mm, c = 400 pm, d = 3mm, e = 6 mm, f = 500 pm.
46
Telopea 11(1): 2005
Kazandjian and Wilson
Indigofera pratensis var. pratensis has a wide range in eastern Queensland. The two taxa
co-occur in the Chillagoe area without any sign of intergradation. Since there is no
indication of any hybridisation between them, we therefore conclude, on both
morphological and biological grounds, that Domin’s var. coriacea should be
recognised at specific rank.
Indigofera scabrella Kazandj. & Peter G. Wilson, nom. et stat. nov.
Indigofera pratensis var. coriacea Domin, Biblioth. Bot. 89 (3): 191 (1926), non
I. coriacea Aiton, 1789.
Type citation: ‘Bei Chillagoe und besonders in den Savannenwaldern zwischen
Crooked Creek und dem Walsh River (Domin II. 1910)’ [near Chillagoe and
particularly in the savanna woodland between Crooked Creek and the Walsh River).
Lectotype (designated here): Queensland: in xerodrymio inter riv. Crooked Creek et
Walsh River, K. Domin 4515, Feb 1910 (PR 527226) (Fig. 3).
Residual syntype: Queensland: in xerodrymio inter fl. Walsh River et opp. Chillagoe,
K. Domin 4514, Feb 1910 (PR 527225).
Fig. 2. Leaflet characters. Indigofera pratensis var. pratensis. a, undersurface showing conspicuous
reticulum; b, upper surface. Indigofera pratensis var. coriacea. c, undersurface showing
inconspicuous reticulum ; d, upper surface. (a,b from Puttock UNSW 14239; b,c from Kazandjian
NSW 705606). Scale bars: 3mm.
Indigofera pratensis complex
Telopea 11(1): 2005
47
National Hir»ai(i<im or n.s.Vi
PK
LC fToT') ft.
I- profiler,;. r.*[v<U. vt*. c.
Nationai. Himariuu orN.S.W.
h j>r*tf, rival/
m m im mnomB m m 527226
Dr. K. DOM IN, Iter Australiensc a.
1909 —1910 Nro.
TndU’nfera pratenel? V.TT^Ll. _
va’i.toriacea DOM IN vur.n,
Queensland: in xerodrymio inter riv.
Crooked Creek et V/alah River
legi II. /$> 10.
aK 0 .no.23/19 60
DETER min A VI r Pcier Ci. W.lw.n . IV
Fig. 3. Lectotype of Indigofera pratensis var. coriacea.
mi
48
Telopea 11(1): 2005
Kazandjian and Wilson
I'ntK MW AWT Pfter G Wilton lV
mmcM ipse? liTioiiUs mm ;~) 2 ; 220 - ^
Aus dein Botanisehen Mut>oum zu Hamburg. /^P; y
(€x musea £odeJ/roy 3{amburgensi)
Indigofern pratensl? MUST,!..
var.anguatifolla DOFiTK.
4Cr*%U-
frope £ rieb < in* r r -rer jTustralian or.
, 1863-1863. legit jTmalia j)!etrich.
ako.no. 23/ 19 60
HEppA Pinw sto HATioaiLfs mm 527229
Aus dera Botanisehen Museum zu Hamburg.
(£x museo $odcjjroy dfcmburgenstj
Ir.digofera pratenals "USLT .var.angua t ifolla DOM.
/* det.Dr K.Domin.
1 _
Prope - nw jTustraliae or.
1863-1863. legit Jtmalla J)ietr!ch.
akc.nc.23/l960
NAnoNAL Hi««»«!' m HI NS W
/’/? £T2.T^
lc
/ prA&4~siT
DFTtRMINAVIT Pn.r r, Wciioo n^f
Fig. 4. Lectotype of Indigofera pratensis var. angustifoliola.
Indigofera pratensis complex
Telopea 11(1): 2005
49
Shrub or sub-shrub 0.5-0.8(-1.5) m high. Young stems angular, densely covered with
hyaline appressed Tl-type biramous hairs. Leaves pinnate, 47-83 mm long, with (5-)
7-11 leaflets; stipules deltoid, 1.0-1.5(-2) mm long, (0.5-)0.7-0.9 mm wide; petiole
c. 5-9 (-12) mm long; rachis furrowed. Leaflets coriaceous, green above and paler
beneath; stipellae inconspicuous, to 0.7 mm long; lamina ovate to elliptical; the basal
leaflets are (9-) 11—23(—30) mm, the lateral leaflets c. 12-25 mm and the apical leaflets
13-29 mm long; upper surface with sparse, appressed hairs, lower surface with
moderately dense hairs; apex nrucronate; secondary veins conspicuous but not
prominent, the reticulum usually inconspicuous. Inflorescence a lax axillary raceme
up to 150(—220) mm long, much longer that the subtending leaf; distance between
adjacent mature flowers more than 4 mm. Bracts deltoid to narrowly triangular,
0.7-1.4 mm long. Flowers purple, 10-12 mm long; pedicels long, 3—5(—7) mm. Calyx
1.7-2 mm long; teeth 0.5-0.9 mm long, slightly shorter than the tube. Standard
purple, 9-11 mm long, 5.5-7 mm wide, moderately hairy on the back, the apex
obtuse-acute when the standard is fully expanded. Wings shorter than or equal to the
standard, darker in colour towards the apex, 7-10.5 mm long, 2-3 mm wide. Keel as
long as the standard, lateral pockets c. 1 mm long; hairs few and scattered at the base,
becoming moderately dense towards the apex and margin. Staminal tube 7.5-8.5 mm
long, purplish. Ovary appressed-hairy. Pods straight, cylindrical, 27^4 mm long, 2-3 mm
wide, moderately densely pubescent; endocarp spotted. Seeds 6-10 per fruit,
cylindrical to almost cubic, 1.5-2 mm long and wide; testa dark brown to light olive,
the surface smooth to rough.
Phenology: flowers mostly in Summer; fruits mostly Autumn.
Distribution and habitat: This species is restricted to an area west of Atherton
Tableland with the known distribution extending from “Nychum”, NNW of Chillagoe,
south to the Mount Surprise area (ie, between c. 16°50' and 18°10'S). It is recorded
from savannah woodland and open forest on red or grey gravelly soils.
Notes: As the epithet ‘coriacea’ is pre-occupied, a new epithet is proposed. The
proposed epithet, ‘scabrella’, is taken from the Latin meaning ‘minutely scabrous’, a
reference to the leaflets being slightly rough to the touch.
Domin described this taxon as having 5-7 leaflets but leaves with 5 leaflets are
uncommon, except for the early leaves of a new season's growth.
Other specimens examined: Queensland: Cook: 13 km from the Walsh River crossing on the road
to Wrotham Park, Clarkson 4255, 14 Jan 1982 (BRI, CANB, NSW, UNSW); Mungana Red Hill,
Cole 5114, 8 Aug 1963 (BRI); Petford-Chillagoe Rd, 11 km W of Almaden., Hacker 397,
16 July 1983 (BRI); 5 km W of Chillagoe, Jackes 9954, 9956, May 1999 (JCT, NSW); 22 km W of
Walsh river, Kazandjian AK0012Au, AK0013Au dr Dowe, 14 May 2001 (JCT, NSW); Nychum
holding on F.lizabeth Creek near homestead, Macdonald 1,4 Dec 1970 (BRI, CANB, MEL, NSW);
29 km W of Mount Surprise, Puttock UNSW 13439, 20 Apr 1982 (NSW, UNSW); Lyndbrook,
Etheridge Line, Towers s.n., 11 Apr 1960 (BRI); 0.8 km E of Mungana Yards, Wilson UNSW 13388
dr Puttock, 17 Apr 1982 (NSW, UNSW); 10.5 km E of Chillagoe, Wilson UNSW 13404 dr Puttock,
17 Apr 1982 (NSW, UNSW); 18.1 km E of Petford, Wilson UNSW 13407 & Puttock, 17 Apr 1982
(NSW, UNSW); 73 km from Almaden on road to Mt. Surprise, 5 km past Bullock Creek Station
turnoff, Forster 9635 ,29 Jan 1992 (BRI, NSW).
Cultivated specimens: James Cook University, Townsville (seed ex Jackes 9954 ) Kazandjian s.n.,
15 Jan 2002 (NSW 705631); James Cook University, Townsville (seed ex Wilson UNSW 13388 dr
Puttock) Kazandjian s.n., 15 Jan 2002 (NSW 705606).
50
Telopea 11(1): 2005
Kazandjian and Wilson
Indigofera pratensis var. angustifoliola
The name I. pratensis var. angustifoliola Domin was based on specimens, collected near
Gladstone by Amalie Dietrich, that were characterised by narrow leaflets 15-20 mm
long and 2.75-3.75 mm wide. When Mueller (1860) described I. pratensis, from
specimens collected in the Burdekin area, he gave the leaflet dimensions as 10-40 mm
long and 6-13 mm wide. More recent measurements, over the geographic range of the
Indigofera pratensis sens, str., indicate that the leaflet size is commonly in the range
10—30(—44) mm long and 5-15(-24) mm wide (Wilson & Rowe, unpublished data).
When specimens that had been collected from the Queensland coast between 20° and
25°S (ex BRI, CANB, MBA, MEL and NSW) were examined, only one had leaflets that
matched Domin’s description. This was a specimen (MEL586535) collected by Eugene
Fitzalan in 1874 at Port Denison (Bowen), a considerable distance north of Gladstone.
Apart from this, the only other specimen to approach the Dietrich collection is one of
Robert Brown’s found in the vicinity of Keppel Bay (near Rockhampton, just north of
Gladstone); this appears to be a depauperate plant with leaflets 3-5.2(-6) mm wide.
A field trip to the Gladstone area did not locate any living plant with leaflets matching
Domin’s description, although plants of typical Indigofera pratensis were commonly
encountered. These observations suggest that Dietrich (and Brown and Fitzalan) may
have collected the specimens in a dry year when the size of the leaflets was affected by
the availability of water. I. pratensis occurs in open Eucalyptus woodland in eastern
Queensland where water and nutrient availability vary markedly from season to
season. Field observations of other species suggest that features such as plant height,
leaflet number and size, pedicel and petiolule length, and fruit size are likely to be
affected by such environmental conditions. Hence, we judge var. angustifoliola to be an
environmentally induced variant not worthy of recognition.
Typification
Domin listed two Dietrich collections in the protologue of this species, so
lectotypification is required.
Indigofera pratensis var. angustifoliola Domin, Biblioth. Bot. 89 (3): 191 (1926)
Type citation: ‘Gladstone, A. Dietrich s.n. und n. N. 16a, beide als I. macrophylla
Venten. bestimmt.’
Lectotype (designated here): Queensland: Gladstone, A. Dietrich s.n. (PR 527229)
Fig. 4. Isolectotype: HBG.
Residual syntype: Queensland: Gladstone, A. Dietrich 16a (PR 527230, HBG).
At PR, the two syntypes are mounted on a single sheet but with separate numbers; the
lectotype is the upper specimen on the sheet. The typed label calls this taxon var.
angustifolia rather than angustifoliola, but there is no way of knowing if this was an
error or if Domin changed his mind about the epithet when publishing the name.
Also, Domin erred in the type citation since Dietrich had, in fact, determined the
species as I. macrostachya Vent, (a synonym of the Himalayan species, Indigofera
hctcrantha Wall, ex Brandis) not I. macrophylla Schumach. & Thonn., an African species.
Indigofera pratensis complex
Telopea 11(1): 2005
51
Acknowledgments
This study represents a small part of a wider analysis of variation within the I. pratensis
complex undertaken by the first author at James Cook University, Townsville. She is
very appreciative of the financial support received from Fondo Nacional de Ciencia,
Tecnologia e Innovacion (FONACIT) in Caracas, Venezuela. She is also grateful for the
assistance and guidance given by Betsy R. Jackes (JCT), and thanks John Dowe and
Juan Cruz for their help with locating and collecting specimens in the field. The second
author is grateful to the Australian Biological Resources Study (ABRS) for grant
funding that assisted in the earlier stages of this study. Both authors thank the relevant
herbaria (PR, FIBG, BRI, MBA, NSW, UNSW) for the loan of specimens.
References
Domin K (1926) Indigofera. Bibliotheca Botanica 89 (3): 741-746.
Hewson H (1988) Plant Indumentum. Australian Flora 8c Fauna Series No. 9. (Australian
Government Publishing Service: Canberra)
Kazandjian AA (2002) Systematics of the Indigofera pratensis Complex (Fabaceae): a Morphological
and Molecular Approach. (Unpublished PhD thesis, James Cook University, Townsville)
Mueller F (1860) Essay on the plants collected by Mr. Eugene Fitzalan, during Lieut. Smith’s
expedition to the estuary of the Burdekin. (Government Printer: Melbourne)
Prabhakar M, Vijay Kumar BK, Ramayya N 8t Leelavathi P (1985) Structure, distribution and
taxonomic significance of trichomes in some Indigofera L. (Fabaceae). Proceedings of the
Indian Acadademy of Science (Plant Science) 95: 309-314.
Schrire BD (1995) Evolution of the tribe Indigofereae (Leguminosae Papilionoideae). Pp. 161-244
in Crisp MD 8c Doyle JJ (eds), Advances in Legume Systematics, vol. 7. (Royal Botanic Gardens:
Kew)
Manuscript received 1 September 2004, accepted 12 May 2005
t
Telopea 11(1): 53-57
Lectotypification of Schoenodum tenax
(Restionaceae) and a note on the type of
Lyginia imberbis (Anarthriaceae)
Barbara G. Briggs
National Herbarium of New South Wales, Mrs Macquaries Road, Sydney, NSW 2000, Australia;
email: barbara.briggs@rbgsyd.nsw.gov.au
Abstract
Robert Brown in 1810 made clear that the original description of Schoenodum tenax Labill. was
based on collections of more than one taxon. The male material is the species currently known
as Lyginia imberbis R. Br. but the female specimens include both the species currently known as
LeptOcarpus tenax (Labill.) R. Br. (basionym S. tenax) and Apodasmia brownii (Hook, f.) B.G. Briggs
& L.A.S. Johnson (basionym Leptocarpus brownii). A female specimen in the Willdenow
Herbarium in Berlin (B), of the species currently known as Leptocarpus tenax, is designated as
lectotype of S. tenax; this matches the illustration and text of the protologue and preserves current
usage. The type specimen of Lyginia imberbis R. Br. is a lectotype rather than a holotype.
Introduction
In a recent publication, Briggs and Johnson (1998) divided the then recognised species
of Leptocarpus among four existing and three new genera. In that work we maintained
the name L. tenax for the commonest, most widespread species and the one that
epitomises Leptocarpus for most Australian biologists. There we pointed out that this
use could only be maintained if the type of the conserved name Leptocarpus R. Br. were
changed. Later I made a formal proposal (Briggs 2001) to change the conserved type
of Leptocarpus from L. aristatus R. Br. to L. tenax (Labill.) R. Br. Since that proposal was
made, it has become clear that there are additional issues concerning the typification
of Leptocarpus tenax and Lyginia imberbis.
Lectotypification of Schoenodum tenax
Labillardiere (1806) based his description of Schoenodum and S. tenax on material of
both male and female plants, and both sexes are illustrated in his Tab. 228. 1 have now
examined the relevant Labillardiere specimens in Florence (FI) and Paris (P) and a
microfiche of sheet 18267 in the Willdenow herbarium in Berlin (B). The male
material in FI belongs to the taxon currently known as Lyginia imberbis R. Br.
Labillardiere’s female specimen in FI (sheet 188031 of the Webb Herbarium) is
referable to the taxon currently known as Apodasmia brownii (Hook, f.) B.G. Briggs &
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Briggs
L.A.S. Johnson (=Leptocarpus brownii Hook. f.). However, the female material in P and
B is referable to the species to which the name Leptocarpus tenax R. Br. is presently
applied. These specimens are thus considered to represent three taxa, so that only
lectotypification can fix the application of the names Schoenodum and S. tenax.
Labillardiere’s statement in the protologue of the source of the collections is ‘Habitat
in capite Van-Diemen’, a designation given to collections from several locations in
Tasmania. This appears to be intended to apply to both the male and female
specimens, but only the females could have been collected there. Nelson (1974) has
drawn attention to a number of specimens of Western Australian endemics cited by
Labillardi&re that are labelled capite van-Dieman’. Using the names currently in use,
Lyginia imberbis is a Western Australian endemic that is common on the southern
coast which was visited on the voyage of d’Entrecasteaux, when Labillardiere’s
collections were made; Apodasmia brownii occurs in Tasmania and Victoria; while
Leptocarpus tenax is found in Tasmania, the south of Western Australia and widely
distributed in eastern Australia.
In determining whether previous authors have lectotypified S. tenax, the following
four publications are relevant.
(1) Brown (1810) was the first to recognize that Labillardiere’s material of
Schoenodum tenax included more than one taxon. In the Prodrotnus, Brown
adopted the name L. tenax for the seventh species of his genus Leptocarpus and
cited Schoenodum tenax femina. Labill. nov. holl. 2. t. 229’ under his entry for that
species. In the protologue of Lyginia imberbis he cited 'Schoenodum tenax mas.
Labill. nov. holl. 2. t. 229’. Brown also referred to his own material, using his
customary style: ‘(M.) v.v.’ and ‘(M. J. D.) v.v.’ respectively. Since he had not
examined Labillardiere’s specimens (D. Mabberley pers. comm.). Brown’s citation
of Labillardiere would refer to the excellent illustrations, rather than directly to
Labillardiere s specimens. Also, Brown did not use the word ‘type’, or equivalent.
(2) In his Enumeratio Plantarum, Kunth (1841) widened the circumscription of
Schoenodum, equating it with ‘Leptocarpi species Brown’ and excluded
Labillardiere’s male plant. Although he referred specifically to Labillardiere’s
collection in the Willdenow herbarium he did not formally refer to it as lectotype.
(3) Rickett 8c Stafleu (1959), in their review of Appendix III of the International
Code of Botanical Nomenclature (Paris 1956), state about Lyginia, ‘Conservation
superfluous: Schoenodum Labillardiere ... need not be rejected because its type
S. tenax is typified by the female plant of Labillardiere, which is Leptocarpus R.
Brown ... and not Lyginia. [In reference to the conserved name Leptocarpus,
Rickett & Stafleu listed the type of the rejected name Schoenodum as 'S. tenax
Labillardiere ( vide Kunth, Enum. 3: 445. Jul 1841)’, a reference that has persisted to
the current edition of the ICBN.]
(4) Johnson and Evans (1966) observed that ‘The female specimen was, in effect,
selected as lectotype by R. Brown when he based L. tenax upon it’.
None of the publications mentioned effected lectotypification since they did not
diffeientiate between the two taxa of Labillardiere’s female material or, in some cases,
distinguish between Labillardiere’s female material and his illustration. Of these
references, (1) and (2) did not use the term ‘type’, whereas (3) and (4) did not refer to
a gathering of a single species.
Schoenodum tenax & Lyginia imberbis
Telopea 11(1): 2005
55
In the Webb Herbarium at FI there arc male and female specimens relevant to S. tenax
mounted separately on two sheets. Sheet FI 188030 bears flowering male culms,
identifiable as Lyginia imberbis, and has a packet attached to the upper right corner
labelled ‘foem. flores’, this contains an inner packet labelled ‘semen cum fragmentiis
capsulae’. The two pieces of material in the inner packet, however, appear to be
staminal columns from male flowers of a Lyginia species. Four small handwritten
sheets in Labillardiere’s hand are pinned to the sheet of this male material (FI 188030);
the writing on two of these matches the text of the protologue of Schoenodum (p. 79)
and S. tenax (p. 80); the other two are descriptive notes that appear to be a first draft
of the description. Sheet FI 188031 bears groups of slender, unbranched culms, with
three of the culms terminating in female inflorescences referrable to what is now called
Apodasmia brownii. Also attached to FI 188031 are (1) a packet containing numerous
female flowers of A. brownii, (2) a small packet labelled ‘masculi flores’ which contains
male flowers of Lyginia and (3) a note ‘facies chondropetali rotb.’ in Labillardiere’s
hand.
The relevant sheet in P, of the taxon currently known as L. tenax, has a handwritten
label (but not by Labillardiere) ‘ Schoenodum tenax ’ and a printed label ‘Van Diemen
Labillardiere Donne par M. Webb’. It bears a female plant with several culms and two
inflorescences.
Sheet number 18267 in the Willdenow herbarium in B also bears a female specimen of
the taxon currently known as L. tenax. It is labelled, in Labillardiere’s hand,
‘Schoenodum tenax <$’ (sic.). The right-hand piece shows an inflorescence; the left-
hand piece is only part of a culm, but its culm sheaths, and the ascending curve of a
culm arising from a rhizomatous base, identify it as the same taxon.
It is now clear that Labillardiere’s female material is a mixture of two species, one
represented by the material in FI and the other by the specimens in B and P. 1 hese are
sufficiently similar to have been taken as conspecific (especially in the context of the
male Lyginia material also being considered conspecific) when parts of the collections
were chosen for close study and when duplicates were distributed. The description of
vegetative structures may be based on all of the original material, of the three taxa, but
with emphasis on the structures of the male, Lyginia, as in the description of the roots.
However, it appears that material of only one of the female species was examined in
detail and illustrated. The protologue of S. tenax describes the female inflorescence as
‘panicula contracta, palmaris, spiculis elliptico-oblongis, sessilibus pedunculatisque,
imbricatis squamis ovato-oblongis, acuminatis, unifloris.’ The reference to elliptic-
oblong, single-flowered pedunculate spikelets is consistent with Labillardiere s
illustration (which shows a female inflorescence of distinct, elongated spikelets) and
with the specimens in both B and P. These features are characteristic of L. tenax, as that
name is presently applied, and are not shown by the taxon known as A. brownii, which
has much more condensed inflorescences, the spikelets not clearly distinguishable
within the densely aggregated compound inflorescences, and the small flower-clusters
each multiflowered.
Although the FI specimen is in the herbarium that houses Labillardiere’s main
collection, it does not agree with the description and illustration of the female
inflorescence in the protologue, which therefore cannot have been based upon it.
Moreover, if the FI female specimen were chosen as lectotype, this would change the
application of the names Leptocarpus tenax and Apodasmia brownii, causing confusion
in the naming of two widespread and ecologically important species.
56
Telopea 11(1): 2005
Briggs
The female specimens in B and P agree with the description and illustration of the
protologue (in accord with Article 9.17 of the ICBN) and choice of either of them
would preserve current usage of the names Leptocarpus and L. tenax. Only the
specimen in B has been annotated by Labillardiere, so this would appear the obvious
choice among them. I therefore here designate as the lectotype of Schoenodum tenax
Labill. the specimen on sheet 18267 of the Willdenow herbarium in B, which is female
plant material of the taxon currently known as Leptocarpus tenax. A photograph of the
lectotype is included in the microfiche set of the Willdenow herbarium (Inter
Documentation Company microfiche set 7440). This lectotypification preserves the
usage established by other authors who identified Schoenodum tenax with
Labillardiere’s female gathering(s).
The lectotype of Lyginia imberbis R.Br.
Lyginia imberbis R. Br„ the species to which Labillardiere’s male material is referred, is
one of three species of the sole genus of Lyginiaceae (Briggs & Johnson 2000), or a
member of one of three genera of Anarthriaceae if a more inclusive family concept is
adopted (Chase et al. 2000, Bremer 2002, APG II 2003), by which the Anarthriaceae
encompasses the Anarthria clade of Briggs et al. (2000). Since the original material
consists of Brown’s collection and also Labillardiere’s illustration,lectotypification was
necessary. Briggs & Johnson (2000) overlooked the need for lectotypification and cited
a sheet of Brown’s collection, bearing both male and female plants, as the holotype.
That incorrect use of a term to describe the type is an error to be corrected (ICBN Art.
9.8); it did not prevent the action of specifying a type among the original material
effecting lectotypification. Thus the lectotype (designated [as ‘holo’J by Briggs &
Johnson, Telopea 8: 496, 2000) is the specimen King George III d S d [Sound], R. Brown
(Bennett No. 5837), 1802-5 (BM, S , 9 mounted together on one sheet, annotated by
Brown ‘Rest/o’, isolectotypes E, K, P).
Acknowledgments
Sincere thanks go to Gillian Perry for drawing my attention to relevant issues, many
very helpful discussions of typification concerning these names and taxa, and
numerous valuable comments on this manuscript. Dr David Mabberley kindly assisted
with information concerning Robert Brown and Labillardiere. Dr. P. Cuccuini
provided photographs, a scan of Labillardiere’s specimens in FI and access to
specimens in that herbarium. The assistance of G. Padovani, C. Nepi and A. Guiliano
at FI, P. Morat and M. Pignal at P, J. Thurlow at MEL and M. Garcia at NSW is
gratefully acknowledged.
Schoenodum tenax & Lyginia imberbis
Telopea 11(1): 2005
57
References
APG II (2003) An update of the Angiosperm Phylogeny Group classification for the orders and
families of flowering plants: APG II. Bol. ]. linn. Soc. 141: 399-436.
Bremer K (2002) Gondwanan evolution of the grass alliance of families (Poales). Evolution
56:1374-1387.
Briggs BG (2001) (1489) Proposal to conserve the name Leptocarpus (Restionaceae) with a
conserved type. Taxon: 50: 919-921.
Briggs BG & Johnson LAS (1998) New genera and species of Australian Restionaceae (Poales).
Telopea 7: 345-373.
Briggs BG & Johnson LA (2000) Hopkinsiaceae and Lyginiaceae, two new families of Poales in
Western Australia, with revisions of Hopkinsia and Lyginia. Telopea 8:477-502.
Briggs BG, Marchant AD, Gilmore S 8c Porter CL (2000) A molecular phylogeny of Restionaceae
and allies. Pp. 661-671 in KL Wilson 8c D Morrison (eds) Monocots-Systematics and
Evolution.! CSIRO: Melbourne)
Brown R (1810) Prodromus Florae Novae Hollandiae et Insulae Van Diemen
Chase MW, Fay MF 8c Savolainen V (2000) Higher-level classification in the angiosperms: new
insights from the perspective of DNA sequence data. Taxon 49: 685-704.
Kunth CS (1841) Enumeratio Plant arum, vol. 3.
Labillardiere JJH de (1806) Novae Hollandiaeplantarum specimen, part 23.
Nelson EC (1974) The locations of collection and collectors of specimens described by
Labillardiere in 'Novae Hollandiae Plantarum Specimen' - additional notes. Pap. 8c Proc. Roy.
Soc. Tasmania 108: 159-170.
Rickett HW 8c Stafleu FA (1959) Nomina generica conservanda et rejicienda spermatophytorum.
Taxon 8: 213-243.
Manuscript received 18 October 2004, accepted 19 June 2005
Telopea 11(1) 59-78
New combinations and synonymies in the
Australian Graphidaceae
A.W. Archer
Botanic Gardens Trust Sydney, Mrs Macquaries Road, Sydney NSW 2000, Australia
Abstract
Species of the Australian Graphidaceae have been re-allocated to the genera proposed in a recent
revision of the family. Forty-one new combinations are made, a new name, Grapliis elixiana, is
published, and the following synonymies are proposed: Diorygma erythrellum replaces Graphina
atramontana ; Graphis leucoparypha replaces G. turgidula var. norstictica ; Hemithecium
chlorocarpoides replaces Graphina repleta var. monospora; Hemithecium chrysentcron replaces
Graphina repleta-, Leiorreuma hypomelaenum replaces Phaeographis necopinata; Phaeographis
lindigiana replaces Phaeographis pseudomelana, and Platygramme pudica replaces Phaeographina
echinocarpica. Keys to the genera and species in Australia are given.
Introduction
The lichen family Graphidaceae has recently been rearranged, with new genera
described, some older genera resurrected and existing genera revised (Staiger & Kalb
1999; Staiger 2002; Kalb, Staiger & Elix 2004). This re-arrangement necessitates a
number of name changes, new combinations and new synonymies in the Australian
Graphidaceae which are listed below, together with keys to the taxa in each genus.
As a result of this revision, the following genera are now known from Australia:
Acanthothecis Clem., Carbacanthographis Staiger & Kalb (new), Diorygma Eschw.,
Dyplolabia A. Massal., Fissurina F£e, Glyphis Ach., Graphis Adans. (revised),
Hemithecium Trevis., Leiorreuma Eschw., Phaeographis Miill. Arg. (revised),
Platygramme Fee, Platythecium Staiger (new), Sarcographa Fee, Sarcographina Mull.
Arg., Thalloloma Trevis., Thecaria Fee. The genus Phaeographina Mull. Arg. has also
been revised but there are now no Australian taxa in the revised genus.
The genus Gymnographa Miill. Arg. is rejected as it is based on an old specimen of
Phaeographis eludens (Stirt.) Shirley with degenerate ascospores; consequently the
genus Sarcographina Mull. Arg., which had been reduced to synonymy with
Gymnographa, is retained.
Many specimens, including a large number of type specimens, were examined in
detail, some for the first time, in the course of this revision of the Graphidaceae
(Staiger 2002); the majority were from one collector and biased in favour of South
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Archer
American material but few Australian specimens were examined. Consequently, the
circumscription of the genera fits the specimens examined and some of the
combinations made below must be regarded as tentative until the revised genera are
better defined.
New synonymies and new reports
In addition to the synonymies recently described (Archer 2004, 2005), the following
synonyms are reported.
Graphina atramontana A.W. Archer is morphologically indistinguishable from
Diorygma erythrellutn (Mont.) Kalb, Staiger & Elix and the two species are considered
to be synonymous.
Graphina pertenella (Stirt.) Shirley and Graphina laevigata (Mull. Arg.) A.W. Archer
possess the same chemistry and similar ascospores and differ only in the degree to
which the lirellae are open. The two species are considered to be synonymous and are
here combined under the earlier name and transferred to the genus Platythecium as
P. pertenellum.
The ascospores in Graphina repleta (Stirt.) Shirley are usually hyaline, with some
gradually becoming pale brown, but all give a red-brown colour with iodine, in
contrast to the hyaline ascospores present in other Graphina species which usually give
a blue or blue-violet colour. Stirton reported the ascospores to react brownish blue
“sporae caerulo-infuscatae” (Stirton 1881). This colour reaction of the ascospores with
iodine, the pale reddish-brown exciple and the presence of stictic acid are identical to
those of Hemithecium chrysenteron (Mont.) Trevis.; the two species are here reduced to
synonymy. Similarly, Graphina repleta var. monospora A.W. Archer, with larger
ascospores, is identical with Hemithecium chlorocarpoides (Nyl.) Staiger, a species
originally described from Java and recently reported from Australia (Staiger 2002). The
two Hemithecium species differ only in the size of the ascospores and may be
synonymous (Staiger op. cit.).
Phaeographina echinocarpica A.W. Archer & Elix is a later name for Platygramme
puciica (Mont. & Bosch) M. Nakan. & Kashiw (Nakanishi et al. 2003). The chemistry of
the latter species was originally reported as’an unknown substance’ (Nakanishi 1977)
but the compound was recently identified as echinocarpic acid (Nakanishi et al. 2003)
so P. pudica is identical to P. echinocarpica.
The morphology and chemistry of Phaeographis necopinata A.W. Archer & Elix are
identical to those of Leiorreuma hypomelaenum (Mull. Arg.) Staiger, recently reported
from Australia. Both contain the uncommon hypostictic acid as the major lichen
compound.
Phaeographis pseudomelana Miill. Arg. is indistinguishable from Phaeographis
lindigiana Miill. Arg., recently reported from Australia.
In addition, the following species in the Graphidaceae have recently been reported
from Australia:
Phaeographis brasiliensis (A.Massal.) Kalb & Matthes-Leicht, P. hypomelaena Miill.
Arg., P. lindigiana Miill. Arg., P. lobata (Eschw.) Miill. Arg. and P. platycarpa Miill. Arg.
New combinations in Australian Graphidaceae
Telopea 11(1): 2005
61
(Kalb 2001) and Hemithecium chlorocarpoides (Nyl.) Staiger [Phaeographina
chlorocarpoides (Nyl.) Zahlbr.] (Staiger 2002).
The key to the genera is adapted from Staiger (op. tit., pp. 62-67) with genera not
found in Australia omitted. Detailed descriptions of the genera are also given in Staiger
(op. tit.).
Key to genera of Graphidaceae in Australia
la. Exciple with distinctly carbonised areas . 2
lb. Exciple uncarbonised with, at the most, small brownish areas . 15
2a. Mature ascospores hyaline, 1+ blue or blue-violet, or I-ve. 3
2b. Mature ascospores brown or brownish, 1+ red, red-brown or red-violet . 9
3a. Lirellae with white powdery cover containing lecanoric acid (C+ red) . Dyplolabia
3b. Lirellae lacking a white powdery cover, or, if present, lacking lecanoric acid (C-ve) . 4
4a. Hymenia in well-developed carbonised stromata; discs open, brownish granular Glyphis
4b. Hymenia not in well-developed carbonised stromata; discs closed or, if open, not brownish
granular ... 5
5a. Labia or exciple divergent; discs visible in surface view. 6
5b. Labia or exciple convergent; discs completely covered by the labia. 7
6a. Carbonisation restricted to the base of the exciple, lateral exciple poorly developed;
ascospores 20 pm long; testacein A and/or B present. Platythecium
6b. Lateral exciple and labia well-developed; ascospores > 20 pm long; lichen compounds absent
. Glyphis sub. gen. Pallidoglyphis
7a. Labia carbonised, often completely, and convergent, with a thalline cover or a white pruinose
layer; ascospores 1- or 1+ weak violet . Carbacanthographis
7b. Carbonised layer lacking a white pruinose layer; ascospores 1+ blue-violet. 8
8a. Lirellae fissurine, apically or laterally carbonised; ascospores ovoid-ellipsoid, 4-locular or
muriform, with or without halo . Fissurina
8b. Lirellae not fissurine; labia distinctly developed and carbonised or, if fissurine, ascospores
not ovoid but elongate and lacking halo . Graphis
9a. Carbonised exciple and excipular labia usually well-developed . 10
9b. Carbonised exciple lacking thalline cover. Phaeographina
10a. Hypothecium becoming carbonised with age, giving a thick carbonised base. 11
10b. Hypothecium not becoming carbonised with age but base may be carbonised . 14
11a. Lirellae embedded in stromata [raised, paler, whitish areas] . 12
1 lb. Lirellae not embedded in stromata but may be crowded or branched . 13
12a. Ascospores septate with lenticular locules. Sarcograplia
12b. Ascospores muriform . Sarcographina
13a. Proper margin and lateral exciple well-developed, discs open, ± sunken, red or white
pruinose [ascospores in known species muriform] . Thecaria
13b. Proper margin and lateral exciple poorly developed or, if well-developed upper part not
covered by thalline layer; discs not sunken, brown to black, epruinose or weakly pruinose
[ascospores in known species transversely septate] . Leiorreuma
62
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14a.
14b.
15a.
15b.
16a.
16b.
17a.
17b.
18a.
18b.
19a.
19b.
20a.
20b.
21a.
21b.
22a.
22b.
23a.
23b.
24a.
24b.
Proper margins well-developed, convergent, apices wedge-shaped and carbonised, or
laterally carbonised; disc sunken and white- or greyish white pruinose; ascocarps large and
prominent . Platygramme
Proper margins poorly developed and weakly carbonised, ± brown, divergent or well-
developed but discs not concealed and not pruinose . Phaeographis p.p.
Ascospores brown . 16
Ascospores hyaline. 18
Labia well-developed, convergent, silicate, disc not visible [stictic acid].
. Hemithecium p.p.
Labia poorly developed, no sulcate, not convergent; discs ± open but narrow or margins
well-developed and slightly striate; discs visible and distinctly open . 17
Ascospores ovoid, < 20 pm long, 4 x 1-2 -locular, discs open, brownish black, epruinose
. Platythecium p.p.
Ascospores elongate, > 20 pm long, > 4 x 1-2 -locular; if ascospores ovoid, then discs not
brownish black and epruinose . Phaeographis p.p.
Paraphysis tips warty . 19
Paraphysis tips not warty. 20
Ascospores ovoid or globose, ± halonate; lirellae fissurine; exciples and margins poorly
developed. Fissurina p.p.
Ascospores elongate; lirellae not fissurine. Acanthothecis
Labia well-developed, crenate, convergent; disc slit-like, not visible, completely concealed
by margins; apothecia raised front thallus . 21
Labia poorly developed or not distinctly convergent, with discs ± open; apothecia usually
not raised from thallus. 22
Ascospores ovoid, 4 -locular or muriform ± halonate . Fissurina p.p.
Ascospores lacking halo, 1+ blue-violet, > 25 pm long, > 6 -locular; labia often distinctly
crenate . Hemithecium p.p.
Apothecia fissurine; thalline margins project over disc; ascospores ovoid ± halonate, 1+ weak
blue or I -ve, rarely 1+ blue violet . Fissurina p.p.
Apothecia otherwise; ascospores distinctly 1+ blue or blue violet. 23
Ascospores small, < 20 pm long, 4-5 x 1-2-locular . Platythecium
Ascospores larger, > 20 pm long . 24
Discs open, sometimes ± narrow, brown or reddish; paraphysis tips brown, granular nor
stictic and stictic acids absent . Thalloloma
Discs open, distinctly white pruinose; norstictic or stictic acids may be present .
. Diorygma
Keys to species of Graphidaceae in Australia
Acanthothecis
la. Thallus saxicolous; ascospores 19-22 pm long, 4-locular [in Australian specimen]
. A. silicicola
lb. Thallus corticolous; ascospores muriform . 2
2a. Ascospores 20-30 pm long; norstictic acid present . A. subaggregans
2b. Ascospores 14-24 pm long; stictic acid present. A. gyridia
New combinations in Australian Graphidaceae Telopea 11(1): 2005 63
Carbacanthographis
la. Exciple laterally carbonised; ascospores 12-17 pm long, 4-5 x 2 -locular .
. C. marcescens
lb. Exciple completely carbonised; ascospores 19-23 pm long, 8 X 1-2 -locular.
. C. salazinica
Diorygma
la. Ascospores septate with lenticular locules; norstictic acid only present . 2
lb. Ascospores muriform; norstictic acid and/or other depsides present . 3
2a. Ascospores 60-90 pm long. D. circumfusum
2b. Ascospores 45-55 pm long. D. wilsonianum
3a. Ascospores < 80 pm long. 4
3b. Ascospores > 80 pm. 5
4a. Ascospores 30-65 pm long. D. erythrellum
4b. Ascospores 18-23 pm long. D. nothofagum
5a. Stictic acid present; ascospores 95-150 pm long. D. hieroglyphicum
5b. Norstictic or protocetraric acid present . 6
6a. Norstictic acid only present; ascospores 80-105 pm long . D.junghuhnii
6b. Protocetraric acid ± norstictic acid present . 7
7a. Protocetraric acid only present; ascospores 95-150 pm long. D. pruinosutn
7b. Protocetraric and norstictic acid present; ascospores 120-150 pm long.
. D. rufopruinosum
Fissurina
la. Thallus saxicolous . 2
lb. Thallus corticolous. 3
2a. Ascospores 14-16 pm long, 4 -locular; psoromic acid absent . F. howeana
2b. Ascospores 16-20 pm long, 4x2 -locular; psoromic acid present.F. streimannii
3a. Ascospores muriform, 4-6 x 1-3 -locular . 4
3b. Ascospores 4 -locular. 6
4a. Lichen compounds absent; ascospores 21-28 pm long, 4-5 x 2 -locular. F. elaiocarpa
4b. Lichen compounds present . 5
5a. Ascospores 28-35 pm long; stictic acid present. F. abdita
5b. Ascospores 8-14 pm long; 2-methoxypsoromic acid present . F. globulifica
6a. 2-Methoxypsoromic acid present.7
6b. Lichen compounds absent. 8
7a. Proper exciple laterally carbonised; ascospores 16-20 pm long. F. elixii
7b. Proper exciple uncarbonised; ascospores 16-26 pm long. F. paradoxica
8a. Lirellae inconspicuous, visible only as a slit. 9
8b. Lirellae conspicuous, raised; ascospores 15-22 pm long. F. insidiosa
9a. Proper exciple laterally carbonised; ascopores 10-11 pm long . F. albonitens
9b. Proper exciple uncarbonised . 10
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10a. Lirellae (slit) with thin black margin; ascospores 18-22 pm long.
. F. ttigririmis var. deficiens
10b. Lirellae lacking black margins; ascospores 11-20 pm long. F. dumastii
Glyphis
la. Ascospores septate with lenticular locules, 30-60 pm long, 8-13 -locular .. G. cicatricosa
lb. Ascospores muriform . 2
2a. Lirellae raised from the thallus; ascospores 30-45 pm long, 8-10 X 2-4 -locular .
. G. scyhpuliferum
2b. Lirellae not raised from thallus; ascospores 40-50 pm long, 12-14 x 2-5 -locular .
. G. montoensis
Graphis
la. Ascospores septate with lenticular locules . 2
lb. Ascospores muriform . 38
2a. Lirellae immersed. 3
2b. Lirellae not immersed . 9
3a. Proper exciple completely carbonised. 4
3b. Proper exciple laterally or apically carbonised . 6
4a. Norstictic acid present; ascospores 30-44 pm long, 8-12-locular . G.inamoena
4b. Lichen compounds absent. 5
5a. Ascospores 20-25 pm long . G. immersicans
5b. Ascospores 55-65 pm long, 13-16 -locular . G. propitiqua
6a. Proper exciple apically carbonised; lichen compounds absent; ascospores 38-55 pm long,
9- 12-locular . G.sayeri
6b. Proper exciple laterally carbonised; lichen compounds present . 7
7a. Lichexanthone and norstictic acid present; ascospores 15-20 pm long, 5-6 -locular
... G. stipitata
7b. Stictic acid present . 8
8a. Ascospores 50-65 pm long, 10-15 -locular. G. crassilabra
8b. Ascospores 24—35 pm long, 8—11 -locular. G. immersella
9a. Lirellae open; norstictic acid present. 10
9b. Lirellae closed; lichen compounds present or absent . 12
10a. Proper exciple completely carbonised; ascospores 20-34 pm long, 6-8 -locular.
. G. semiaperta
10b. Proper exciple laterally carbonised . 11
11a. Ascospores 28-40 pm long, 8-11 -locular . G. apertella
lib. Ascospores45-60 pm long, 10-16-locular . G. streimannii
12a. Lirellae sulcate . 13
12b. Lirellae not sulcate . 19
13a. Norstictic acid present; proper exciple laterally carbonised; ascospores 60-72 pm long,
10- 14-locular. G.elegans
13b. Norstictic acid absent. 14
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65
14a. Lichen compounds absent. 15
14b. Stictic acid present . 18
15a. Proper exciple completely carbonised; ascospores 40-55 pm long, 8-13 -locular .
. G. rimulosa
15b. Proper exciple laterally carbonised . 16
16a. Ascospores 36-50 pm long, 9-14-locular . G.leptoclada
16b. Ascospores <35 pm long. 17
17a. Lirellae large, conspicuous, predominantly simple, terminally rounded; ascospores 23-35 pm
long, 6-8 -locular . G. endoxantha
17b. Lirellae small, inconspicuous, branched, terminally acute; ascospores 21-34 pm long
8-10-locular .. G. subtenella
18a. Proper exciple laterally carbonised; ascospores 26-40 pm long, 7-10 -locular .
... G. stenotera
18b. Proper exciple apically carbonised; ascospores 40-55 pm long, 10-16 -locular.
. G. treubii
19a. Lichen compounds absent. 20
19b. Lichen compounds present . 28
20a. Proper exciple completely carbonised. 21
20 b. Proper exciple apically or laterally carbonised . 23
21a. Lirellae weakly silicate; ascospores 62-80 pm long, 13-16 -locular. G. longula
21b. Lirellae smooth. 22
22a. Ascospores 28—42 pm long, 8-11 -locular . G. anfractuosa
22b. Ascospores 50—65 pm long, 12—15-locular . G. catherinae
23a. Proper exciple apically carbonised . 24
23b. Proper exciple laterally carbonised . 25
24a. Lirellae inconspicuous; ascospores 40-60 pm long, 10-12-locular . G.epimelaena
24b. Lirellae conspicuous; ascospores 30-40 pm long, 8-10-locular . G. xanthospora
25a. Ascospores > 50 pm long. 26
25b. Ascospores < 50 pm long. 27
26a. Lirellae 1-3 mm long; ascospores 54-70 pm long, 12-16 -locular .
. G. stenospora var. deficiens
26b. Lirellae short, simple, < 1 mm long; ascospores 50-60 pm long, 12-14 -locular.
. G. subregularis
27a. Ascospores 20-30 pm long, 6-8 -locular . G. tenella (auct.)
27b. Ascospores 30-41 pm long, 10-12-locular . G.albissima
28a. Norstictic acid present . 29
28b. Stictic or protocetraric acid present . 35
29a. Proper exciple completely, or almost completely, carbonised . 30
29b. Proper exciple laterally carbonised; ascospores 40-50 pm long, 10-14 -locular.
. G. vinosa
30a. Lirellae much-branched; ascospores 15-26 pm long, 6-8 -locular . G. intricata
30b. Lirellae simple or little branched. 31
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31a. Lirellae completely lacking a thalline margin [cf. Opegrapha]; ascospores 30-40 pm long,
8-11 -locular . G. emersa
31b. Lirellae with a thalline margin. 32
32a. Ascospores 55-85 pm long, 15-20-locular . G. leucoparypha
32b. Ascospores < 50 pm long. 33
33a. Lirellae 2-6 mm long; ascospores 25-35 pm long, 8-11-locular . G. kakaduensis
33b. Lirellae < 3 mm long . 34
34a. Proper exciple completely carbonised; ascospores 25-35 pm long, 6-9 -locular .
. G. desquatnescens
34b. Proper exciple completely, or almost completely carbonised; ascospores 15-30 pm long,
6-8 -locular . G. librata
35a. Protocetraric acid present; proper exciple laterally carbonised; ascospores 25-32 pm long,
8-10-locular. G. supracola
35b. Stictic acid present . 36
36a. Proper exciple laterally carbonised; ascospores 24-33 pm long, 6 -locular.
. G. leptocarpa
36b. Proper exciple completely carbonised. 37
37a. Ascospores 28-40 pm long, 8-11 -locular . G. descissa
37b. Ascospores 66-84 pm long, 14-18 -locular . G. rustica
38a. Thallus saxicolous . 39
38b. Thallus corticolous .. 40
39a. Proper exciple laterally carbonised; ascospores 70-90 pm long; lichen compounds absent
. G. celata
39b. Proper exciple completely carbonised; ascospores 40-5 5 pm long; norstictic acid present
. G. saxicola
40a. Carbonised exciple concealed in thalline margin; norstictic or hirtiructic acid present ....
. 41
40b. Carbonised exciple visible; lichen compounds present or absent . 43
41a. Exciple completely carbonised; ascospores terminally muriform only . 42
41b. Exciple laterally carbonised; ascospores 120-140 pm long, fully muriform; G.atrocelata
42a. Ascospores 85-105 pm long; norstictic acid present. G. aquilonia
42b. Ascospores 102-130 pm long; hitifructic acid present . G. elixiana
43a. Lirellae short and simple, 1-2 mm long . 44
43b. Lirellae > 2 mm long, simple or branched . 46
44a. Proper exciple completely carbonised; ascospores 95-150 pm long . 45
44b. Proper exciple laterally carbonised; ascospores 90-100 pm long; norstictic acid present
. G. hiascens
45a. Norstictic and protocetraric acids present; ascospores 115-150 pm long .... G. lumbschii
45b. Lichen compounds absent; ascospores 95-120 pm long . G. lumbschii var. deftciens
46a. Lichen compounds present; proper exciple laterally carbonised. 47
46b. Lichen compounds absent; proper exciple laterally or completely carbonised. 51
47a. Norstictic acid present . 48
47b. Stictic acid present . 50
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Telopea 11(1): 2005
67
48a. Ascospores < 60 pm long. 49
48b. Ascospores 100-130 pm long . G. subserpentina
49a. Lirellae immersed, visible as a thin black line; ascospores 37-50 pm long . G. borealis
49b. Lirellae conspicuous, black; ascospores 25-35 pm long . G. gradlescens
50a. Ascospores 37-50 pm long . G. polyclades
50b. Ascospores 70-90 pm long . G. streblocarpa
51a. Proper exciple completely carbonised; ascospores 100-145 pm long, terminally muriform
only. G. vestitoides
51b. Proper exciple laterally carbonised . 52
52a. Ascospores > 35 pm long. 53
52b. Ascospores < 35 pm long. 54
53a. Ascospores 60-80 pm long. G. daintriensis
53b. Ascospores 35-45 pm long. G. subvelata
54a. Lirellae closed; ascospores 2-seriate, 19-23 pm long . G.tenuirima
54b. Li—rellae open; ascospores 1-seriate, 20-28 pm long. G dimidata
Hemithecium
la. Ascospores septate with lenticular locules, . 2
lb. Ascospores muriform . 3
2a. Lirellae sessile; ascospores 80-95 pm long, 15-24 -locular. H. aphanes
2b. Lirellae immersed; ascospores 28-32 pm long, 6-10 -locular. H. argopholis
3a. Lirellae with grooves; stictic acid present . 4
3b. Lirellae lacking grooves . 3
4a. Ascospores 50-75 pm long. H. chrysenteron
4b. Ascospores 80-100 pm long; . H. ddorocarpoides
5a. Ascospores 8 per ascus, 35-40 pm long; stictic acid present . H. radidcola
5b. Ascospores 1 per ascus; stictic acid absent . 6
6 a. Ascospores 155-225 pm long . H. hadrospora
6 b. Ascospores >125 pm long. 2
7a. Ascospores 80-100 pm long. H.incerta
7b. Ascospores 57-80 pm long. H. contorta
Leiorreuma
la. Lichen compounds absent; ascospores 20-33 pm long, 6 -locular . L. exaltum
lb. Lichen compounds present . 2
2a. Stictic or hypostictic acid present . 3
2b. Nornotatic acid present; ascospores 21-25 mm long, 6 -locular . L. nornotaticum
3a. Hypostictic acid present; ascospores 25-40 pm long, 7-8 -locular. L. hypomelaenum
3b. Stictic acid present; ascospores 25-37 pm long, 8-9 -locular . L. melanostalazans
68
Telopea 11(1): 2005
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Phaeographis
la. Ascospores muriform . 2
lb. Ascospores septate with lenticular locules. 7
2a. Ascospores 1 per ascus, 100-135 pm long; norstictic acid present. P. atromaculata
2b. Ascospores 8 per ascus; norstictic acid absent . 3
3a. Stictic acid present . 4
3b. Lichen compounds absent. 5
4a. Ascospores 36-53 pm long, 8-11 x 2-5 -locular; proper exciple carbonised .. P. wilsonii
4b. Ascospores 25-35 pm long, 6-8 x 2-3 -locular; proper exciple yellow-brown.
. P. montiscalvi
5a. Ascospores 15-18 pm long, 4x2 -locular. P. exilior
5b. Ascospores > 20 pm long.6
6 a. Ascospores 40-60 pm long, 10-14 x 2-3 -locular. P. litoralis
6 b. Ascospores 23-35 pm long, 6-8 x 2-3 -locular. P. caesioradians
7a. Thallus saxicolous; ascospores 4 -locular . 8
7b. Thallus corticolous; ascospores > 4-locular. 10
8 a. Lirellae open, disc visible; ascospores 12-15 pm long . P. Itypoglaucoides
8 b. Lirellae closed or only slightly open . 9
9a. Thallus smooth; thalline margins absent; ascospores 12-15 pm long . P. eludens
9b. Thallus tuberculate; thalline margins conspicuous; ascospores 10-12 pm long.
. P. tuberculifera
10a. Ascospores 4-locular. 11
10 b. Ascospores > 4-locular. 16
11a. Norstictic acid present . 12
lib. Norstictic acid absent.13
12a. Ascospores 15-15 pm long. P. subtigrina
12 b. Ascospores 14—23 pm long . P. brasiliensis
13a. Carbonised exciple present;. 14
13b. Carbonised exciple absent; ascospores 8-12 pm long . P. ceratoides
14a. Proper exciple completely carbonised; ascospores 15-22 pm long. P. elaeina
14b. Proper exciple laterally or apically carbonised . 15
15a. Proper exciple laterally carbonised; ascospores 14-24 pm long . P. subintricata
15b. Proper exciple apically carbonised; ascospores 17-20 pm long . P.lindigiana
16a. Ascospores 4-6 -locular.17
16b. Ascospores > 6-locular . 19
17a. Norstictic acid present; ascospores 16-20 pm long. P. intricans
17b. Norstictic acid absent. 18
18a. Lichen compounds absent; ascospores 16-22 pm long . P. subdividens
18b. Neotricone present; ascospores 12-23 pm long . P. neotricosa
19a. Norstictic acid present . 21
19b. Norstictic acid absent; lirellae carbonised. 20
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69
20a. Lichen compounds absent; lirellae apically carbonised; ascospores 27-47 pm long,
8-10 -locular . ft lobata
20b. Stictic acid present; thin carbonised exciple present; ascospores 20-37 pm long, 6-8 -locular
. ft dendroides
21 a. Lirellae carbonised; ascospores 30-55 pm long, 7-11-locular . P.mucronata
21b. Lirellae uncarbonised; ascospores 6-8-locular . 22
22a. Ascospores 23-36 pm long, . ft nardiensis
22b. Ascospores 15-31 pm long . ft platycarpa
Platygramme
la. Ascospores 8 per ascus.2
lb. Ascospores 1 per ascus.3
2a. Carbonised exciple visible; ascospores 20-40 pm long, 4-6 x 2-3 -locular.
.ft nrechavelatae
2 b. Carbonised exciple concealed; ascospores 13-18 pm long, 4x2 -locular.ft fuscescens
3a. Carbonised exciple visible; lichen compounds absent .4
3b. Carbonised exciple concealed; echinocarpic acid present; ascospores 162-200 pm long
.ft pudica
4a. Lirellae conspicuously open; ascospores.5
4b. Lirellae not conspicuously open; ascospores 135-180 pm long.ft impudica
5a. Ascospores 145-180 pm long . ft muelleri
5b. Ascospores 40—75(—100) pm long . ft australiensis
Sarcographa
la. Lichen compounds absent; ascospores 14-18 x 5-6 pm, 4-locular . S. subtricosa
lb. Stictic acid present. 2
2a. Ascospores 7-10 -locular, 25-37 pm long . S. ocidata
2b. Ascospores < 6 -locular. 3
3a. Ascospores 17-22 pm long, 4-locular . S. labyrintliica
3b. Ascospores 23-32 pm long, 6 -locular . S. verrucosa
Thecaria
la. Disc red; hymenium with red pigment [isohypocrellin]; ascospores 125-175 pm long,
muriform . T. montagnei
lb. Disc white pruinose; hymenium lacking red pigment; ascospores 75-100 pm long .
. T. quassiicola
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List of species in Australia
Acanthothecis Clem.
1. Acanthothecis gyridia (Stirt.) A.W. Archer, comb. nov.
Graphis gyridia Stirt., Trans. Proc. R. Soc. Vic. 17: 77 (1881)
Graphina gyridia (Stirt.) Zahlbr., Cat. Lick. Univ. 2: 412 (1923)
2. Acanthothecis cf. silicicola (Redinger) Staiger 8c Kalb, Mycotaxon 73: 112 (1999)
Graphis cf. silicicola Redinger, Ark. Bot. 27A (3): 56 (1935)
3. Acanthothecis subaggregans (Mull. Arg.) A.W. Archer, comb. nov.
Graphina subaggregans Mull. Arg. Bull. Herb. Boissier 1: 58 (1893)
Acanthothecis gracilis Staiger & Kalb, Mycotaxon 73: 99 (1999), syn. nov.
Carbacanthograpliis Staiger 8c Kalb
1 . Carbacanthograpliis marcescens (Fee) Staiger 8c Kalb, Biblioth. Lichenol. 85: 109
( 2002 )
Graphis marcescens Fee, Essai Crypt.: 38 (1825)
2. Cabacanthographis salazinica (A.W. Archer) A.W. Archer, comb. nov.
Graphina salazinica A.W. Archer, Mycotaxon 77: 176 (2001)
Diorygma Eschw.
1. Diorygma circumfusum (Stirt) Kalb, Staiger 8c Elix, Symb. Bot. Ups. 34(1): 145
(2004)
Graphis circutnfusa Stirt., Trans. & Proc. Roy. Soc. Victoria 17: 73 (1881)
2. Diorygma erythrellum (Mont. & Bosch) Kalb, Staiger & Elix, Symb. Bot. Ups. 34(1):
150 (2004)
Uslalia erythrella Mont. Ik Bosch, Plant, junghuhn., Fasc. IV: 478 (1855)
Graphina erythrella (Mont. & Bosch) Zahlbr., Cat. Licit. Univ. 2: 405 (1923)
Graphina incisa A.W. Archer, Mycotaxon 77: 169 (2001)
Graphina atramontana A.W. Archer, Mycotaxon 77: 161 (2001), syn. nov.
3. Diorygma junghuhnii (Mont. 8c Bosch) Kalb, Staiger 8c Elix, Symb. Bot. Ups. 34(1):
157(2004)
Ustalia junghuhnii Mont. 8c Bosch, Plant, junghuhn., Fasc. IV: 477 (1855)
Graphis mendax Nyl., Ann. Sci. Nat. Bot. ser. 4, 11: 244 (1859)
4. Diorygma hieroglyphicum (Pers.) Staiger 8c Kalb, Symb. Bot. Ups. 34( 1): 151 (2004)
Opegrapha hieroglyphica Pers., Ann. Wetterauischen Ges. Gesammthe Naturk. 2:16
(1811)
Graphis pallido-ochracea Kremp., Nuovo. Giorn. Bot. Ital. 7: 32 (1875)
5. Diorygma nothofagi (A.W. Archer) A.W. Archer, Australasian Lichenology 56:
10(2005)
Graphina nothofagi A.W. Archer, Mycotaxon 77:172 (2001)
6 . Diorygma pruinosum (Eschw.) Kalb, Staiger 8c Elix, Symb. Bot. Ups. 34(1): 166
(2004)
Leiogramma pruinosum Eschw., in Martius, Icon, select, cryptogam. Fasc. I: 12 (1828)
New combinations in Australian Graphidaceae
Telopea 11(1): 2005
71
Graphis platyleuca Nyl., Syr;. Lick. Nov. Cal.: 75 (1868)
7. Diorygma rufopruinosum (A.W. Archer) Kalb, Staiger & Elix, Symb. Bot. Ups. 34 (1):
169 (2004)
Graphina rufopruinosa A.W. Archer, Mycotaxon 77: 175 (2001)
Graphina boweniana A.W. Archer, Mycotaxon 77: 164 (2001)
8 . Diorygma wilsoniana (Mull. Arg.) A.W. Archer, Australasian Lichenology 56: 10
(2005)
Graphis wilsoniana Miill. Arg., Bull. Herb. Boissier 1: 57 (1893)
Dyplolabia A. Massal.
1. Dyplolabia afzelii (Ach.) A. Massal. Neagenea lichenum: 6 (1854)
Graphis afzelii Ach., Syn. Lich.: 85 (1814)
Fissurina Fee
1. Fissurina abdita (A.W.Archcr) A.W. Archer, comb. nov.
Graphina abdita A.W. Archer, Mycotaxon 77: 160 (2001)
2. Fissurina albonitens (Mull. Arg.) A.W. Archer, comb. nov.
Graphis albonitens Mull. Arg., Hedwigia 30: 53 (1891)
3. Fissurina dumastii Fee, Essai Crypt.: 1-59 (1825)
Fissurina glauca (Mull. Arg.) Staiger, Biblioth. Lichenol. 85: 159 (2002)
Graphis glauca Mull. Arg., Bull. Herb. Boissier 1:58 (1893)
4. Fissurina elaiocarpa (A.W. Archer) A.W. Archer, comb. nov.
Graphina elaiocarpa A.W. Archer, Mycotaxon 77: 167 (2001 )
Fissurina marginata Staiger, Biblioth. Lichenol. 85: 144 (2002), syn. nov.
5. Fissurina elixii (A.W. Archer) A.W. Archer, comb. nov.
Graphis elixii A.W. Archer, Australasian Lichenology 43: 16 (1998)
6 . Fissurina globulifica (Nyl.) Staiger, Biblioth. Lichenol 85: 137 (2002)
Graphis globulifica Nyl., Bull. Soc. Linn. Normandie, ser. 2, 2: 117 (1868)
7. Fissurina howeatia (A.W. Archer) A.W. Archer, comb. nov.
Graphis howeana A.W. Archer, Aus. Syst. Bot. 14: 259 (2001)
8. Fissurina insidiosa C. Knight & Mitt., Trans. Linn. Soc. London 23: 102 (1860)
Graphis insidiosa (C.Knight 8( Mitt.) J.D.Hooker, Handbook NZ Flora: 586 (1867)
Fissurina subcontexta (Nyl.) Nyl., Lich. Nov. Zel: 125 (1888)
Graphis subcontexta Nyl., Bull. Soc. Linn. Normandie, ser. 2, 2: 118 (1868)
Graphis robustior Miill. Arg., Nuovo Giorn Bot. Ital. 23: 398 (1891)
9. Fissurina nigririmis var. deficiens (A.W. Archer) A.W. Archer, comb. nov.
Graphis nigririmis (Nyl.) Miill.Arg. var. deficiens A.W.Archer, Aus. Syst. Bot. 14: 264
( 2001 )
10. Fissurina paradoxica (A.W. Archer) A.W. Archer, comb. nov.
Graphis paradoxica A.W. Archer, Mycotaxon 80: 367 (2001)
11. Fissurina psoromica (A.W. Archer) A.W. Archer, comb. nov.
Graphina psoromica A.W. Archer, Mycotaxon 77: 173 (2001)
72
Telopea 11(1): 2005
Archer
12. Fissurina streimannii (A.W. Archer) A.W. Archer, comb. nov.
Graphina streimannii A.W. Archer, Mycotaxon 88: 143 (2003)
Glyphis Ach.
1. Glyphis cicatricosa Ach., Syti. Lick: 107 (1814)
2. Glyphis montoensis (A.W. Archer) Staiger, Biblioth. Lichenol. 85: 173 (2002)
Graphina montoensis A.W. Archer, Mycotaxon 77: 172 (2001)
3. Glyphis scyphulifera (Ach.) Staiger, Biblioth. Lichenol. 85: 175 (2002)
Lecidea scyphulifera Ach., Syn. Licit.: 27 (1814)
Gyrostomum scyphulifereum (Ach.) Nyl. Bull. Soc. Linn. Normandie, ser. 2, 2: 78 (1867)
Graphis Adans.
1. Graphis albissima Miill. Arg., Bull. Herb. Boissier 3: 319 (1895)
2. Graphis anfractuosa (Eschw.) Eschw., in C.F.P. von Martius, FI. Bras. enum. pi 1 : 86
(1833)
Scaphis anfractuosa Eschw., Syst. Licit.: 25 (1824)
3. Graphis apertella A.W. Archer, Aus. Syst. Bot. 14: 258 (2001)
4. Graphis aquilonia (A.W. Archer) Staiger, Biblioth. Lichenol. 85: 209 (2002)
Graphina aquilonia A.W. Archer, Mycotaxon 77: 160 (2001)
5. Graphis atrocelata (A.W. Archer) A.W. Archer, comb. nov.
Graphina atrocelata A.W. Archer, Mycotaxon 77: 163 (2001)
6. Graphis borealis (A.W. Archer) A.W. Archer, comb. nov.
Graphina borealis A.W. Archer, Mycotaxon 77: 164 (2001)
7. Graphis catherinae A.W. Archer, Aus. Syst. Bot. 14: 259 (2001)
8. Graphis celata (A.W. Archer) A.W. Archer, comb. nov.
Graphina celata A.W. Archer, Mycotaxon 77: 166 (2001)
9. Graphis crassilabra Miill. Arg., Flora 65: 502 (1882)
10. Graphis daintriensis (A.W. Archer) A.W. Archer, comb. nov.
Graphina daintriensis A.W. Archer, Mycotaxon 77: 166 (2001)
11. Graphis dimidata Vain., Acta Soc. Fauna Flora Fenn. 7(2): 108 (1890)
12. Graphis descissa Miill. Arg., Bull. Herb. Boissier 3:318 (1895)
13. Graphis desquantescens (Fee) Zahlbr ., Denkschr. Akad. Wiss. Wien math.-naturwiss.
Kl. 83: 108 (1909)
Opegrapha desquantescens Fee, Bull. Soc. Bot. France 21: 24 (1874)
14. Graphis elegans (Smith) Ach., Syn. Licit: 85 (1814)
Opegrapha elegans Smith, in J.E. Smith & J. Sowerby English Botany: 16 (1807)
New combinations in Australian Graphidaceae
Telopea 11(1): 2005
73
15. Graphis elixiana A.W. Archer, ttotti. nov.
[The name Graphis elixii already exists; see Fissurina elixii]
Phaeographina elixii A.W. Archer, Biblioth. Lichenol. 78: 13-16 (2001)
16. Graphis emersa Miill. Arg., Hedwigia 32: 132 (1893)
17. Graphis endoxantha Nyl., Bull. Soc. Linn. Normandie, ser. 2, 2: 110 (1868)
18. Graphis epimelaena Mull. Arg., Bull. Herb. Boissier 3: 319 (1895)
19. Graphis gracilescens Vain., Ann. Acad. Sci. Fenn. ser. A, 15, 6: 203 (1920)
20. Graphis hiascens (Fee) A.W. Archer, comb. nov.
Opegrapha hiascens Fee, Suppl. Ess. Crypt. Ecorc.\ 25 (1837)
21. Graphis immersella Miill. Arg., Bull. Herb. Boissier 3: 319 (1895)
22. Graphis immersicans A.W.Archer, Aust. Syst. Bot. 14: 262 (2001)
23. Graphis inamoena Zahlbr., Ann. Crypt. Exot. 1: 126 (1928)
24. Graphis intricata Fee, Essai Crypt., 42 (1825)
Graphis centrifuga Riis., Arch. Soc. Zool. Bot. Fenn. “Vanamo 3:187 (1949)
25. Graphis kakaduensis A.W. Archer, Aus. Syst. Bot. 14: 264 (2001)
26. Graphis leptocarpa Fee, Essai Crypt.: 36 (1824)
27. Graphis leptoclada Miill.Arg., Flora 65: 335 (1882)
28. Graphis leucoparypha Kremp., Nuovo Giorn. bot. ital. 7: 35 (1875)
Graphis turgidula var. norstictica A.W. Archer, Aus. Syst. Bot. 14:267 (2001), syn. nov.
29. Graphis librata C. Knight, Trans. N.Z. Instit. 16: 404 (1884)
30. Graphis longula Kremp., Flora 59: 414 (1876)
31. Graphis lumbschii (A.W. Archer) A.W. Archer, comb. nov.
Graphina lumbschii A.W. Archer, Mycotaxon 77: 166 (2001)
31 a. Graphis lumbschii var. deficiens (A.W. Archer) A.W. Archer, comb. nov.
Graphina lumbschii var. deficiens A.W. Archer, Mycotaxon 77: 167 (2001)
32. Graphis pertricosa (Kremp.) A.W. Archer, comb. nov.
Enterographa pertricosa Kremp., Nuovo. Giorn. Bot. Ital. 7: 39 (1875)
33. Graphis polyclades Kremp., Verb. K.K. Zool.-Bot. Ges. Wien 30: 341 (1880)
34. Graphis propinqua Miill. Arg., Flora 65: 502 (1882)
35. Graphis rimulosa (Mont.) Trevis., Spighe e Paglie: 11 (1853)
Opegrapha rimulosa Mont., Ann. Sci. Nat. Bot., ser.2, 18: 271 (1842)
36. Graphis rustica Kremp., Nuovo Giorn. bot. ital. 7: 61 (1875)
Graphis turgidula Mull. Arg., /. Linn. Soc. Bot. London 30: 457(1895)
74
Telopea 11(1): 2005
Archer
37. Graphis saxicola (Mull. Arg.) A.W. Archer, comb. nov.
Graphina saxicola Mull. Arg., Flora 70: 401 (1887)
38. Graphis sayeri Mull. Arg., Flora 70: 401 (1887)
39. Graphis semiaperta Mull. Arg., Nuovo Giorn. bot. ital. 23: 397 (1891)
40. Graphis stenospora Miill. Arg. var. deficiens A.W. Archer, Mycotaxon 80: 370
( 2001 )
41. Graphis stenotera Vain., Ann. Acad. Sci. Fenn. ser. A, 15: 209 (1920)
42. Graphis stipitata A.W. Archer, Mycotaxon 80: 368 (2001)
43. Graphis streblocarpa (Bel.) Nyl., Flora 49: 133 (1866)
Opegrapha streblocarpa Bel., Voy. Indies. Or., Botanique 11, Cryptogamie: 134 (1834)
Graphis fissofurcata Leight., Trans. Linn. Soc. London, Bot. 27: 177 (1869)
Graphina streblocarpa (Bel.) Mull. Arg., Flora 65: 502 (1882)
44. Graphis streimannii A.W. Archer, Aus. Syst. Bot. 14: 265 (2001)
45. Graphis subregularis A.W. Archer, Aus. Syst. Bot. 14: 266 (2001)
46. Graphis subserpentina Nyl., Acta Soc. Sci. Fenn. 7: 465 (1863)
Graphina subtartarea Mull. Arg., Flora 70: 402 (1887)
Graphina palmicola Mull. Arg., Flora 70: 402 (1887)
Graphina subserpentina (Nyl.) Mull. Arg., Bull. Bot. Soc. Belgique 32: 152 (1893)
47. Graphis subtenella Miill. Arg., Flora 70: 400 (1887)
48. Graphis subvelata Stirt., Queensland Agric. J. 5: 488 (1899)
Graphina subvelata (Stirt.) Zahlbr., Cat. Licit. Univ. 2: 428 (1923)
49. Graphis supracola A.W. Archer, Aus. Syst. Bot. 14: 267 (2001)
50. Graphis tenella auct, non Ach., Syn. Lich.: 81 (1814)
51. Graphis tenuirima (Shirley) A.W. Archer, comb. nov.
Graphina tenuirima Shirley, Bot. Bull. Dept. Agric. Qld., Bot. Bull. V:34 (1892)
52. Graphis treubii Zahlbr., Ann. Cryptog exot. 1: 129 (1928)
53. Graphis vestitoides (Fink) Staiger, Biblioth. Lichenol. 85: 263 (2002)
Graphina vestitoides Fink, Mycologia 19: 218 (1927)
Graphina acharii auct.
54. Graphis vinosa Miill. Arg., Bull. Herb. Boissier 3: 318 (1895)
55. Graphis xanthospora Miill. Arg., Bull. Herb. Boisier 3: 320 (1895)
Hemithecium Trevis.
1. Hemithecium aphanes (Mont. & Bosch) M. Nakan. & Kashiw.,
Bull. Natn. Sci. Mus., Tokyo, Ser. B, 29(2): 88 (2003)
Graphis aphanes Mont. & Bosch, Plant. Junghuhn. 4, 474 (1855)
Graphis vennifera Miill. Arg., Flora 70, 401 (1887)
2. Hemithecium argopholis (C.Knight in Miill. Arg) A.W. Archer, comb. nov.
Graphis argopholis C.Knight in Miill. Arg., Flora 70: 401 (1887)
New combinations in Australian Graphidaceae
Telopea 11(1): 2005
75
3. Hemithecium chlorocarpoides (Nyl.) Staiger, Biblioth. Lichenol. 85: 283 (2002)
Graphis chlorocarpoides Nyl. Flora 49: 133 (1866)
Graphina repleta var. macrospora A.W. Archer, Telopea 8: 291 (1999), syn. nov.
4. Hemithecium clirysenteron (Mont.) Trevis, Spighe e Paglie 1: 13 (1853)
Phaeographina clirysenteron (Mont.) Miill. Arg., Hedwigea 30: 52 (1891)
Graphis clirysenteron Mont., Ann. Sci. Nat., Bot. 18(2): 268 (1842)
Graphis repleta Stil t., Trans. & Proc. Roy. Soc. Victoria 17: 73 (1881), syn. nov.
5. Hemithecium contorta (Miill. Arg.) A.W. Archer, comb. nov.
Graphina contorta Mull. Arg., Rev. Mycol. 9: 81 (1887)
6. Hemithecium hadrospora (A.W. Archer) A.W. Archer, comb. nov.
Phaeographina hadrospora A.W. Archer, Telopea 9: 337 (2001)
7. Hemithecium incerta (Redinger) A.W. Archer, comb. nov.
Graphina incerta Redinger, Ark. Bot. 26A( 1): 59 (1933)
8. Hemithecium radicicola (A.W. Archer) A.W. Archer, comb. nov.
Graphina radicicola A.W. Archer, Mycotaxon 77: 175 (2001)
Leiorreuma Eschw.
1. Leiorreuma exaltatum (Mont. & Bosch) Staiger, Biblioth. Lichenol. 85: 298 (2002)
Phaeographis exaltata (Mont. & Bosch) Miill. Arg., Flora 65: 381 (1882)
Lecanactis exaltata Mont.Sc Bosch, in Junghuhn, Plant, junghuhn., Fasc. IV: 475 (1855)
2. Leiorreuma hypomelaenum (Miill. Arg.) Staiger, Biblioth. Lichenol. 85: 300 (2002)
Paheographis hypotnelaena Mull. Arg., Flora 69: 313 (1886)
Phaeographis necopinata A.W. Archer & Elix, Mycotaxon 72: 92 (1999), syn. nov.
3. Leiorreuma melanostalazans (Leight.) A.W. Archer, comb. nov.
Phaeographis melanostalazans (Leight.) Miill. Arg., Flora 65: 336 (1882)
Platygrapha melanostalazans Leight., Trans. Linn. Soc. London 27: 180 (1869)
4. Leiorreuma nornotaticum (A.W. Archer) A.W. Archer, comb. nov.
Phaeographis nornotatica A.W. Archer 8< Elix, Mycotaxon 72: 93 (1999)
Phaeographis Miill. Arg.
1. Phaeographis atromaculata (A.W. Archer) A.W. Archer, comb. nov.
Phaeographina atromaculata A.W. Archer, Telopea 9: 331 (2001)
2. Phaeographis brasiliensis (A. Massal.) Kalb & Matthes-Leicht, Biblioth. Lichenol. 78:
148 (2001)
3. Phaeographis caesioradians (Leight.) A.W. Archer, comb. nov.
Phaeographina caesioradians (Leight.) Redinger, Ark. Bot. 26 A: 99 (1933)
Graphis caesioradians height., Trans. Linn. Soc. London 27: 176 (1869)
4. Phaeographis ceratoides (Vain.) Zahlbr. Cat. Licit. Univ. 2: 365 (1923)
Graphis ceratoides Vain., Ann. Acad. Sci. Fenn. ser. A. 15 (6): 227 (1920)
5. Phaeographis dendroides (Leight.) Miill. Arg., Flora 65: 336 (1882)
Platygrapha dendroides Leight., Trans. Linn. Soc. London 27: 179 (1869)
76
Telopea 11(1): 2005
Archer
6. Phaeographis elaeina (C. Knight) Mull. Arg., Bull. Herb. Boissier 3: 321 (1895)
Graphis elaeina C. Knight, Trans. Linn. Soc. London, Bot. 2: 41 (1882)
7. Phaeographis eludens (Stirt.) Shirley, Proc. Roy. Soc. Queensland 6: 197 (1889)
Graphis eludens Stirt., Trans. Proc. Roy. Soc. Victoria 17: 72 (1881)
8. Phaeographis exilior (Vain.) A.W. Archer, comb. nov.
Phaeographina exilior (Vain.) Zahlbr., Cat. Lich. Univ. 2: 438 (1923)
Graphis exilior Vain., Ann. Acad. Sci. Fenn., ser.A, 15, 6: 200 (1920)
9. Phaeographis hypoglaucoides, K.P. Singh 8c Awasthi, Bull. Bot. Survey India 21: 109
(1979)
10. Phaeographis intricans (Nyl.) Staiger, Biblioth. Lichenol. 85: 329 (2002)
Sarcographa intricans (Nyl.) Mull. Arg., Flora 70: 77 (1887)
Graphis intricans Nyl., Acta Soc. Sci. Fenn. 7: 473 (1863)
11. Phaeographis lindigiana Mull. Arg., Flora 65: 383 (1882)
Phaeographispseudomelana Miill. Arg. Bull. Herb. Boissier 3: 321 (1895), syn. nov.
12. Phaeographis litoralis (A.W. Archer) A.W. Archer, comb. nov.
Phaeographina litoralis A.W. Archer, Telopea 9: 339 (2001)
13. Phaeographia lolmta (Eschvv.) Miill. Arg., Flora 65: 383 (1882)
Lecanactis lobata Eschw., Syst. Lich.: 25 (1824)
14. Phaeographis montiscalvi (A.W. Archer) A.W. Archer, comb. nov.
Phaeographina montiscalvi A.W. Archer, Telopea 9: 341 (2001)
15. Phaeographis mucronata (Stirt.) Zahlbr., Cat. Lich. Univ. 2:382 (1923)
Graphis mucronata Stirt., Trans. Glasgow Field Naturalists 4: 95 (1876)
16. Phaeographis nardiensis A.W. Archer, Telopea 9; 674 (2001)
17. Phaeographis neotricosa Redinger, Ark. Bot. 27A(3): 93 (1935)
18. Phaeographis platycarpa Mull. Arg., Bot. Jahrb. Syst. 20: 284 (1894)
19. Phaeographis subdividens (Leight.) Mull. Arg., Flora 65: 383 (1882)
Graphis subdividens Leight., Trans. Linn. Soc. London 27: 177 (1869)
20. Phaeographis subintricata (C. Knight) Miill. Arg., Bull. Herb. Boissier 3: 320
(1895)
Graphis subintricata C. Knight, Trans. Linn. Soc. London, Bot: 2: 40 (1882)
21. Phaeographis subtigrina (Vain.) Zahlbr., Cat. Lich. Univ. 2: 287 (1923)
Graphis subtigrina Vain., Hedwigia 46: 177 (1907)
22. Phaeographis tuberculifera A.W. Archer, Telopea 9: 675 (2001)
23. Phaeographis wilsonii (A.W. Archer) A.W. Archer, comb. nov.
Phaeographina wilsonii A.W. Archer, Telopea 9: 343 (2001)
Platygramme Fee
1. Platygramme arechavaletae (Miill. Arg.) A.W. Archer, comb. nov.
Phaeographina arechavaletae Miill. Arg., Rev. Mycol. 10: 5 (1888)
Phaeographina banksiae Miill. Arg., Bull. Herb. Boissier 1: 59 (1893)
New combinations in Australian Graphidaceae
Telopea 11(1): 2005
77
2. Platygramme australiensis Staiger & Matthes-Leicht, Biblioth. Lichenol. 85: 355
( 2002 )
[ Phaeographitta caesiopruinosa (Fee) Mull. Arg., auct. australis].
3. Platygramme fuscescens (A.W. Archer) A.W. Archer, comb. tiov.
Phaeographina fuscescens A.W. Archer, Telopea 9: 337 (2001)
4. Platygramme impudica (A.W. Archer) A.W. Archer, comb. nov.
Phaeographina impudica A.W. Archer, Telopea 9: 339 (2001)
5. Platygramme muelleri (A.W. Archer) Staiger, Biblioth. Lichenol. 85: 364 (2002)
Phaeographina muelleri A.W. Archer, Telopea 8: 473 (2000)
Phaeographina caesiopruinosa (Fee) Mull. Arg. var. monospora Mull. Arg., Bull. Herb.
Boissier3 : 322 (1895)
6. Platygramme pudica (Mont. & Bosch) M. Nakan. & Kashiw., Bull. Natn. Sci.
Mus., Tokyo , ser.B, 29(2): 89 (2003)
Graphis pudica Mont. 8c Bosch, Plant, junghuhn., Fasc.4: 474 (1855)
Phaeographina echinocarpica A.W. Archer 8c Elix, Mycotaxon 72: 91 (1999), syn. nov.
Platythecium Staiger
1. Platythecium pertenellum (Stirt.) A.W. Archer, comb. nov.
Graphis pertenella Stirt. , Trans. & Proc. Roy. Soc. Victoria 17: 72 (1881)
Graphina brachyspora Miill. Arg., Flora 66: 79 (1883)
Graphis laevigata Miill. Arg., Nuovo Giorn. Bot. Ital. 23:398 (1891)
Sarcographa Fee
1 . Sarcographa labyrinthica (Ach.) Mull. Arg., Mem. Soc. Phys. Geneve 29: 62 (1887)
Glyphis labyrinthica Ach. Syn. Lich.: 107 (1814)
2. Sarcographa oculata Miill. Arg., Bull. Herb. Boissier 3:323 (1895)
3. Sarcographa subtricosa (Leight.) Miill. Arg., Flora 70: 78 (1887)
Glyphis subtricosa Leight., Trans. Linn. Soc. London (Botany) 27: 181 (1869)
4. Sarcographa verrucosa (Mont. 8< Bosch) Zahlbr., Cat. Lich. Univ. 2: 467 (1923)
Glyphis verrucosa Mont. 8c Bosch, in Miquel, Plant, junghuhn. Fasc. IV: 489 (1855)
Sarcographina Miill. Arg.
1. Sarcographina cyclospora Miill. Arg., Flora 70: 425 (1887)
Glyphis cyclospora (Mull. Arg.) Shirley, Proc. Roy. Soc. Qld. 6: 215 (1889)
Thalloloma Trevis.
1. Thalloloma atronitens (A.W. Archer) A.W. Archer, comb. nov.
Graphina atronitens A.W. Archer, Mycotaxon 77: 163 (2001)
Tltecaria Fee
1. Tltecaria montagnei (Bosch) Staiger, Biblioth. Lichenol. 85: 446 (2002)
Graphis montagnei Bosch, in Miquel, Plant, junghuhn., Fasc. IV: 472 (1855)
2. Thecaria quassiicola Fee, Essai Crypt: 97 (1824) as ‘quassiaecola’,
Phaeographina quassiicola (Fee) Miill. Arg., Mem. Soc. Phys. Geneve 29: 47 (1887)
78
Telopea 11(1): 2005
Archer
References
Archer AW (2004) Additional synonymy in the Australian Graphidaceae. Australasian Lichenology
55: 16-27.
Archer AW (2005) Australian species in the genus Diorygma (Graphidaceae). Australasian
Lichenology 56: 10-11.
Kalb K (2001) New or otherwise interesting lichens.l. Bibliotheca Lichenologica 78: 141-167.
Kalb II, Staiger B, & Elix JA (2004) A monograph of the lichen genus Diorygma- a first attempt.
Symb. Bot. Ups. 34(1): 133-181
Nakanishi M (1977) Notes on Japanese species of Phaeographina. Hikobia 8: 91-100.
Nakanishi M, Kashiwadani H & Moon KH (2003) Taxonomical notes on Japanese
Graphidaceae (Ascomycotina), including some new combinations. Bull. Natn. Sci. Mus.,
Tokyo, ser.B, 29: 83-90.
Staiger B & Kalb K (1999) Acanthothecis and other graphidioid lichens with warty periphysoids
or paraphysis tips. Mycotaxon 73: 69-134.
Staiger B (2002) Die Flechtenfamilie Graphidaceae. Bibliotheca Lichenologica 85: 1-526.
Manuscript received 27th January 2005, accepted 30th June 2005
Telopea 11(1): 79-86
A new species of Pomaderris (Rhamnaceae)
from the Central Tablelands of
New South Wales
J.C. Millott 1 and K.L. McDougall 2
' Sydney Water Corporation, ‘Darien’ Mile End Road, Rouse Hill, NSW 2155 Australia;
2 Department of Environment and Conservation, PO Box 2115, Queanbeyan, NSW 2620 Australia
Abstract
Pomaderris walshii, a shrub of the Central Tablelands of New South Wales, is described and
illustrated. The species is apparently restricted to the upper Kangaroo River, south-east of
Robertson. It is threatened because of its small population size and possible future changes in land
use, and fire and flood frequencies. As a result of morphometric analyses on P. walshii and related
taxa, P. argyrophylla subsp. graniticola is raised to specific rank (as P. graniticola).
Introduction
Pomaderris comprises 70 species throughout Australia and New Zealand. Of the 65
species that occur in Australia, 45 of these occur within New South Wales (Harden
2000). Much of the genus has been reviewed over the last two decades resulting in a
significant increase in the number of known species, particularly in south eastern
Australia (Walsh 1988a; Walsh 1988b; Walsh 1990; Walsh 1992; Walsh & Coates 1997).
Pomaderris parrisiae N.G. Walsh, P. nitidula (Benth.) N.A. Wakef. and P. argyrophylla
N.A. Wakef. form a closely related group within Pomaderris (Walsh and Coates 1997)
and can be difficult to distinguish as they are morphologically very similar.
Collectively, they are found along the east coast of Australia from south-east NSW to
Queensland and are usually found in small, disjunct and isolated populations both
within and between species (Pig. 1).
A small population of a Pomaderris species was recently discovered near Carrington
Falls in the Central Tablelands of NSW and was found to have close affinities to the
above group of taxa. Surveys of the nearby area revealed that its distribution appears
to be limited to two small populations in the riparian zone in the upper Kangaroo
River catchment (Millott 2003). The smaller population, located in Budderoo National
Park, consists of 13 individual plants. The larger population is located on private land
approximately 2.5 km upstream, and consists of approximately 30 plants.
© 2005 Royal Botanic Gardens and Domain Trust
ISSN0312-9764
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Telopea 11(1): 2005
Millott and McDougall
A morphometric analysis was undertaken to compare plants from the populations in
the upper Kangaroo River with P. parrisiae, P. nitidula, P. argyrophylla subsp.
argyrophylla and P. argyrophylla subsp. graniticola to assess its level of differentiation
and taxonomic status.
Methods
A total of 76 dried collections were used in the morphometric analysis: 17 specimens
of the Pomaderris from Carrington Falls were measured and compared against 13
specimens of P. argyrophylla subsp. argyrophylla, 10 of P. argyrophylla subsp.
graniticola, 29 of P. parrisiae and 7 of P nitidula. The sample sizes reflect availability of
material.
The nine morphological characters measured were selected from a larger set used in
preliminary analyses (Millott 2003). These nine characters were found to fully describe
the variation amongst the selected taxa. Five of these characters were foliar and the
remaining four were floral (Table 1).
Each character was measured on either 10 leaves or 10 flowers per specimen and the
mean used in the statistical analysis. The two exceptions to this were leaf hair length
and lateral vein overtopping. These characters were measured once per specimen
because they were taken in a single visual inspection of the leaf. Only leaves that were
more than two nodes removed from the growth apex were chosen for measurement to
avoid taking measurements on juvenile leaves; this method appeared to be the most
reliable for choosing mature leaves. The leaf base angle was taken as the angle at the
base of the leaf between the two leaf margins at points approximately 5 mm along the
margins from the base. The inflorescence diameter was taken as the widest point across
the inflorescence which was at or near the base of the inflorescence for these species.
The morphometric data were analysed using the CLUSTER program of Primer for
Windows 5.2.8. A matrix of Bray-Curtis similarity coefficients from log (x+1)
transformed data was constructed to enable CLUSTER. The significance of differences
in characters between species groupings was determined by multiple ANOVA.
Results and Discussion
In the CLUSTER analysis, all but one collection was grouped with the other collections
of the taxon it had previously been determined as (Fig. 2). The aberrant collection
(ag5) had the highest values for five of the eight quantitative measures within that
taxon — the values were 1.8 to 2.5 standard deviations above the mean. As a
consequence, collection ag5 was omitted from subsequent analyses. Further collections
from the ag5 population would be useful to clarify its identity and status.
Although five clusters appear to be naturally separable, there was less similarity within
P. argyrophylla subsp. graniticola collections than between the other taxa groups.
Despite this, subsp. graniticola as an entity is distinct from the other taxa and a new
status is proposed here for it. The plants from Carrington Falls appear to be most
closely allied to P. nitidula. Pomaderris parrisiae is most closely allied to P. argyrophylla
subsp. argyrophylla.
A new species of Pomaderris (Rhamnaceae)
Telopea 11(1): 2005
81
O Pomaderris argyrophyll;
A. Pomaderris argyrophyll;
■fa Pomaderris nitidula
■ Pomaderris parrisiae
Pomaderris sp. nov.
subsp.
subsp.
argyrophylla
graniticola
Fig. 1. Distribution of the Pomaderris species that are the subject of this paper: • =
P. argyrophylla subsp. argyrophylla ; A = P. argyrophylla subsp. graniticola ; * = P. sp. nov.; ☆ = P.
nitidula-, ■ = P. parrisiae.
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Millott and McDougall
There were significant differences between taxa for each of the characters measured
(Table 1). The plants from Carrington Falls are separable from P. argyrophylla subsp.
argyrophylla by their larger leaf base angle, shorter leaves, and leaf vein indumentum,
from P. argyrophylla subsp. graniticola by their broader inflorescences, longer and
broader leaves, and presence of leaf vein indumentum, from P. nitidula by their shorter
anthers, narrower hypanthia, larger leaf base angle, and shorter sepals, and from
P. parrisiae by all characters except hypanthium diameter.
Because of the differences between the Carrington Falls Potnaderris and the four
related taxa, we believe it warrants recognition at the species level.
Table 1. 99% confidence intervals for each quantitative character measured, and
presence/absence of overtopping of abaxial vein indumentum.
All measurements used to determine character means were used to calculate the confidence intervals.
Within each character, identical superscript letters indicate that the means are not significantly different as
determined by single classification ANOVA (P < 0.05).
Character
P. argyrophylla
subsp.
argyrophylla
P. argyrophylla
subsp.
graniticola
P. nitidula
P. parrisiae
P. walshii
Anther
length (mm)
0.9 - 1,1 a
0.8- 1.0 a
1.2 - 1 ,5 b
1.3-1.4 b
0.9- 1.0 a
Hypanthium
diameter (mm)
0.9- 1.2 a
1.0- 1.1®
1.2-1,4 b
1.2 - 1.3 b
1.0- 1.1 a
Inflorescence
diameter (mm)
43 - 58 ac
19 — 41 b
26 - 49 ab
49 - 57 c
45 - 59 ac
Leaf base angle ( c )
45 - 61 a
74 - 121 b
56 - 74 a
63 - 68 c
71 - 84 b
Leaf simple
hair length
0.1 - 0.3 a
0.1 -1.1*
0.2 - 0.5 a
1.1 - 1.3 b
0.8- 1.0 a
Leaf length (mm)
54 - 77 a
19 - 36 b
38 - 52 c
61 - 66 a
43 - 52 c
Leaf width (mm)
16 - 21 a
9- 14 b
14- 19 a
21 - 23 c
15 - 19 a
Sepal length (mm)
1.7-1.9®
1.9 - 2.0 b
2.3- 2.6 C
2.5 - 2.6 C
1.8 - 2.0 ab
Overtopping
absent
absent
present
absent
present
of leaf lateral vein
hair bylacuna hair
Potnaderris walshii J.C. Millott & K.L. McDougall, sp. tiov.
Potnaderris nitidula (Benth.) N.A. Wakef. proxime affinis sed statura maiore, floribus
minoribus, lamina basi obtusiore et distributione geographica magis ad meridiem
differt.
Holotype: New South Wales: Central Tablelands: Budderoo National Park, Douglas
Creek, c. 0.7 km upstream of Carrington Falls, S of disused quarry, /. Millott , 1 Nov
2003 (NSW656890).
Shrub or small tree to 3 m high. Young stems and petioles with appressed, silvery to
rusty simple hairs and medium to dense, white stellate hairs. Leaves narrowly ovate;
(36—)43—52(—60) mm long; 14—19(—22) mm wide; base cuneate to obtuse; margins
A new species of Pomaderris (Rhamnaceae)
Telopea 11(1): 2005
83
ag7
Fig. 2. Cluster analysis of 79 specimens of five apparent Pomaderris taxa. The herbarium determination is indicated by the following labels: aa = P. argyrophylla
subsp. argyrophylla; ag = P. argyrophylla subsp. graniticola; cf = P. sp. nov.; n = P. nitidula; p = P. parrisiae.
84
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Millott and McDougall
entire, plane or slightly recurved; apex acute to acuminate; adaxial surface green and
glabrous; abaxial surface moderately hairy with loosely appressed, white-silvery simple
hairs and dense white stellate hairs; lateral veins not or slightly impressed above, clearly
visible below and covered with an indumentum of moderately dense, appressed rusty
simple hairs and sparsely to moderately dense white stellate hairs; abaxial lateral vein
indumentum overtopped by, or level with, lacuna indumentum; petiole 4-10 mm
long. Stipules narrowly triangular, apex acute, 2-6 mm long, soon deciduous.
Inflorescence of 20- c. 100 flowers, pyramidal to hemispherical, terminal, 4-6.5(-7.5)
cm long and wide at base; bracts deciduous; pedicels 1.5-4.3 mm long. Flowers cream-
coloured to yellow; externally pubescent to villous with loosely appressed, silvery
simple hairs and dense stellate hairs (sepals less densely indumented than
hypanthium); hypanthium (0.8-) 1-1.4mm in diameter, 0.8-1.2 mm long; sepals 1.8-
2.0 mm long; petals present, 1.7-1.9 mm long, spreading, spathulate; stamens 2-2.5 mm
long; anthers (0.7)-0.8-1.2 mm long; ovary inferior, summit simple-pubescent; style
glabrous, 1.6 - 1.9 mm long, branched in lower or middle third. Fig. 3.
Etymology: Named in recognition of botanist Neville Walsh of the National
Herbarium of Victoria for his work on the revision of this genus.
Distribution: Currently only known from the upper Kangaroo River and its tributaries
(above Carrington Falls) on the Central Tablelands of New South Wales.
Habitat and ecology: Populations have been recorded in riparian shrubland
dominated by Callicoma serratifolia, Ceratopetalum apetalum and Grevillea rivularis,
and open grassy forest (partly cleared for grazing) dominated by Eucalyptus fastigata.
An unnamed Pomaderris species was recorded close to Carrington Falls on the
Kangaroo River by Jordan (1989) in Eucalyptus piperita / E. sieberi forest but this has
not been relocated in recent times. The upper Kangaroo River is underlain by
Hawkesbury Sandstone and the soils in populations are sandy alluvium. Annual
rainfall is about 1800 mm and populations range in elevation from about 550 - 600 m
above sea level. The Budderoo population was last burnt in 1983, however, the
response of the species to fire is unknown. Plants possibly resprout following flood
damage. Old, dead prostrate stems were observed on small plants closest to the river, a
possible consequence of the last major flood event in 1999.
Conservation status: Two populations of this species have been recorded over a lineal
range of about 3 km. One population of approximately 13 plants is reserved in
Budderoo National Park but most of these plants are apparently young. The remaining
plants are on freehold land. Populations may be threatened by changes in fire or flood
frequencies, or by future use of private land. A conservation code of 2ECi is suggested
for this species.
Other specimens examined: New South Wales: Central Tablelands: Douglas Creek
(a tributary of Kangaroo River) above Carrington Falls, SE of Robertson, K. McDougall 567, June
1998 (MEL 2117356).
New status for Pomaderris argyrophylla subsp. graniticola
As a result of our analysis we elevate P. argyrophylla subsp. graniticola to specific rank
because it is clearly morphometrically distinct from the other four taxa analysed.
85
A new species of Pomaderris (Rhamnaceae)
Telopea 11(1): 2005
Fig. 3. Pomaderris walshii. a, habit; b, flower; c, petal; d, stamen, showing petal attachment;
e, abaxial leaf indumentum; f, leaf venation; g, cross section of leaf. Scale bar: a = 5 cm; b = 0.4 cm;
c = 0.25 cm; d = 0.25 cm; e = 0.5 cm; f = 3 cm; g = 0.25 cm.
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Millott and McDougall
Pomaderrisgraniticola (N.A. Wakef.) K.L. McDougall & J.C. Millott, comb. & slat. nov.
Pomaderris argyrophylla N.A. Wakef. subsp. graniticola N.G. Walsh & F. Coates,
Muelleria 10: 46 (1997).
Type: Queensland, Girraween National Park, c. 50 m W of Dr Roberts Waterhole,
towards its southern reach, N.G. Walsh 3883, 15.ix. 1994 (holotype MEL; isotypes BRI,
CANB, NSW, UNE).
Notes: The apparent preference of this taxon for growing on granitic substratum is
retained in the new name. Notes on the distinctions between P. graniticola and
P. argyrophylla are given in Walsh and Coates (1997), who indicate that in the area cast
of Stanthorpe in Queensland, the two taxa may be difficult to distinguish. The
significant diagnostic characters identified above — inflorescence diameter (19-41 mm
in P. graniticola , 43-58 mm in P. argyrophylla), leaf base angle (74-121° in
P. graniticola, 45-61° in P argyrophylla), leaf length (19-36 mm in P. graniticola ,
54-77 mm in P. argyrophylla), leaf width (9-14 mm in P. graniticola, 16-21 mm in
P. argyrophylla), and sepal length (1.9-2.0 mm in P. graniticola, 1 . 7-1.9 mm in
P. argyrophylla) — will hopefully assist in identification.
Acknowledgments
The authors wish to thank Neville Walsh for much useful advice on Pomaderris
taxonomy. Sam Demuth assisted in both locating and helping to protect the
populations of this particularly vulnerable species. Belinda Pellow and Louisa Murray
gave their valuable assistance in curation of specimens at WOLL and NSW,
respectively. Many thanks also to Peter Wilson for the latin diagnosis and to
Catherine Wardrop for the illustration.
References
Harden GJ (2000) Pomaderris. Pp. 590-604 in GJ Harden (ed.), Flora of New South Wales. Volume
1 (Revised edition). (University of New South Wales Press: Kensington)
Jordan P (1989) Grevillea rivularis. Unpublished report. (NSW National Parks and Wildlife
Service: Hurstville)
Millott )C (2003) The taxonomy and ecology of a Pomaderris population near Carrington Falls,
NSW, (Honours Thesis: University of Wollongong)
Walsh NG (1988) Two new species of Pomaderris Labill. (Rhamnaceae) from south-eastern New
South Wales. Muelleria 6(6): 429-435.
Walsh NG (1988) Two new species of Pomaderris Labill. (Rhamnaceae) from south-eastern
Australia. Muelleria 7(1): 81-87.
Walsh NG (1990) Two new species of Pomaderris Labill. (Rhamnaceae) from New South Wales.
Muelleria 7(2): 207-212.
Walsh NG 8c Coates F (1997) New taxa, new combinations and an infrageneric classification in
Pomaderris (Rhamnaceae). Muelleria 10: 27-56.
Manuscript received 27 January 2005, accepted 11 July 2005
Telopea 11(1): 87-89
A colligate Spirogyra (Zygnemataceae,
Zygnematophyceae) in Australia
Stephen Skinner and Timothy J. Entwisle
Botanic Gardens Trust Sydney, Mrs Macquaries Road, Sydney NSW 2000, Australia
email: stephen.skinner@rbgsyd.nsw.gov.au
Abstract
A species of Spirogyra, Spirogyra yuin, with external collar-like sleeves holding cells in filaments
is described for the first time in Australian freshwater systems. Similar in form and dimensions
to S. silesiaca, it is distinct in possessing an irregularly honeycombed mesospore wall and golden
mesospore.
Introduction
No reports of colligate Spirogyra have been published for the southern hemisphere. In
describing Spirogyra colligata, Hodgetts noted a ‘most curious feature : the H shaped
piece of membrane’ between each touching pair of cells. This connecting-clamp is a
thin cylindrical piece of cell-wall...’ Hodgetts (1920). Hodgetts saw this taxon as
unique in the genus. Spirogyra colligata with vegetative cells 29-40 pm diam. and
lenticular zygospores with verrucose, golden brown mesospore walls is reported from
Europe and North America (Kadlubowska 1984). Kadtubowska (1969) investigated
further the connecting clamp, or collar, and amended Hodgetts description to point
out that the collar does not have a septum as described by Hodgetts but rather a
furrow or band, and was completely detachable, lwo further colligate taxa have been
described for Spirogyra: Spirogyra silesiaca Kadi, has vegetative cells 43—52 pm diam.,
and lenticular zygospores with verrucose, brown mesospores, described from Poland
(Kadlubowska 1967). Spirogyra subcolligata Bi has vegetative cells 37-41 pm diam.,
and smooth-walled, lenticular to ovoid zygospores, the mesospore wall quite thick,
and sulcate, described from Huaiyang, China (Bi 1979).
Spirogyra yuin Skinner & Entwisle sp. ttov.
Type: NSW: South Coast: Dry River, below bridge at Quaama, Skinner 0522 ,2 Jan 2002
(holo NSW).
Spirogyrae silesiacae similis sed mesosporis bilaminatis, rugosis externe, et inaequaliter
faveolatis atque aureis interne.
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Fig. 1. a-c. Spirogym yuin, Skinner 0522 (NSW), a, vegetative cell, with and without collar;
b, receptor gametangium and sigmoid connecting tube; c, zygospore (part) with sculpturing.
Scales 20 pm.
A new species of Spirogyra
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Vegetative cells cylindrical, L/D 4.5-12, 45—50(—55) pm diam., end-wall colligate,
chloroplasts ribbon-like, 4-5, straight or weakly spiralled; pyrenoids numerous, with
wide starch haloes. Conjugation scalariform, occasionally terminal; donor cell as
vegetative cells, receptor cell inflated ovoid in the middle of the cell, 79-95 pm across
the inflation; conjugation tubes from both sides, cylindrical to vermiform, funnel
ended with mucilage collar at junction, 23-27pm diam. Zygospore lenticular, ovoid in
side view, 68-82 pm diam., c. 50 pm thick, exospore smooth, clear, thin, mesospore
two-layered, outer pearly, finely furrowed, inner wrinkled, honeycombed, golden
yellow. Fig. 1 a-c.
Etymology; named in recognition of the Yuin people, the traditional owners of the
area in which the collection was made and the river rises.
Distribution: New South Wales, in fluctuating river with cobblestone bed, endemic.
Specimen examined: New SouthWales: South Coast: Dry R., below bridge at Quaama, Skinner
0522 ,2 Jan 2002 (NSW).
While the specimens found at Quaama were similar to the published species in general
form, and are closest to S. silesiaca in dimensions, the furrowed, irregularly
honeycombed mesospore, golden in colour, is distinctive.
Acknowledgments
We thank Ms Alice Cheung for Chinese translations. Representatives of New South
Wales south coast aboriginal peoples agreed to the use of the Yuin name.
References
Bi Liejiu (1979) New Zygnemataceous algae from Henan Province. Oceanologica Limnologica
Sinica 10: 354-361.
Hodgetts W) (1920) A new species of Spirogyra. Annals of Botany 34: 519-524, PI. 22.
Kadtubowska JZ (1967) Spirogyra silesiaca sp. n. Fragmenta Floristica Geobotanica 13: 163-164.
Kadlubowska JZ (1969) Structure of cell-wall o (Spirogyra colligata I lodgetts (1920) and changes
of diagnosis of this species — Budowa sciany komorkowej Spirogyra colligata Hodgetts (1920)
oraz zmiana diagnozy tego gatunku. Fragmenta Floristica Geobotanica 15: 255-257.
Kadlubowska JZ (1984) Chlorophyta VIII Conjugatophyceae I: Zygnentales Siifiwasserflora von
Mitteleuropea 16 (Gustav Fischer Verlag: Jena)
Manuscript received 16 February 2005, accepted 20 April 2005
Telopea 11(1): 91-93
A new Zygnemopsis species (Zygnemataceae,
Zygnematophyceae), with mature zygospores,
from Australia
Stephen Skinner, Hannah McPherson and Gillian M. Towler
Botanic Gardens Trust Sydney, Mrs Macquaries Road, Sydney NSW 2000, Australia
email: Stephen.skinner@rbgsyd.nsw.gov.au
Abstract
A species of Zygnemopsis, Zygnemopsis faveoscrobictilata, with a distinctive cup-like concave horn
in the exospore at each corner of the zygospore is reported from the Dumaresq River. It is similar
in other respects to Z. quadrata and Z. areolata from China in vegetative dimensions and
mesospore sculpturing, but there is little or no separation between the exospore and mesospore
in the Australian material.
Introduction
As part of a recent survey of macroalgae of the Dumaresq, Macintyre, Severn and
Gwydir Rivers in northern New South Wales adjacent to the Queensland border, the
authors made numerous collections of various Zygnemataceae. Among these we found
a Zygnemopsis species as the dominant macroalga in riffle banks in the Dumaresq at
the point where the low-level bridge marks the border between the states, two
kilometres north-east of Mingoola. Here it formed numerous yellowish to apple green
rafts, especially in the pools between the larger cobblestones, and among the Azolla
pinnata and Ludwigia peploides ssp. montevidensis plants in the same shallow, warm-
water pools. It was notable, not only as the dominant species, fertile and so
distinguishable from Zygnema, but also in that many of the zygospores in most
conjugation ladders were mature. There have been eleven collections of Zygnemopsis
lodged at NSW from Western Australia, Northern Territory, New South Wales and
Queensland but these have rarely contained sufficient mature zygospores to
distinguish them with confidence to more than genus. No descriptions of taxa from
this genus have been published from Australian material (Kadfubowska 1984).
Zygnemopsis faveoscrobiculata Skinner, H. McPherson & Towler sp. tiov.
Type: NSW: North Western Slopes: Dumaresq R., at low level bridge on NSW-Qld
border, north of Mingoola, Skinner 0684, McPherson & Towler, 7 Oct 2004 (holo NSW
910473).
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Skinner, McPherson and Towler
Fig. 1 a-d. Zygnetnopsis faveoscrobiculata, Skinner 0684, McPherson & Towler (NSW),
a-c, various stages of maturity of zygospores, top and side views; d, mature zygospore and
gametangia. Scales = 20 pm.
A new species of Zygnemopsis
Telopea 11(1): 2005
93
Zygnemopsis areolatae similis sed membrana exosporae ad mesosporae adiacenta
cornuibusque concavatis, atque mesospora aspectu Iaterali globosa.
Vegetative cells narrow cylindrical, 11-14 pm diam, L/D 1.2-3, endwall plane;
chloroplasts stellate, (1—)2(—3), each with central pyrenoid; basal attachment of
filament not seen. Conjugation scalariform, zygospore filling conjugation canal and
reaching outer walls of both gametangia; gametangia pectin-filled post-zygote, not
markedly laminate. Zygospore rectangular ovoid in face view, subglobose in side view,
34-42 pm long, 25-27 pm diam., 20-25(—27) pm thick, with pectinate halo (most
visible in side view); exospore wall clear, close-fitting, wrinkled with four concave
corners, mesospore wall thick, faveolate-scrobiculate, golden brown, scrobiculae 2-2.8 pm
diam. Fig. 1 a-d.
Etymology: the name derives from the honeycomb patterns of dimples on the
mesospore.
Distribution: New South Wales-Queensland border, in river riffle banks.
Other specimen examined: New South Wales: North Western Slopes: Bakers Creek, near Bundara,
Skinner 0790, McPherson & Towler, 13 Oct 2004 (NSW).
Notes: The four cup-like concave horns of the exospore clearly distinguish this taxon
from similar species including Zygnemopsisstephaniae 1 ranseau ( I ranseau et al. 1934),
Z. quadrata )ao (Jao 1935), and Z. areolata Zhu (Zhu 1980). Zygnemopsis stephaniae
has a more quadrate zygospore, without distinctive corners on the exospoie.
Zygnemopsis quadrata has an exospore which is widely separate from the mesospoie,
and which has convex corner processes (or horns) and the mesospore has an equator ia
ridge. Zygnemopsis areolata has a wide separation between mesospore and exospoie,
and is more narrow, and lozenge-like in side view. The significance and developmenta
origin of these remarkable features are unknown.
Acknowledgments
The authors wish to acknowledge the support of the Hermon Slade Foundation foi
sponsoring the field work, the Botanic Gardens Trust, and Ms Alice C teung or
Chinese translations.
References
Jao Chin-Chih (1935) Studies on the freshwater algae of China, I. Zygnemataceae from Szechwan.
Sinensia 6:551-564.
Kadlubowska JZ (1984) Chlorophyta VIII Conjugatophyceae I: Zygnemales. Sufiwasserflora von
Mitteleuropea 16 (Gustav Fischer Verlag: Jena)
Transeau EN, Tiffany LH, Taft CE & hi LC (1934) New species of Zygnemataceae. Irons. Am.
Microscopical Soc. 53: 208-230.
Zhu Wan-Jia (1980) Some new species of the Zygnemataceae from Guangdong (Kwangtung). Acta
Phytotaxonomica Sinica 18: 106-109.
Manuscript received 16 February 2005, accepted 20 April 2005
Telopea 11(1): 95-98
Justice for Justice Barron Field
Helen Hewson
Centre for Plant Biodiversity Research, Plant Industry, CSIRO,
Canberra ACT 2601
Abstract
Botanical tributes to Mr Justice Barron Field are reviewed, a correction in Senna is pointed out,
and the new combination Senna barronfieldii (Colla) Hewson is made.
Introduction
It is overdue that we revisit the botanical tributes given to Mr Justice Barton Field
(1786-1846), judge in the Supreme Court of New South Wales (1816— 1824), to review
them and to make a correction. Two genera and one species were named in his honour.
Mr Justice Barron Field took a wide interest in the Colony during his posting. I bis
included observing the plants and animals, composing poetry about them, collecting
and drawing them, communicating collections to scientists in Britain and Eutope,
being a founding member of the Philosophical Society of Australasia and being the
inaugural President of the Agricultural Society of New South Wales. Much evidence of
this is revealed in his book, Geographical Memoirs on New South Wales (1825) (see also,
Desmond (1977) and Currey (1966)). While he seems to have been a controversy
figure at judicial, political, sociological and literary levels, his contributions to natura
history were sufficiently valued for some scientists to commemoiate him y naming
some plants in his honour.
Fieldia A.Cunn., in Field, Geographical Memoirs on New South Wales 364, t. (Apr. 1825).
Type: Fieldia australis A.Cunn.
Syn: Basileophyta F.Muell. Type: B. friderici-augusta F.Muell., J 5 ' Gen. Rept. Veg of the
Colony 16 (1853), nom invah, nom. nud.
Fieldia australis A.Cunn., Geographical Memoirs on New South Wales 364, t. (Apr. 1825).
Type: “Blue Mountains ... about twenty-two miles from the entrance of the new route
which has recently been traced out by Mr Bell, jr., 1823, A.Cunningham. It is foun
likewise at the Five Islands or Red-point of the charts.
Cunningham’s tribute to Barron Field is as follows: “The name now proposed for the
genus is intended to commemorate that of a gentleman who has, in his judicial
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capacity, much aided the advancement of the colony of New South Wales to its present
flourishing state; and whose important researches there, in various branches of
physical science, will materially tend to confer that interest upon our distant settlement
which it so richly deserves, and which yet remains in a great measure to he
appreciated.”
This was reinforced by Hooker in 1827 when he quoted Cunningham’s tribute and
stated in a concluding remark regarding his three volume work, Exotic Flora
(1821-1827): “To this work do I owe the acquaintance of the gentleman after whom
the present plant is named, together with the possession of a valuable collection of
New Holland plants, and the use of many excellent drawings made in that country.”
Another genus, Fieldia, was described by Gaudichaud-Beaupre and named in Field’s
honour: Fieldia Gaud., in Freycinet, Voy. Uranie 10: 424, t.36 (1829).
Type: Fieldia lissochiloides Gaud. Type: “In insulis Moluccis (Rawak).”
The name of this genus is a later homonym. The taxon is now regarded as a species of
the orchid genus Vandopsis (V lissochiloides (Gaud.) Pfitzer). Gaudichaud’s tribute to
Barron Field is as follows: “J’ai consacre ce genre, comme un temoignage de ma
profonde reconnaissance, a M. Barron Field, juge de la cour supreme au Port-Jackson,
qui nous a iacilite les moyens de traverser les Montagnes-Bleus.”
From this it is clear that Barron Field assisted members of Louis Freycinet’s team to
cross the Blue Mountains when they visited the Colony in 1819.
A species in the genus Cassia was also named in Field’s honour.
Cassia barronfieldii Colla [as barrenfieldii], Flort. Ripul. App. 2: 343 (Jan. 1826); App.
4: 23, t.l 1 (July 1830). [Dates follow Stafleu and Cowan (1976).]
Type: “Botany Bay.” Syrnon (1966) indicated that a holotype exists at TO. A specimen
at K (bearing no collector detail), transferred from TO, is regarded by Randell (1989)
as a syntype. Randell lectotypified the name by this specimen.
Named in “honorem Cl. Barrenfieldii hanc novam pulcherrimamque stirpem dicavi e
seminibus absque nimine specifico a Schultesio nissis enatam.” And, “in honorem cl. J.
C. Barrenfieldio utpote primus qui illam invenerit in regionibus Botany-Bay, ac semina
sine nominee praeclaro Schrankio miserit quae nobis humanissimus hie Professor
communicaverat.”
Bentham (1864: 285) treated this as a taxonomic synonym of C. australis Sims, Dot.
Mag. 53: t.2676 (Aug. 1826), and recorded the synonym as “afterwards corrected to
C. Fieldii ”. The correction, noted by Bentham, has not been located. Syrnon (1966: 102)
stated that he found no record of the change; Randell (1989) stated that it was a
“nomen nudum”; and Chapman (1991) does not record the name. Similarly any such
“correction” is not treated by Vogel (1837). The name C. Fieldii Colla is, however,
recorded by Steudel (1841) where he clearly refers back to the C. barrenfieldii of Colla
(without specified reference). He also refers to the record of C. barrenfieldii in Vogel.
We do not know if Steudel was first to “correct” the name. Nor do we know if Colla
published an earlier “correction”, as indicated in Index Kewensis (though the record in
Index Kewensis is erroneous). Until a specified correction is located in the literature, it
is impossible to be sure if C. fieldii is a nom. inval., norn. mid. or, a nom. illeg., nom.
Justice for Justice Barron Field
Telopea 11(1): 2005
97
superfl. It is also possible that the name is based on a different type, though Bentham’s
use of the term “corrected” does not lend support to this alternative.
More recently, the taxon to which this nomenclatural confusion applies has been
treated by Symon (1966) as a taxonomic synonym of Cassia odorata Morris,
FI. Conspicua: t. 57 (Sept. 1826) [date from Stafleu & Cowan (1976)]. This is currently
treated as Senna odorata (Morris) Randell (1989: 202); Harden (1991: 322); and
Orchard (1998: 98). The type citation for C. odorata is: “this plant has very recently
been introduced from New South Wales; and the present specimen was kindly
forwarded by Aylmar Bourke Lambert, Esq., from Boynton House, Wilts, where it
flowered in the greenhouse in the month of May.” Symon (1966) and Randell (1989)
have lectotypified this with the plate, no type specimen having been located. Whilst
there is no overt evidence, it is possible that the original material of this taxon was
collected by Field. Field did send collections to Lambert (see Miller 1970).
Conclusion
The Colla epithet predates that of Morris and must take precedence based on the dates
indicated by Stafleu and Cowan (1976) The correct name for this taxon is:
Senna barrotifieldii (Colla) Hewson comb. nov.
Basionym: Cassia barronfieldii Colla [as “barrenfieldii”], Hort. Ripul. App. 2: 343 (Jan.
1826). Type: “Botany Bay”; lecto: K ,fidc Randell (1989).
Cassia australis Sims, Bot. Mag. 53: t.2676 (Aug. 1826) nom. illeg. (non Vellosa (1825))
Type: “from New Holland”; lecto: plate 2676, fide Randell (1989).
Senna odorata (Morris) Randell, /. Adelaide Bot. Card. 12(2): 202 (1989), Cassia
odorata Morris, FI. Conspicua: t.57 (Sept. 1826). Type: “introduced from New South
Wales”; lecto: plate 57, fide Symon (1966), Randell (1989).
The spelling of Field’s Christian name is here corrected for the first time. Inadvei tently,
a nasty jibe was perpetuated by Colla. Amongst Field’s enemies in Australia he was
taunted with being “barren” of ideas (Elliott 1947), a cruel play on the name of a man
who was quite the opposite, as attested in his essays in the Geographical Memoirs. I he
man may have been irascible but his intellect is defendable. Indications in the literal y
world (Byrne 1961) that Field “hated botanists”, cannot be borne out. His conti ibution
to the knowledge of Australia’s flora through botanists, scientists and horticulturists
such as Allan Cunningham, William Hooker, Aylmer Lambert, Luigi Colla, Chailes
Gaudichaud, Josef Schultes and Franz Schrank (representing Australia, Britain, Italy,
France and Germany), demonstrate that his contribution was tangible and valued.
Acknowledgments
I wish to acknowledge the assistance provided by the facilities, library, staff and
associates of the Centre for Plant Biodiversity Research where I am an Honorary Associate.
98
Telopea 11(1): 2005
Hewson
References
Bentham G (1864) Flora Australimsis, vol. 2. (Lovell, Reeve & Co.: London)
Byrne JV (1961) Barron Field — Recultivated. Southerly 21(3): 6-18.
Chapman AD (1991) Australian Plant Name Index. (AGPS: Canberra)
Currey CH (1966) In Pike D. (ed.) Australian Dictionary of Biography 1: 73-376. (Melbourne
University Press: Melbourne)
Desmond R (1977) Dictionary of British and Irish Botanists and Horticulturists. (Taylor & Francis:
London)
Elliott B (1947) The First Austral Harmonist in Singing to the Cattle. (Georgian House: Melbourne)
Field B (1825) Geographical Memoirs on New South Wales. (Murray: London)
Harden GJ (1991) Flora of New South Wales, vol. 2. (New South Wales University Press: Kensington)
Hooker W) (1825-1827) Exotic Flora, vol. 3. (Printed for William Blackwood: Edinburgh; and T.
Cadell: London)
Miller HS (1970) The Herbarium of Aylmer Bourke Lambert. Taxon 19: 522.
Orchard AE (1998) (ed.) Flora of Australia, vol. 12. Mimosaceae (excl. Acacia), Caesalpiniaceae.
(CSIRO Publishing: Collingwood)
Randell BJ (1989) Revision of the Cassiinae in Australia. J. Adelaide Bot. Card. 12(2): 165-272.
Stafleu F & Cowan RS (1976) Taxonomic Literature 2 nd edn 1: 523.
Steudel EG (1841) Nomenclator botanicus 2 nd edn. (Stuttgardiae et Tubingae)
Symon D (1966) A revision of the genus Cassia in Australia. Trans. Roy. Soc. S. Australia 90: 73-146.
Vogel T (1837) Synopsis Generis Cassiae. (Logier: Berlin)
Manuscript received 26 May 2004, accepted 1 February 2005
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Telopea
Journal of Plant Systematics
11 ( 1 ): 1-98 • 2005
ISSN 0312-9764
Paetwla storyi, a new genus and species related to Ixodia and Haeckeria
(Asteraceae: Gnaphalieae) A.E. Orchard 1-9
Morphological and ontogenetic studies on the gynostemium of some
Australian members of Diurideae and Cranichideae (Orchidaceae)
H. Kurzwdl, P.H. Weston and A.]. Perkins 11-33
A new species of Goodenia (Goodeniaceae) from Nocoleche Nature
Reserve, Far Western Plains, New South Wales
Belinda }. Pellow and John L. Porter 35-41
Reassessment of Indigofera pratensis var. coriacea Domin and var.
angnstifoliola Domin (Fabaceae: Faboideae) with the recognition of a
new species Aniuska A. Kazandjian and Peter G. Wilson 43-51
Lectotypification of Schoenodum tenax (Restionaceae) and a note on
the type of Lyginia imberbis (Anarthriaceae) Barbara G. Briggs 53-58
New combinations and synonymies in the Australian Graphidaceae
A. W. Archer 59-78
A new species of Pomaderris (Rhamnaceae) from the Central Tablelands
of New South Wales J.C. Millott and K.L. McDougall 79-86
A colligate Spirogyra (Zygnemataceae, Zygnematophyceae) in Australia
Stephen Skinner and Timothy J. Entwisle 87-89
A new Zygnemopsis species (Zygnemataceae, Zygnematophyceae), with
mature zygospores, from Australia
Stephen Skinner, Hannah McPherson and Gillian M. Towler 91-93
Justice for Justice Barron Field Helen Hewson 95-98
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