Resolution of the Thelymitra variegata (Lindl.) F.Muell. (Orchidaceae) complex of southern Australia and New Zealand

2009·6.3 MB·English

by  J A Jeanes

Resolution of the Thelymitra variegata
(Orchidaceae) complex of southern
Australia and New Zealand
Jeffrey A. Jeanes
Royal Botanic Gardens Melbourne, Birdwood Avenue, South Yarra, Victoria 3141, Australia;
e-mail: [email protected]
Introduction Abstract
The eight currently known taxa in
Thelymitra J.R.Forst. & G.Forst. is a complex genus of orchids consisting
the Thelymitra variegata (Lindl.)
of about 100 described species, several described natural hybrids and
F.Muell. complex are reviewed and
at least 13 undescribed taxa. It is concentrated in higher rainfall areas of descriptions are presented for each.
temperate Australia, but a few species occur in tropical north-eastern Thelymitra pulcherrima Jeanes,
Australia, about 20 species occur in New Zealand (15 endemic) and a T. speciosa Jeanes and T. uliginosa
Jeanes from southwestern Western
few species occur in Indonesia, New Caledonia, New Guinea and the
Australia are described as new and
Philippines.
illustrated. Thelymitra spiralis (Lindl.)
This is one of a series of papers by the author reviewing the various
F.Muell. var. pulchella Nicholls is,
morphologically distinct groups or complexes within Thelymitra. raised to species rank and given the
During the course of my studies of the Thelymitra variegata (Lindl.) new name T. maculata Jeanes as the
F.Muell. complex it became evident that several undescribed species epithet'pulchella'is unavailable. The
key diagnostic features relating to
were present within the group. I take this opportunity to descibe these
the size and colour of the perianth,
new species and review the entire complex.
the size, colour and shape of the
There are a number of morphological features that, in combination, lateral lobes of the column and
distinguish members of the T. variegata complex from all other leaf characteristics are elucidated.
Thelymitra species. The leaf is usually variously auriculate at the base Information on distribution, habitat,
pollination biology, flowering time
and the blade is usually spirally twisted or at least somewhat curved.
and conservation status is given for
The leaf is also pubescent at the base with the hairs mainly on the veins
all eight taxa.The main distinguishing
and margins. The flowers usually have strong lustrous colours, often in
features of Thelymitra apiculata (A.S.
combinations of pink, red, yellow, mauve or purple. The column has a George) M.A.CIem. & D.LJones,
rudimentary post-anther lobe with an arc of small digitate to globose T. matthewsii Cheesem., T. spiralis,
glands arranged between the bases of the lateral lobes. The lateral T. variegata and the four new species
are tabulated. A key is provided to
lobes are well-developed and prominent, stipitate, suborbicular to
distinguish all eight members of the
narrow-elliptic and smooth to rugulose. The anther has an elongate
I variegata complex.
terminal beak that is essentially an extension of the connective (see
Muelleria 27(2): 149-170 (2009)
Fig. 1). Although six of the eight species in the complex are apparently
insect-pollinated, it is interesting to note that no floral odours have
ever been attributed to the group.The likely mechanism for pollination
is floral mimicry of other wildflowers (A. Brown pers. comm.)
Gardens
Melbourne
Muelleria 149
Jeanes
Taxonomic history Hoffman and Brown (1998) followed their earlier work
but also recognised two informal taxa, both as T. aff.
The first two species in the group to be recognised,
variegata, one from north of Perth, the other from the
Thelymitro variegata and T. spiralis (Lindl.) F.Muell.,
south coast of Western Australia. Brown et al. (2008)
were described by Lindley (1839-40) as Macdonaldia
recognised and illustrated the seven Western Australian
variegata Lindl. and M. spiralis Lindl. respectively
species described in this paper, either by their currently
(see Discussion for notes on Macdonaldia). Mueller
accepted names or the manuscript names ascribed by
(1865: pp 97-98) described T. porphyrosticta F.Muell.
the current author.
and commented that his new species was similar to
T. variegata and T. spiralis thereby effectively creating
Discussion
these two new combinations. Bentham (1873)
recognised only T. variegata while reducing T. spiralis Historically there has been significant confusion about
and T. porphryosticta to synonymy. Mueller (1882) the correct application of names and the interpretation
followed Bentham's classification. Cheeseman (1911) of species within the Thelymitra variegata complex.
described T.matthewsii Cheesem.from plants found on At least some of this confusion has persisted to the
the North Island of New Zealand and later Rogers (1930), present day and has revealed itself in the many floras
apparently unaware of Cheeseman's species, described and orchid books published over the years.
T. daltonii R.S.Rogers (as T. D'Altonii) from Halls Gap in Lindley (1839-40) described the first two species
the Grampians, Victoria. Nicholls (1949) created four in the group as Macdonaldia variegata and M. spiralis
varieties of T. spiralis—T. spiralis var. scoulerae Nicholls, respectively and distinguished these, and several
T. spiralis var. pallida Nicholls, T. spiralis var. punctata other apparently unrelated species, from Thelymitra
Nicholls and T. spiralis var. pulchella Nicholls—all of in having "the anther incumbent, and not parallel with
which, as well as the type variety, are well illustrated the stigma". The generic name Macdonaldia Gunn
in colour in Nicholls (1951). George (1971) recognised ex Lindl. was coined by the notable plant collector
all of Nicholls' varieties except var. pulchella, which Ronald Gunn in honour of Mrs Smith nee Macdonald
he relegated to synonymy under the type variety. He an orchid collector from Tasmania. Hooker (1858)
also relegated T. porphryosticta to synonymy under reduced Macdonaldia to a section within Thelymitra
T. variegata and recognised T. matthewsii as occurring differing from his other sections in having the "Column
in Western Australia. George (1984) described bifid at the apex, its sides not produced into toothed
T. variegata var. apiculata A.S.George from plants or feathery arms. Anther projecting, generally more
collected between Eneabba and Mogumber north or less granular or villous". Mueller (1865, p. 98) did
of Perth. Jones (1988) followed George (1971), but not recognise Macdonaldia and created the new
in a brief entry in the supplement at the back of the combinations T. variegata and T. spiralis. In the same
book T. variegata var. apiculata was raised informally article, Mueller (1865) described T. porphyrosticta
to species rank as T. apiculata (A.S.George) M.A.CIem. from specimens collected by Maxwell near Salt
& D.LJones. Following Clements' study of the various River and Kalgan in Western Australia. Following
type specimens, Clements and Jones in Clements my examination of the two type specimens at MEL, I
(1989) formally raised T. variegata var. apiculata to have concluded that one (Kalgan River) is referable to
species rank as T. apiculata. Otherwise Clements (1989) T. variegata (lectotypified below) the other (Salt River)
recognised only T. matthewsii, T. spiralis var. spiralis, to T. spiralis (rejected syntype). A specimen at Kew (not
T. spiralis var. pulchella and T. variegata as distinct seen by me) collected by Maxwell at Kalgan River is an
taxa. Hoffman and Brown (1992) recognised only isolectotype (George 1971). Bentham (1873) recognised
T. apiculata, T. spiralis and T. variegata as occurring in only T. variegata while relegating T. spiralis (and
Western Australia, although they do allude to T. spiralis T. porphyrosticta) to synonymy therein. Bentham (1873)
var. pulchella and the taxon from Western Australia that recognised Macdonaldia as a section within Thelymitra
was equated by George (1971) with T. matthewsii as and defined it as "Column-wing broadly produced
possibly being distinct entities worthy of recognition. behind the anther, but much shorter than it, and not
150 Vol 27(2) 2009
Table 1. Taxonomic history of the T. variegata complex showing the names in Macdonaldia and Thelymitra used by various authors for the eight taxa recognised in the current treatment
M
u Author Name used
e
lleria Lindley (1839-1840) M. variegata M. spiralis C '—. TO n/a C TO C — TO c — (cid:9632)to n/a
Lindley (1840) M. variegata M. spiralis C 1—. TO n/a C — TO n/a C X TO C — TO
Mueller (1865) T. variegata T. spiralis c 1— TO C \ TO n/a C — TO C TO C 1— TO
T. porphryosticta T. porphryosticta
Bentham (1873) T. variegata T. variegata C TO n/a C TO C TO n/a C TO
Mueller (1882) T. variegata I variegata n/a n/a n/a c C TO n/a
Cheeseman (1911) I variegata none T. matthewsii n/a C — TO n/a C —. TO C "X TO
Rogers (1930) T. variegata T. spiralis T. D'Altonii n/a C TO n/a C — TO n/a
Nicholls (1949), Nicholls T. variegata *T. spiralis var: spiralis T. matthewsii I spiralis var. n/a C TO n/a n/a
(1951) & Nicholls (1969) T. spiralis var. scoulerae pulchella
T. spiralis var. pallida
T. spiralis var. punctata
George (1971) T. variegata T. spiralis var. spiralis T. matthewsii T. spiralis var. n/a T. matthewsii n/a C TO
T. spiralis var. scoulerae spiralis
T. spiralis var. pallida
T. spiralis var. punctata
T. spiralis var. pulchella
Hoffman & Brown T. variegata T. spiralis none none T.sp T. matthewsii C '— TO C "** (cid:9632)TO
(1984)
George (1984) T. variegata var. none none none T. variegata none n/a C TO
variegata var. apiculata
Jones (1988) T. variegata *T. spiralis var. spiralis T. matthewsii none T. variegata T. matthewsii n/a C TO
T. spiralis var. scoulerae var. apiculata
T. spiralis var. pallida T. apiculata T
h
T. spiralis var. punctata e
ly
m
Clements (1989) T. variegata TT.. ssppiirraalliiss vvaarr.. sspciorualliesr ae T. matthewsii. pT.u slpcihreallilsa var. T. apiculata T. matthewsii n/a n/a itra v
T. spiralis var. pallida arie
T. spiralis var. punctata g
a
ta
Hoffman & Brown T. variegata T. spiralis none *T. spiralis var. T. apiculata ?T. matthewsii n/a n/a (Q
(1992) ?T. matthewsii pulchella rc
h
Hoffman & Brown T. variegata T. spiralis none *T. spiralis var. T. apiculata ?T. matthewsii T. aff. variegata T. aff. variegata ida
c
(1998) ?T. matthewsii pulchella ea
e
Brown etal. (2008) T. variegata T. spiralis none T. maculata ms T. apiculata T. uliginosa ms T. speciosa ms T. pulcherrima ms ) c
o
m
15 Jeanes (this paper) T. variegata T. spiralis T. matthewsii T. maculata T. apiculata T. uliginosa T. speciosa T. pulcherrima ple
1 * z o *-> E a> c o C QJ T3 io Q. a» u U TO >> X 1c io C TO E dJ -Q -* (cid:9830)_c (V T3 <V c >> o <4— jc <V TO X o c E "d. ;g c sz <v u o c OI X x
Jeanes
hood-shaped. Slender flexuose herbs, with the habit T. spiralis) that I believe it warrants recognition at species
of T. antennifera". Thelymitra carnea R.Br. and T. flexuosa rank. The epithet 'pulchella' is, however, unavailable at
Endl. were the only species Bentham included within species rank due to the presence of T. pulchella Hook,
this section, while he placed I variegata in his section f. from New Zealand so a new name had to be coined
Biaurella along with four apparently unrelated species. (see T. maculata Jeanes below).
Few authors since Bentham have given Macdonaldio George (1984) described T. variegata var. apiculata
any formal recognition. from plants collected near Badgingarra homestead
The description of T. motthewsii (Cheeseman 1911) north of Perth. These differed from the type variety
brought the number of generally recognised species primarily in the presence of apiculate lateral lobes on
within the group to three. Apparently Cheeseman the column and the less obviously spiralled leaf blade.
believed his new species to be endemic to New Zealand The distinctiveness of this taxon was so significant that
as there is no mention in his article about it occurring in it was soon raised to species rank (Clements 1989) as
Australia. When Rogers (1930) described T. daltonii (as T. apiculata.
T. D'Altonii) from the Grampians in Victoria, he was either Hoffman and Brown (1998) illustrated and informally
unaware of Cheeseman's article (he makes no mention described two apparently distinct taxa related closely
of it in his protologue) or he believed T. motthewsii to T. variegata, both as T. aff. variegata. My own research
to be a New Zealand endemic and chose to ignore has shown these two allopatric taxa to be sufficiently
it. Whatever the case may be it soon became evident distinct morphologically, ecologically and by flower
that the two names were synonymous, as indicated colour to warrant recognition at species rank (see
by Nicholls (1949, 1951; 1969). George (1971) alluded T. speciosa Jeanes and T. pulcherrima Jeanes below).
to the presence of T. motthewsii in Western Australia Szlachetko (1995) recognised Macdonaldia as a
based on plants discovered to the east of Perth and genus distinct from Thelymitra based on differences
illustrated later in Hoffman and Brown (1984). My study in the column structure. He recognised 13 species in
of these plants and others from several localities in the Macdonaldia, and created nine new combinations (one
Albany area has shown them to be of an undescribed of which, M. venosa (R.Br.) Szlach., was superfluous),
species distinct from T. motthewsii (see T. uliginoso but failed to list all the species belonging to that
Jeanes below). genus according to his concepts. Szlachetko and
Nicholls (1949) described and illustrated four new Rutkowski (2000) again recognised Macdonaldia as
varieties of T. spiralis - var. scouleroe, var. pallida, var. distinct from Thelymitra, but did not list the species
punctata and var. pulchella - the first three based on belonging to either genus according to their concepts.
plants collected by Mrs E. Scouler at Yarloop, the last It is clear from their generic key that some species,
from plants collected near Bolgart by Mrs R. Erickson, traditionally regarded as belonging to Thelymitra,
all from Western Australia. George (1971) and Clements have characteristics of both their Macdonaldia and
(1989) differed in their interpretation of the holotype Thelymitra and it would be difficult to place these
of T. spiralis hence their interpretation of Nicholls' species satisfactorily in either genus. The classification
varieties also differed. The former interpreted the of Szlachetko and Rutkowski has not been taken up by
type of T. spiralis as being the same taxon as Nicholls' later workers on the group including the current author.
var. pulchella and hence relegated that variety to All the definitions of Macdonaldia by the variousauthors
synonymy, while maintaining Nicholls' other three over the years, whether at generic or sub-generic level,
varieties as distinct. Clements (1989) interpreted the have varied considerably. Significantly, the type species
type of T. spiralis as being distinct from Nicholls' var. of Macdonaldia was designated by Clements (1989) as
pulchella but rather similar to the other three varieties M. smithiana Gunn ex Lindl. (=T. flexuosa), a species
of Nicholls, which he consequently did not recognise not belonging to the T. variegata complex. Preliminary
as distinct taxa. In this matter my interpretation of phylogenetic studies of Thelymitra based on ribosomal
the types follows that of Clements. In fact Nicholls' DNA confirm that Macdonaldia is paraphyletic and
var. pulchella is so distinctive (and different from that all its elements are embedded within Thelymitra,
152 Vol 27(2) 2009
Thelymitro variegata (Orchidaceae) complex
and that species in the T. variegoto complex form a the point of insertion of the anther, with an arc of small
separate clade embedded deeply within the genus. (M. ovoid to digitate glands on the dorsal surface.
Clements pers. comm.) Lateral lobes (column-arms or lateral staminodes):
These two structures lie one on each side of the
Explanation of the terminology used post-anther lobe and extend forward or upward and
often converge. They are each supplied by a single
Thegenus Thelymitra is unusual in theOrchidaceaein that
unbranched vascular bundle and are thought to
the six perianth segments generally differ very little from
represent staminodes. They may be flat and ribbon¬
each other in terms of size, shape and ornamentation.
like, terete and finger-like, straight, curved, twisted
The labellum does not bear any hairs, calli, glands, ridges,
spirally or bent sharply, and are usually ornamented
lobes, teeth or fringes and is apparently not involved in
with lobes, teeth, tubercles or trichomes. The lateral
pollination. Since the perianth is virtually actinomorphic
lobes are suborbicular to elliptic in members of the
and generally lacks characters by which to distinguish
T. variegata complex, are more or less straight, parallel
the species, traditionally the structure of the column
or divergent and point in the same general direction as
has supplied most of these distinguishing characters.
the body of the column or slightly forward.
Over the years a terminology has evolved to describe
Auxiliary lobes (accessory lobes or side lobules):
the column structure in Thelymitra, but some of these
Several species of Tfie/ym/fra have a pair of distinct lobes
terms are poorly understood and some have never been
between the post-anther lobe and the lateral lobes.
defined adequately. Below is an explanation of some of
These have no vascular strand and are most accurately
the terms commonly used in this paper; most have a
described as being part of a tripartite post-anther lobe.
traditional usage, although this has often not been well
They tend to be fleshy with irregularly jagged margins
understood.
and sometimes have small surface tubercles. In the
Column (gynostemium): The column is exposed in
T. variegata complex the auxiliary lobes are completely
the centre of the flower, it lacks a free filament and style,
absent.
is short and thick and broadly winged from below the
Anther. In Thelymitra, the anther is usually small,
stigma to the level of the anther or beyond. The apex
ovoid, and situated entirely between the column
is usually 3-5-lobed and is often ornamented with
wings. The connective extends beyond the pollinia
trichomes, fringes, teeth, calli, glands, tubercles or lobes.
into an apical beak-like projection of varying size. The
In members of the T. variegata complex the apex of the
anther may be entirely above the stigma or variously
column has a pair of prominent suborbicular to elliptic
obscured behind it. In the T. variegata complex the
lateral lobes and a rudimentary post-anther lobe.
anther is usually inserted near the apex of the column,
Post-anther lobe (mid-lobe): This structure lies
the pollinia usually held mostly above the stigma. The
beyond the point of insertion of the anther and of the
anther beak is very well developed, usually being about
lateral lobes, and it is usually of a different colourto the
as long as the pollinia, is fleshy and more or less straight
rest of the column. It has a complex vascular supply
or slightly curved forward when seen in profile.
always associated with that of the functional anther
Pollinia: Members of the genus Thelymitra possess
and may be regarded as an outgrowth of the filament.
four pollinia in two groups of two. In the T. variegata
In some species it is represented only by a short flap
complex the pollinia are usually tightly bound with the
or a band of small glands crowded across the back
pollinarium being removed by insects as a single unit.
of the anther. In most species it extends well beyond
However, autogamy has been noted in T. matthewsii
the anther with a free margin that may be plain,
and is believed to occur in T. uliginosa.
undulate, toothed, notched or variously ornamented
Stigma: The stigma in Thelymitra is more or less
with tubercles. At its maximum development it forms
bi-lobed at the apex, usually quadrate or transverse-
a fleshy, tubular hood that is variously open on the
elliptic in shape and located at the base of the column
ventral side and overhangs and obscures the anther.
on a thick stalk.
In the T. variegata complex the post-anther lobe is
rudimentary, extending only a short distance beyond
Muelleria 153
Jeanes
Lateral lobes
Glands
Viscidiu
Figure 1-3.1. Column features of Thelymitra variegata. From left to right: ventral, lateral and dorsal views
(photographs by Jeff Jeanes); 2. Thelymitra spiralis (photograph by Ron Heberle);
3. Thelymitra maculata (photograph by Viv Holly).
154 Vol 27(2) 2009
Thelymitra variegato (Orchidaceae) complex
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Muelleria 155
Jeanes
Key to the known members of the T. variegata complex.
1 Lateral lobes of column <3 mm long; anther connective produced into a beak to 1.5 mm long.
1: Lateral lobes of column 3-6 mm long; anther connective produced into a beak usually >1.5 mm long.
2 Perianth segments mostly to 12 mm long, rarely to 15 mm long, unspotted.
2: Perianth segments mostly >12 mm long, often spotted ..~.
3 Flower solitary, very rarely 2, deep purplish, strongly striated; petals and sepals usually to 10 mm
long, of contrasting colour; dry open forests; South Australia, Victoria and New Zealand.4. T. matthewsii
3: Flowers 1 or 2, pink, mauve or pale purplish, not strongly striated; petals and sepals often more
than 10 mm long, of similar colour; swamp margins; Western Australia.3. T. uliginosa
dark pink or purplish in longitudinal lines; leaf-base not strongly auriculate, lamina often curved
but not tightly spiralled; lateral lobes often suborbicular, orange or yellow; dry inland habitats.2. T. maculata
4: Flowers 1 -3, colour not as above; leaf-base strongly auriculate, lamina usually tightly spiralled;
lateral lobes usually elliptic, yellow; more mesic near-coastal habitats.1- T. spiralis
5 Lateral lobes of column with a terminal needle-like point; leaf-base not strongly auriculate,
lamina curved, not tightly spiralled.5. T. apiculata
5: Lateral lobes of column lacking a terminal needle-like point; leaf-base strongly auriculate,
lamina tightly spiralled or not...—.6
6 Flower usually solitary, rarely 2; perianth segments predominantly purplish, with broad,
deep reddish gold edges and yellowish margins; found in heavy clay-loam soils.7. T. speciosa
6: Flowers 1-6; perianth segments not coloured as above; found in lateritic soil or deep sand.7
7 Perianth segments mostly to 18 mm long, of contrasting colour; sepals predominantly
reddish brown with yellow margins; petals predominantly purplish; found in lateritic soil;
flowers June to early August ....8. T. pulcherrima
7: Perianth segments mostly >18 mm long, more or less of similar colour, predominantly reddish,
purplish or violet, variegated, mostly with darker spots or blotches, often with yellow margins;
found in deep sand; flowers August to early October.6. T. variegata
Materials and methods and spirit-preserved herbarium material (including
type specimens) much easier.
This paper is the result of a qualitative and quantitative
When collecting Thelymitra for study it is essential
study of the pertinent type material (or photographic
that the entire above ground parts of the plant be taken,
reproductions thereof), hundreds of herbarium
with the majority of the material being preserved in spirit.
specimens (both dry and spirit-preserved) from AD, BM,
Plants preserved in the pressed state are often difficult
BRI, CANB, E, HO, MEL, NSW, P, PERTH, QRS, SUNIV and
to identify to species level in the absence of additional
WELT, and numerous freshly collected specimens, all of
information. Spirit-preserved specimens, on the other
which were vouchered and deposited at the relevant
hand, are generally much more easily identified to
herbaria. Orchid taxa in general, and Thelymitra taxa
species level. The observation of plants growing in-situ is
in particular, are much more readily identified from
the ideal method of study for Thelymitra in general, and
fresh living material where characters of the perianth,
often it is only by this method that cryptic new species
the column, flower colour and fragrance are still intact.
can be identified. For this reason the importance of field
Familiarity with the taxa gained from field study and
work in the study of species complexes within Thelymitra
the study of freshly collected specimens sent to me by
cannot be overstated and should form an integral part of
field operatives has made the identification of dried
156 Vol 27(2) 2009
Thelymitra variegota (Orchidaceae) complex
any future studies of the group. It is possible that other acute to acuminate. Pedicels 7-22 mm long, slender.
taxa worthy of recognition exist within this complex, Ovary narrow-obovoid, 5-15 mm long, 1.5-4 mm
but adequate information and collections of these are wide. Flowers 1-2(-3), 21-40(-50) mm diameter, pink,
lacking at present. reddish, purplish or blue, sometimes with darker spots
or longitudinal veins, opening freely in warm weather.
Taxonomy Perianth segments 8-20(-26) mm long, 2-10 mm wide,
concave to almost flat, stiffly spreading, often shortly
1. Thelymitra spiralis (Lindl.) F.Muell., Fragm. 5:
apiculate; dorsal sepal ovate to ovate-lanceolate, acute
98(1865)
to subacute, usually broader than other segments;
Macdonaldiospiralis Lindl., Edwards's Bot. Reg. appendix lateral sepals ovate-lanceolate to lanceolate, slightly
to vols 1-23 [Sketch l'eg. Swan /?.]: 50 (1839-40). asymmetric, acute to acuminate; petals ovate-
Type: Swan River, 1839, J. Drummond s.n. (holotype K- lanceolate to lanceolate, slightly asymmetric, acute to
LINDL!, isotype BM!). acuminate; labellum lanceolate to linear-lanceolate,
Thelymitra porphyrosticta F.Muell., Fragm. 5: 97-8 acute to acuminate, smaller than other segments.
(1865) p.p. not as to type (see under T. variegata). Column erect from the end of ovary, 4.5-7 mm long,
Thelymitra spiralis (L i n d I.) F. M u e 11. va r. pallida N i c h o 11 s, 2.5-3.5 mm wide, broadly winged, similarly coloured
Victorian Naturalist 66:55 (1949). Type: Yarloop, c. 1946, to perianth; post-anther lobe vestigial, apical margin
E. Scoulers.n. (holotype MEL!). covered with an arc of digitate to globose glands 0.2-
Thelymitra spiralis (Lindl.) F.Muell. var. punctata 0.6 mm long; auxiliary lobes absent; lateral lobes more
Nicholls, Victorian Naturalist 66:55 (1949). Type: Yarloop, or less parallel, ovate or elliptic, 1.5-2.8 mm long, 0.8-
c. 1946, E. Scoulers.n. (holotype MEL, not found). 1.6 mm wide, erect or obliquely erect, stipitate, fleshy,
Thelymitra spiralis (Lindl.) F.Muell. var. scoulerae rugulose, yellow. Anther inserted at apex of column,
Nicholls, Victorian Naturalist 66:55 (1949). Type: Yarloop, obloid, 2.4-3.7 mm long, 1.5-2.5 mm wide, projecting
ix.-x.l 946, E. Scoulers.n. (holotype MEL!). forward at about 90° to column, yellow, the connective
Illustrations: Fitzgerald (1891) 2: 4 (as T. variegata); produced into a fleshy beak 0.5-1.5 mm long, dorsal
Nicholls (1951) plate 45, all except figs c & g; Nicholls surface pubescent, ventral surface channelled;
(1969) plate 47, all except figs c&g; Jones (1988) page pollinarium 1.5-2.5 mm long; viscidium elliptic, c. 0.7
304; Hoffman and Brown (1984) page 42; Hoffman and mm long, c. 0.5 mm wide; pollinia coherent, cream.
Brown (1992) page 253; Hoffman and Brown (1998) Stigma situated c. midway along column, orbicular to
page 253; Heberle (2000) page 288; Jones (2006) page transversely broad-elliptic, c. 2.5 mm long, c. 2.5 mm
254; Brown etal. (2008) page 321. wide, concave, margins irregular. Capsules obovoid,
10-20 mm long, 4-8 mm wide, erect, ribbed. (Fig. 2,
Virtually glabrous terrestrial herb. Tubers ovoid, 1-3 cm
long, 4-10 mm wide, fleshy. Leaf ovate-auriculate at Fig. 10 a-c)
base, narrowing abruptly and linear above, 5-10 cm long, Selected specimens examined: WESTERN AUSTRALIA:
4-10 mm wide, usually spirally twisted, canaliculate, Kelmscott, 15.ix.1924, G.Coghills.n. (MEL 2016031); Armadale,
fleshy, dark green with a purplish base, sheathing 22.ix.1946, W.H. Nicholls s.n. (MEL 643870); Two People Bay-
Nanarup, 14.viii.1963, R. Oliver s.n. (PERTH 276723); 6 miles N
at base where margins often lobed and undulate,
of Porongurups, on Stirling Rd, 14.viii.1958, AS. George s.n.
pubescent towards base with hairs mostly on veins
(PERTH 277185); NW slopes of Mt Manypeaks, 11 .viii.1986, G.J.
and margins, apex subacute. Inflorescence 12—30(—45) cm
Keighery 9660 (PERTH 855162); Kenwick, 1919, A.H. Mann s.n.
tall, more or less straight. Scape 0.6-2 mm diam., wiry,
(PERTH 306541); Cannington, 3.X.1900, A. Purdie s.n. (PERTH
green or purplish. Sterile bract solitary, lanceolate to
276758); Swamp at Cannington, 14.ix.1920, G.N.O. s.n. (PERTH
ovate, 15-55 mm long, 3-9 mm wide, green or purplish,
672343); Serpentine, 23.ix.1904, A. Purdie s.n. (PERTH 294985);
closely sheathing at base, apex diverging from scape,
North Dwellingup, peaty flats W of Railway Station, x.1932,
acute to acuminate. Fertile bracts ovate to obovate, B.T. Goadby s.n. (PERTH 276790); Albany, 8.viii.1963, R. Oliver
6-22 mm long, 3-7 mm wide, green or purplish, s.n. (PERTH 276774); West Walpole, opposite Jarrah Road,
sheathing the pedicels, margins connate at base, apex 21.X.1993, D.L Jones 12459 (CANB 9710428); E of Alexandra
Muelleria 157
Jeanes
Bridge, over Blackwood River, on Brockman Highway, 2. Thelymitra maculata Jeanes nom. etstat. nov%
21 .x.1975, D.F. Blaxell W75/206 & A.S. George (NSW 462466).
Basionym: Thelymitra spiralis (Lindl.) F. Muell. var.
Distribution and habitat: Endemic to southwestern
pulchella Nicholls, Viet. Naturalist 66: 56 (1949).
Western Australia, from just north of Perth to Israelite
Type: Open plains or sand covered gravel, near Bolgart,
Bay, generally within about 100 km of the coast, with
18.viii.1934, R. Erickson s.n. (lectotype QRS 044565! hie
isolated northern populations in the Geraldton to
designates). Syntype: Open plains or sand covered gravel,
Kalbarri region. Grows mostly in and around seasonal
near Bolgart, viii.1949, R. Erickson s.n. (MEL not found).
swamps in moist sandy clay soils. Altitude: 10-300 m.
Illustrations: Erickson (1951) plate 6:20 (as T. spiralis
Conservation status: Uncommon to rare, but
var. pulchella); Nicholls (1951) plate 45, figs c & g (a$
extremely widespread and well conserved. Suggest
T. spiralis var. pulchella); Nicholls (1969) plate 47, figs
3RC by criteria of Briggs and Leigh (1996) and Near
c & g (as T. spiralis var. pulchella); Heberle (2000) pag$
Threatened (NT) by criteria of IUCN (2001).
250 (as T. spiralis); Jones (2006) page 253; Brown et al.
Flowering period: July to October, but mostly
(2008) page 321.
August and September.
Pollination biology: The large, freely opening Virtually glabrous terrestrial herb. Tubers obloid to
flowers, functional viscidium, coherent pollen and ovoid, 1-1.5 cm long, 3.5-6 mm wide, fleshy. Leaf
ovate at base, narrowing abruptly and linear to linear-
sporadic production of seed capsules would indicate
lanceolate above, 5-8 cm long, 3-6 mm wide, curved
that this species is probably entomophilous.
Notes: Thelymitra spiralis is extremely variable, or somewhat spirally twisted, canaliculate, fleshy, dark
particularly in terms of flower size and colour and green with a purplish base, sheathing at base where
degree of spotting and veining of the perianth. It is margins sometimes shallowly lobed and undulate,
most closely related to T. maculata but the latter usually pubescent towards base with hairs mostly on veins
has a single, generally smaller flower that is strongly and margins, apex subacute. Inflorescence 9-20 cm
spotted, particularly on the sepals, somewhat deflexed tall, more or less straight. Scape 0.5-1 mm diam., wiry,
petals and lateral sepals, a narrower column often green or purplish. Sterile bract solitary, lanceolate
with suborbicular, orange lateral lobes, a less spirally to ovate, 15-30 mm long, 3-5 mm wide, green or
twisted leaf that is barely auriculate at the base and a purplish, closely sheathing at base, apex diverging
preference for drier inland habitats. Thelymitra spiralis from scape, acute to acuminate. Fertile bract ovate
is also related to 7*. matthewsii and T. uliginosa, but the to obovate, 6-15 mm long, 3-5 mm wide, green or
latter two species have generally smaller flowers that purplish, margins connate at base, sheathing the
are non-spotted and are self-pollinated. pedicels, apex acute to acuminate. Pedicels 5-16 mm
Four varieties of T. spiralis have been described long, slender. Ovary narrow-obovoid, 4-10 mm long,
(Nicholls 1949).The var. pallida only differs from typical 1.5-3 mm wide. Flower usually solitary, 21 -35(-45) mm
plants in having pale flowers, the var.pt/nctata in having diameter, pale to deep pink or purplish, sepals often
highly spotted flowers and the var. scoulerae in having darker than petals, usually with darker pink to purplish
larger, brightly coloured flowers with broad, somewhat spots and blotches arranged in longitudinal lines on
flaccid perianth segments. These three varieties were sepals and sometimes also on petals, opening freely
later reduced to synonymy (Clements 1989) as they in mild weather. Perianth segments 8—18(—25) mm
show no great departure in floral morphology from long, 2-8 mm wide, concave to almost flat, petals and
typical plants and are not distinct ecologically. The var. lateral sepals often somewhat deflexed; dorsal sepal
pulchella is ecologically and morphologically distinct ovate to ovate-lanceolate, acute to subacute, usually
from T. spiralis, and hence was recognised by Clements broader than other segments; lateral sepals ovate-
(1989) as a variety of T spiralis and is herein raised to lanceolate to lanceolate, slightly asymmetric, acute
species rank as T. maculata (see below). to subacute; petals ovate-lanceolate to lanceolate,
slightly asymmetric, acute; labellum lanceolate to linear-
lanceolate,acute toobtuse,smallerthanothersegments.
Column erect from the end of ovary, 4.5-6.5 mm long,
158 Vol 27(2) 2009