See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/236705597
A revision of Pultenaea (Fabaceae: Mirbelieae).
4. Species occurring in Western Australia
Article in Australian Systematic Botany · May 2005
DOI: 10.1071/SB04029
CITATIONS
READS
4
66
3 authors, including:
Lindy Orthia
Australian National University
17 PUBLICATIONS 100 CITATIONS
Rogier de Kok
70 PUBLICATIONS 380 CITATIONS
SEE PROFILE
SEE PROFILE
All content following this page was uploaded by Rogier de Kok on 27 November 2016.
The user has requested enhancement of the downloaded file. All in-text references underlined in blue
are linked to publications on ResearchGate, letting you access and read them immediately.
CSIRO PUBLISHING
www.publish.csiro.au/journals/asb
Australian Systematic Botany 18, 149–206
A revision of Pultenaea (Fabaceae: Mirbelieae).
4. Species occurring in Western Australia
L. A. OrthiaA,B,D , R. P. J. de KokB,C and M. D. CrispA
A School
B Centre
of Botany and Zoology, Australian National University, Canberra, ACT 0200, Australia.
for Plant Biodiversity Research, Australian National Herbarium, CSIRO, GPO Box 1600,
Canberra, ACT 2601, Australia.
C Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AD, UK.
D Corresponding author. Email: lindy.orthia@anu.edu.au
Abstract. Twenty-five species of Pultenaea endemic to Western Australia are revised at and below species level.
Three species (P. daena Orthia & Chappill, P. indira Orthia & Crisp, P. wudjariensis Orthia), three subspecies
(P. calycina subsp. proxena Orthia & Chappill, P. indira subsp. monstrosita Orthia, P. indira subsp. pudoides Orthia)
and two informal taxa [P. ericifolia ‘southern’ (Orthia 39), P. sp. Mt Lesueur (Beard 7827)] are described for the
first time. The status of all varieties of P. verruculosa is changed: P. verruculosa var. brachyphylla (Turcz.) Benth. is
reinstated at species level, P. verruculosa Turcz. var. verruculosa and P. verruculosa var. pilosa Benth. are placed in
synonymy of Pultenaea verruculosa Turcz., and P. verruculosa var. recurva Benth. is found to be a nomen dubium.
Three names in common usage are found to be illegitimate: P. conferta Benth. [now P. purpurea (Turcz.) Crisp &
Orthia], P. obcordata (R.Br.) Benth. (now P. quaerita Orthia) and P. drummondii Meisn. (older name P. brachytropis
Benth. reinstated). The Pultenaea species occurring in both western and eastern Australia are discussed and addenda
made to the treatments previously published by de Kok and West (2003, 2004). The status of P. vestita R.Br. and
P. juniperina Labill. as naturally occurring WA species is questioned.
Introduction
Pultenaea Sm. is an Australian endemic genus of shrubs
restricted to south-west Western Australia and the temperate
and subtropical regions of the eastern states. Its species are
found in sclerophyllous vegetation types varying from forests
to heathlands, and are absent from the arid interior, the tropics
and rainforest tracts. Commonly known as the ‘bush peas’ or
‘egg and bacon’, Pultenaea species can play an important
ecological role as understorey dominants, nitrogen fixers and
as a food source for terrestrial invertebrates (Froggatt 1892;
Clemson 1985; Auld 1991). Thirty-one Pultenaea taxa are
listed on commonwealth- or state-based threatened species
lists, so the genus also has conservation significance.
Pultenaea has a high degree of east v. west endemism,
with approximately 26 species restricted to the south-west,
approximately 87 species restricted to areas east of the
Nullarbor Plain and only approximately four species found
in natural populations on both sides of the Nullarbor. The
Pultenaea species found in the eastern states were revised
by de Kok and West in three recent papers (2002, 2003,
2004). The present paper completes the species-level revision
of Pultenaea, by giving full taxonomic treatment to the
species endemic to the south-west, and updating WA-specific
© CSIRO 20 May 2005
information for the species that occur in both western and
eastern Australia.
Relationships of the study group
It should be noted from the outset that a recent cladistic study
with DNA sequences from three regions of the genome found
no support for the monophyly of Pultenaea (Orthia et al.
2005). Most Pultenaea species fell into six well-supported
lineages, each equivalent in genetic diversity and time of
origin to several other Mirbelieae genera (Orthia et al. 2005).
Given this, there is an argument to be made for recognising
each of the distinct phylogenetic units in classification, e.g. to
split Pultenaea into different genera. In the past, anomalous
groups of Pultenaea species have been transferred to
resurrected or novel genera: Urodon Turcz. (Ewart and White
1908; Crisp and Weston 1987), Almaleea Crisp & P.H.Weston
(Crisp and Weston 1991) and Stonesiella Crisp & P.H.Weston
(Crisp et al. 1999).
Genetic data also suggest that two Pultenaea species might
be better placed in other genera: P. adunca in Almaleea and
P. barbata in Phyllota (D.C.) Benth. (Orthia et al. 2005).
A previous study suggested that P. neurocalyx Turcz. might
be better placed in Eutaxia R.Br. (Crisp et al. 1999), a move
10.1071/SB04029
1030-1887/05/020149
150
L. A. Orthia et al.
Australian Systematic Botany
weakly supported by the more recent work (Orthia et al.
2005). Ideally, these affiliations, too, would be recognised
in classification.
However, Crisp and Cook (2003) recently proposed that
19 Mirbelieae genera, collectively known as the ‘Mirbelia
group’, be lumped into a single genus. That large genus
would be called Pultenaea, as it is the oldest name. The
implication of that proposal is that current genera such as
Dillwynia, Jacksonia and Gastrolobium will become part of
Pultenaea. Regardless of this, the present paper is written
with the understanding that a species-level revision for the
current Western Australian Pultenaea is useful irrespective
of classification changes above species level.
The species included for revision in this paper were
initially grouped together by default, because de Kok and
West’s revision series (2002, 2003, 2004) only dealt with
the species found east of the Nullarbor Plain. However,
genetic data support the treatment of the western species
separately from the eastern species to some extent. The
six main Pultenaea lineages mentioned above are largely
restricted to either the east (three lineages) or the west
(three lineages, plus a monotypic P. brachytropis); no clade
contains both endemic eastern and endemic western species
(Orthia et al. 2005). In other words, all endemic western
species revised here are genetically distinct from all endemic
eastern species.
The genetically anomalous taxa P. adunca and P. barbata
are included here, along with brief addenda to the previously
revised species P. elachista, P. tenuifolia, P. vestita and
P. juniperina, in order to complete the revision of Pultenaea.
Pultenaea neurocalyx was not included in the revision
because a previous study indicated that it is probably a
Eutaxia (Crisp et al. 1999; Orthia et al. 2005), and its
species level taxonomy is currently the subject of a study
by G. Henderson and J. Chappill (Henderson 1998).
Previous taxonomic work on the WA Pultenaea species
Understanding of the species-level taxonomy of WA
Pultenaea has changed little since Bentham’s Flora
Australiensis (1864). In that work, Bentham revised
19 WA species. In the process, he lumped two Western
Australian endemic genera, Euchilus R.Br. and Urodon, into
Pultenaea. He also reduced P. brachyphylla to a variety
of P. verruculosa, and described two further varieties of
the species, P. verruculosa var. pilosa and P. verruculosa
var. recurva. The next author to revise the WA Pultenaea
species was Williamson (1920, 1921, 1922), who included
five species described after Bentham’s time, as well as
describing a new species, Pultenaea radiata, but otherwise
did not stray far from Bentham’s treatment. Williamson
excluded Urodon spp. from Pultenaea without comment,
presumably being familiar with the work of Ewart and White
(1908), who reinstated the genus. The next two treatments
of WA Pultenaea were both brief: a census (Gardner 1930)
and an illustrated key in How to know Western Australian
wildflowers (Blackall and Grieve 1959). Both works made
some interesting decisions—naming some new Pultenaea
species, and transferring Urodon and Phyllota species
to Pultenaea—but neither included detailed descriptions
of species, authorities for names or explanations for the
taxonomic decisions made. A new treatment of Pultenaea
in the revised edition of ‘How to know Western Australian
wildflowers’ (Grieve 1998) was more comprehensive than its
predecessor, with brief descriptions accompanying a revised
key, including distribution, phenology and conservation
status information. This text eliminated the invalid species
included in the first edition, recognised Urodon as a separate
genus and reinstated Phyllota. The list of species included
under Pultenaea in this text is almost identical to Bentham’s
(1864) list, with one major exception: the reinstatement
of Urodon.
There is evidence to suggest that Bentham’s classification
is no longer adequate. Some workers have informally
proposed changes to the classification but not formalised
them, for example T. Macfarlane informally lumped
P. ericifolia and P. strobilifera as subspecies of P. ericifolia but
never published the proposal. Also, conservation biologists
have speculated that the endangered species P. pauciflora
may have infraspecific taxa (T. Macfarlane, pers. comm.).
Williamson’s revision (1920, 1921, 1922) included five
western species described after Bentham’s time, but his
descriptions generally are very brief, as are the descriptions
of Blackall and Grieve (1959) and Grieve (1998). Therefore
there is no comprehensive taxonomic treatment available
for the WA Pultenaea. de Kok and West (2002) note that
over 350 taxa have been described under Pultenaea, and
their treatment described several new taxa, reduced many
names to synonymy and found several nomenclatural and
typification problems. There is no reason to assume that
the WA Pultenaea should be exceptional, so a new revision
is timely.
Species concept used to delimit taxa
It was considered ideal to delimit species on the basis of
reproductive isolation (see Lee 2003), i.e. by means of
the biological species concept (Mayr 1963), but it was
beyond the scope of this study to measure reproductive
isolation / compatibility directly. Instead, morphological
data were used as proxies to estimate reproductive isolation.
Morphotypes that were consistently morphologically
distinct from all other taxa in at least two independent
traits were considered to be reproductively isolated, and
therefore different species. Specimens that were intermediate
between two different morphotypes were considered to be
evidence that the morphotypes were reproductively
compatible, and therefore the same species. If the
intermediate specimens showed a cline or continuum
of variation, preventing the identification of a clear
Revision of Pultenaea: Western Australian species
boundary between morphotypes, then no infraspecific
distinctions were made. If the two morphotypes maintained
their distinctness most of the time, with only occasional
intermediates to indicate reproductive compatibility, then
they were classified as subspecies. DNA sequences were
consulted for morphotypes whose classification status
at species level remained uncertain, for the additional
information they contributed about the level of genetic
exchange between putative taxa or between populations.
While paraphyletic species were accepted in some cases, in
other cases cryptic taxa were hypothesised to explain the
phylogenetic relationship between accessions.
Materials and methods
Species descriptions
Specimens from the CANB, PERTH, AD, AD-U, BRI, MEL, MJP,
NSW, SYD, BM, K and OXF herbaria were examined and measured,
and any information they retained about species’ morphology,
phenology, habit, ecology or distribution was used to construct species
descriptions. Unfortunately, not all the specimens requested on loan
from PERTH and none of the specimens requested from MJP were
actually received on loan, leaving descriptions of the rare taxa P. adunca,
P. brachyphylla, P. calycina subsp. proxena and P. daena incomplete.
Some of the collection localities of these taxa (acquired from a cursory
look at specimen labels while at PERTH or MJP) were nonetheless
used to construct distribution maps, although the specimens are not
cited here as having been viewed. Additional collections and notes
from living plants were obtained on a field trip made to south-west
Western Australia in October–November 2002. Sheet collections
made during the trip are housed at CANB and PERTH, with some
duplicates available at MEL and K. Additional inflorescences of
some collections were stored in a 70% ethanol solution and housed
at CANB. Inflorescences and flowers from 1–20 specimens per taxon
were dissected for accurate measurement and description. Up to five
specimens per taxon were used for the measurement of vegetative
characters, with additional specimens measured if their organs
seemed outside the size range of previously measured specimens.
A dichotomous key to the Pultenaea taxa found in WA was constructed
by hand, taking account of the phylogenetic groups found by Orthia et al.
(2005). Additional characters that were obviously consistent within
these phylogenetic groups were also noted. Distribution data from
herbarium specimens were used to create distribution maps in ArcMap
(© 1999–2001 ESRI).
A selection of specimens examined is listed at the end of each
taxon treatment, with priority given to those collections mentioned
in the text, those used for DNA work, Sands’ collections [used
in her cytological work (Sands 1966, 1975)] and collections from
underrepresented herbaria. A complete list, sorted by both taxon name
and by herbarium / collector name, is available from the website of
Australian Systematic Botany.
Collection localities for rare or threatened taxa, including type
localities, have been abbreviated or omitted to protect the taxa. No
localities, dates or collection numbers are available for Williamson’s
‘Pultenaea Reference Set’ specimens because the reference set was
acquired by taking pieces from others’ collections, without note being
taken of the collections from which they came.
Characters used in descriptions
Some morphological character states used in descriptions are defined by
de Kok and West (2002, 2003, 2004), e.g. levels of hairiness (glabrous,
sparsely hairy, densely hairy, velutinous), hair types (grooved, terete)
Australian Systematic Botany
151
and bract types (inflorescence-subtending, flower-subtending). Bracts
and bracteoles are considered to be made up of three parts, homologous
to two stipules and a leaf, which are free or fused to each other to varying
degrees. Bract ‘leaves’ (the leaf part of the leaf + stipule structure)
are leaf-like in texture or scarious, and are leaf-like in size to very
reduced. Where a single measurement is given for an organ (except
the stigma), it is the length unless otherwise specified. Measurements
given for stigmas are the diameter. Leaf measurements refer only to
the lamina, including any mucro or arista, and excluding the petiole.
When measurements are given for the standard petal, the full length
including the claw is given first, and the length of the claw is also given
separately. Measurements for components of the gynoecium (stipe if
present, ovary, style) are all given separately. Botanical districts follow
Beard (1980).
New calyx measurements were introduced because of the importance
of calyx shape and size in species identification. The first group of
measurements is denoted as ‘tube’, ‘U–U’, ‘U–L’ and ‘L–U’ (Fig. 1).
‘Tube’ represents the length of the tube from the base of the lower
lobes to the receptacle. ‘U–U’ represents the distance from the upper
lobe apex to the point where upper lobes fuse; ‘U–L’ the distance from
the upper lobe apex to the point where upper and lower lobes fuse;
‘L–U’ the distance from the lower lobe apex to the point where upper
and lower lobes fuse. Measurements were made in a straight line and
included lobe aristae. For species with greatly enlarged upper calyx
lobes, the additional measurement of ‘coherence’ was introduced: this
measurement represents the percentage of upper lobe length that is
coherent with the lower lobe in bud (Fig. 2).
DNA sequencing component
DNA sequences were obtained for the P. ericifolia / P. indira /
P. empetrifolia group of taxa, the P. quaerita / P. arida / P. calycina group
and for P. pauciflora. Sequences from the cpDNA trnL intron / trnL–F
intergenic spacer region were obtained in the first instance, and a
subset of accessions was sequenced for the chloroplast ndhF and
nuclear ITS regions as well, to ensure that the results were not
a peculiarity of trnL–F. Sequences of closely related species were
included, and the type of the genus, P. stipularis Sm., plus either
Fig. 1. Diagramatic representation of a Pultenaea calyx, illustrating
the measurements tube, U–U, U–L and L–U used in species descriptions.
Tube represents the distance from the receptacle to the base of the lower
lobes. U–U represents the distance in a straight line from the upper lobe
apex to the point where upper lobes fuse; U–L represents the distance
from the upper lobe apex to the point where upper and lower lobes fuse,
L–U represents the distance from the lower lobe apex to the point where
upper and lower lobes fuse.
152
L. A. Orthia et al.
Australian Systematic Botany
a
b
c
d
e
f
g
Fig. 2. Representation of Pultenaea calyces, illustrating the meaning of the new measurement, ‘coherence’. (a, b) Each calyx lobe is coherent
with the lobe adjacent to it when the flower is in bud. (c, d) Coherence can also be estimated after the flower opens because calyx lobes
have a prominent ‘ripped’ look where they were coherent in bud. (e, f ) Detail of ‘ripped’ area on upper lobe. (g) For taxa with aristate lobe
apices, the aristae are not coherent with the upper lobes, but remain free. All scale bars = 1 mm. Illustration by Tony Orchard. Vouchers:
a, c, e Pultenaea arida Archer 1211952 (CANB); b, d, f Pultenaea quaerita Hoyle 1396 (CANB); g Pultenaea elachista Wrigley s.n.
(CANB 038043).
Revision of Pultenaea: Western Australian species
P. daphnoides Wendl. or P. adunca, were included as outgroups (see
Table 1 for list of accessions used). Ingroup and outgroup species were
chosen according to relationships recovered by phylogenetic research
conducted in tandem with the taxonomic revision (Orthia et al. 2005),
and sequence alignments, indels, gaps, character weights were treated
as for that study. Maximum parsimony tree estimations were performed
in PAUP* version 4.0b10 (Swofford 2002); each analysis comprised
100 random addition sequence starts, saving 10 trees for each replicate.
The saved trees were then searched, with the Maxtrees option set to
20 000, and a strict consensus tree was constructed from the trees
generated by this search. Data sets were subjected to 1000 bootstrap
replicates (Felsenstein 1985), each with 10 full heuristic searches
starting from a random addition sequence and saving no more than
100 trees in each search.
Although DNA sequencing does not necessarily resolve infraspecific
questions, the use of finer-scale genetic or phenetic quantitative
methods to determine taxonomic status was beyond the scope of
the project.
Results and discussion
Over 1100 Pultenaea sheets were examined and used in
the taxonomic treatment. Two of the species to be included
in the study, P. adunca and P. vestita, plus the new taxa
P. wudjariensis and P. indira subsp. pudoides, were not
collected on the field trip, but all other taxa were. All but
four of the taxa collected were seen in full flower.
While most attempts to sequence plants for the trnL–F
region were successful, some taxa were unable to be
sequenced for the ndhF and in particular the ITS
regions despite repeated attempts. The only taxon for
which we were unable to obtain a trnL–F sequence was
P. indira subsp. pudoides, possibly because we were using
herbarium material. Phylograms from the DNA sequencing
component are presented in Fig. 3, with key statistics
in Table 2.
Paraphyletic species—cryptic taxa or maternal lineage
sorting?
With the exception of P. empetrifolia, all taxa for which
more than one population was included in the phylogenetic
study were found to be paraphyletic, including P. arida,
P. ericifolia, P. indira, P. calycina and P. quaerita. In the
cases of P. arida and P. ericifolia the paraphyly possibly
indicates the presence of cryptic taxa. Pultenaea arida may
contain two or more taxa that maintain their distinctiveness by
flowering at different times, although the evidence supporting
this hypothesis is weak and requires further investigation
(see P. arida treatment for detailed discussion).
Pultenaea ericifolia and allies presented a wealth
of difficulties for revision. The problems arose mainly
because of conflict between inferred relationships based
on morphology and those based on DNA sequences.
For example, sequence data provided evidence that
the morphologically unified P. ericifolia is paraphyletic
(Fig. 3b–d), falling into two distinct clades that include
other taxa, making it difficult to accept the unity of the
species. Genetic data strongly supported the presence of
Australian Systematic Botany
153
two cryptic taxa within P. ericifolia, corresponding to a
north–south geographic disjunction, but this too requires
further investigation. TrnL–F, ndhF and ITS trees also
showed some degree of conflict for close relatives of
P. ericifolia, primarily the placement of P. verruculosa and
P. strobilifera in relation to the two P. ericifolia clades
(Fig. 3b–d). As a result, the five taxa in this group—
P. ericifolia, P. verruculosa, P. strobilifera and two novel
taxa—are here described as part of the ‘P. ericifolia species
complex’. No changes have been formally proposed to ranks
or names in this group, pending more detailed study. The
problems presented by the group are discussed in detail in the
taxonomic treatment.
The north–south divide in the P. ericifolia situation seems
similar to that of the endemic WA species Lambertia orbifolia
C.A.Gardner (Proteaceae), in which two morphologically
indistinguishable disjunct populations were found to belong
to two long-diverged lineages that possibly represented
cryptic taxa (Byrne et al. 1999). Certainly, cryptic taxa
have been detected among many other organisms, including
other legumes, monocots, bryophytes, ferns, fungi and
invertebrates (Parker 1996; Condon 1997; Trewick 1998;
Hauk and Haufler 1999; Robinson and Hoffmann 2000; Shaw
2001; Horton 2002; Orthia et al. 2004), so it should not come
as a surprise to find them in Pultenaea as well.
The other paraphyletic taxa found in the study were
treated as single species. Although under a strict cladistic
approach to classification only a set of terminals forming a
monophyletic group can comprise a taxon (Donoghue 1985;
de Queiroz and Donoghue 1988), this approach does not
credit the possibility of peripatric speciation (Omland 1997),
whereby a geographically isolated population of a widespread
species undergoes a speciation event, ‘budding off’ from
the parent species. Paraphyletic species are inevitable if the
parent species is not considered to be ‘extinct’ as soon
as this ‘budding off’ has occurred (Nixon and Wheeler
1990; Baum 1992; Patton and Smith 1994; Crisp and
Chandler 1996; Omland et al. 2000). For example, this
may explain the placement of P. brachyphylla within the
P. indira clade: P. brachyphylla may have recently budded
off from the more widespread P. indira (distribution of
P. brachyphylla is peripheral to that of P. indira), forming
some barrier to genetic exchange that facilitated speciation
followed by morphological changes, while P. indira continued
its existence without interruption.
Numerous studies (e.g. Melnick et al. 1993; Brown
et al. 1994; Patton and Smith 1994; Funk et al. 1995;
Crisp and Chandler 1996; Omland 1997; Slade and
Moritz 1998; Friesen et al. 1999; Kelemen and Moritz
1999; Omland et al. 2000; Kimball et al. 2003) have
found paraphyletic species among organisms as diverse as
monocots, dicots, insects, amphibians, birds and mammals.
There are competing views in the scientific community about
their taxonomic validity, with debate centred on the conflict
154
Species and subspecies
Standard morphology, similar to type
Mt Ney population, unusual morphology
Toompup Rd, SE of Borden population
Hassell Hwy, Boxwood Hills population
West of Newdegate population
Tarin Rock population
North of Bindoon population
Perth Hills population
Whicher Range population
Yornup, near Manjimup population
Mount Barker population
East of Denmark population
West of Cranbrook population
North of Stirling Range population
Lucky Bay, Cape Le Grand population
South of Ravensthorpe population
Mt Burdett N of Esperance population
Newdegate population
Narrogin population
Quindanning population
Wandering population
Standard, large leaf morphology similar to type
Cape Arid population, unusual leaf morphology
Collector
Collector’s number
GenBank no. trnL–F
GenBank no. ndhF
GenBank no. ITS
Chappill
Chappill
Orthia
Orthia
Orthia
Orthia
Orthia
Gauba
Orthia
Orthia
Orthia
Orthia
Orthia
Crisp
Orthia
Orthia
Orthia
Orthia
Orthia
Orthia
Orthia
Orthia
Orthia
Orthia
Crisp
Orthia
Orthia
Orthia
Orthia
Orthia
Orthia
Chappill
J. Taylor
Orthia
Orthia
Orthia
de Kok
Orthia
Woolcock
6544
6272
71
56
51
63
57
22264
66
53
54
79
80
8451
88
39
41
46
48
44
52
75
59
70
9178
78
86
83
84
85
60
4279
1569
38
61
68
701
45
W2250
AY883190
AF518162
AY883192
AY883197
AY883384
AY883199
AY883201
AF113786
AY883205
AY883385
AY883206
AY883386
AY883387
AF113788
AY883207
AY883388
AY883389
AY883390
AY883391
AY883392
AY883393
AY883394
AY883215
AY883395
AY883396
AY883397
AY883398
AY883400
AY883226
AY883399
AY883230
AY883231
AY883401
AY883232
AY883235
AY883241
AY883242
AY883253
AY883261
AY883284
–
–
AY883290
–
–
–
–
–
–
AY883299
–
AY883402
–
AY883300
–
–
AY883403
–
–
AY883404
–
AY883308
AY883405
–
AY883406
AY883407
–
–
–
–
–
–
AY883325
–
–
AY883335
AY883345
–
AY883362
–
–
–
–
–
–
–
–
–
–
–
–
AF113770
AY883367
AY883408
–
–
–
AY883409
–
AY883410
AY883369
–
–
–
–
–
–
–
–
–
–
–
–
–
AY883378
–
–
L. A. Orthia et al.
P. adunca Turcz.
P. arida E.Pritz.
P. arida E.Pritz.
P. brachyphylla Turcz.
P. calycina (Turcz.) Benth. ssp. calycina
P. calycina ssp. proxena Orthia & Chappill
P. daena Orthia & Chappill
P. daphnoides Wendl.
P. elachista (F.Muell.) Crisp
P. empetrifolia Meisn.
P. empetrifolia Meisn.
P. empetrifolia Meisn.
P. empetrifolia Meisn.
P. ericifolia Benth. s.s.
P. ericifolia Benth. s.s.
P. ericifolia ‘southern’ (Orthia 39)
P. ericifolia ‘southern’ (Orthia 39)
P. ericifolia ‘southern’ (Orthia 39)
P. ericifolia ‘southern’ (Orthia 39)
P. strobilifera Meisn.
P. strobilifera Meisn.
P. strobilifera Meisn.
P. indira Orthia & Crisp ssp. indira
P. indira Orthia & Crisp ssp. indira
P. indira Orthia & Crisp ssp. indira
P. indira ssp. monstrosita Orthia
P. sp. Mt Lesueur (Beard 7827)
P. pauciflora M.B.Scott
P. pauciflora M.B.Scott
P. pauciflora M.B.Scott
P. purpurea (Turcz.) Crisp & Orthia
P. quaerita Orthia
P. quaerita Orthia
P. radiata H.B.Will.
P. rotundifolia (Turcz.) Benth.
P. spinulosa (Turcz.) Benth.
P. stipularis Sm.
P. verruculosa Turcz.
P. wudjariensis Orthia
Notes
Australian Systematic Botany
Table 1. Accessions used in DNA sequencing component of this study or referred to in text, plus GenBank accession numbers of sequences used
Dashes indicate that there was no sequence of the relevant gene for the taxon
Revision of Pultenaea: Western Australian species
Australian Systematic Botany
Fig. 3. Phylogenetic trees constructed in this study. Maximum-parsimony bootstrap phylograms are used to illustrate branch lengths.
Branch lengths are shown above the line and bootstrap values over 50 below the line in all trees. (a) trnL–F tree for the taxa with greatly
enlarged upper calyx lobes (P. quaerita group). (b) trnL–F tree for the taxa with prominent stipules not fused (P. ericifolia / P. indira
group). Pultenaea strobilifera, P. verruculosa and P. sp. Mt Lesueur are part of the P. ericifolia species complex. (c) ndhF tree for the
P. ericifolia / P. indira group. (d) ITS tree for a selection of taxa from the P. ericifolia species complex.
155
156
L. A. Orthia et al.
Australian Systematic Botany
Table 2. Phylogenetic tree key statistics
Statistic
No. of nucleotide characters
No. of recoded indel characters
No. of parsimony informative
characters
CI
RI
No. of most parsimonious trees
P. quaerita
group trnL–F
(a)
P. ericifolia / P. indira
group trnL–F
(b)
P. ericifolia / P. indira
group ndhF
(c)
P. ericifolia
species complex
ITS (d)
996
120
29
996
120
45
579
11
46
662
12
40
0.865
0.904
12
0.806
0.939
565
0.933
0.963
18
0.800
0.804
2
between biological and phylogenetic species concepts.
Some authors have maintained the conviction that species
cannot be paraphyletic, seeing reproductive compatibility
as a plesiomorphic character that obscures dichotomously
branching phylogenetic relationships (Donoghue 1985;
Baum 1992; Kimball et al. 2003). However, if populations
of a sexual species diverge for a time then meet up again
to interbreed, a reticulate phylogenetic branching pattern
will result. Under the phylogenetic species concept, such
populations could be prematurely split into separate species
if they were split before they re-established contact (Baum
1992; Hoelzer and Melnick 1994). Indeed, reproductive
compatibility is a trait that distinguishes the rank of
species from all higher taxonomic ranks, in both its
impact on phylogenetic patterns and its impact on the
lives of individual organisms, a fact that some workers
believe should be recognised in classification (Mayr 1963;
Lee 2003). For these reasons, paraphyletic taxa have been
accepted here.
Perhaps more interesting in the case of the
P. indira / P. brachyphylla clade is that despite a wide
range of morphological variation within the clade, a
geographic rather than morphological split was reproduced
in the phylogenetic analysis. The two morphologically
indistinguishable Ravensthorpe and Esperance populations
of P. indira subsp. indira were split up genetically,
with the Esperance population sitting as sister to the
rest of the clade, while the geographically proximal
but morphologically diverse Ravensthorpe, Newdegate,
P. indira subsp. monstrosita and P. brachyphylla populations
formed a monophyletic group in cpDNA trees (Fig. 3).
Without an nrDNA tree for comparison or a more detailed
genetic study, it is difficult to be certain of what is occurring
within P. indira subsp. indira, but the observed pattern
could be explained by the geographic restriction of cpDNA
haplotypes: Esperance and Ravensthorpe populations
could be exchanging pollen but not seed. This may
also explain the paraphyly observed in P. calycina and
P. quaerita. Such maternal lineage sorting has been observed
in other organisms (e.g. Joseph et al. 1995; Freeman et al.
2001; Eggert et al. 2002), and Pultenaea pollen is probably
more mobile than its large seeds. However, it is
noted that while chloroplasts are usually maternally
inherited in angiosperms, it has not been demonstrated
for Pultenaea, and some distantly related legumes
inherit chloroplasts biparentally or paternally, as do
species from other families (Harris and Ingram 1991;
McKinnon et al. 2001). More detailed genetic studies
are required for some or all of these Pultenaea taxa
to consider more effectively the history of gene flow
between populations.
Concordance with phylogenetic study
The three main groups of WA Pultenaea recovered in
the molecular phylogeny (Orthia et al. 2005) have many
morphological correlates, which are recognised in the
key to species presented here: the species with greatly
enlarged upper calyx lobes (couplets 4–11) form one
lineage (Clade D), the species with hairs inside their pods
(couplets 13–17) form another (Clade F) and the species
with prominent stipules that are not fused behind the petiole
(couplets 23–26) form a third lineage (Clade E) (clade names
from Orthia et al. 2005). Pultenaea brachytropis Benth.
forms a monotypic lineage (couplet 19). Table 3 details
other characters that can be used to differentiate between
these four groups.
Outstanding issues
This paper completes the species level revision of
Pultenaea with one major exception: further work is
required to determine the relationships, history and
delimitation of taxa in the difficult P. ericifolia group.
The status of P. strobilifera in particular has been
in doubt for years as various authors (including one
of us) have informally designated it a subspecies of
P. ericifolia. Some of the approaches that could be taken
to researching this group are detailed under the P. ericifolia
complex treatment.
Further research is also needed into the possibility
of cryptic taxa within P. arida. This species appears
paraphyletic in molecular trees, has a variable
morphology and morphotypes can flower at different
Cladistic group
(see Orthia et al. 2005)
Habit
Leaf margins
Stipules
Inflorescence type
Inflorescence
bracts
Flower
bract texture
Upper
calyx lobes
Clade D
(P. quaerita group)
prostrate to 1.5 m
variable
not fused, material
present behind petiole
absent
herbaceous, leaf like
greatly enlarged,
elliptic–obovate
Clade E
(P. ericifolia /
P. indira group)
Clade F
(P. skinneri /
P. reticulata group)
P. brachytropis
to 1 m
involute
not fused, no material
behind petiole
weakly terminal or
axillary / solitary
indeterminate
terminal indeterminate
usually scarious
narrow triangular
1–3(–5) m
flat or recurved
fused
terminal determinate
scarious
scarious, rarely
herbaceous and
leaf like
scarious
usually 1–1.5 m
recurved
fused
terminal determinate
absent
scarious
rounded, fused
for length
Cladistic group
(see Orthia et al. 2005)
Standard
petal shape
Size of claw
on standard
Wing arrangement
Wing and
keel arrangement
Keel size
relative to wing
Ovary shape
Inside of
ovaries and pods
Clade D
(P. quaerita group)
Clade E
(P. ericifolia /
P. indira group)
Clade F
(P. skinneri /
P. reticulata group)
P. brachytropis
reniform or round
short
variable
variable
± equal
laterally flattened
glabrous
reniform or round
long
often coherent and / or
overlapping at apex
wings conceal keel
much smaller
or ± equal
elongate ovoid
glabrous
round
short
not coherent except
sometimes at apex
keel protrudes
through wings
larger
ovoid
hairy
strongly reniform
short
held apart, not coherent
wings conceal keel
much smaller
ovoid
glabrous
Revision of Pultenaea: Western Australian species
Table 3. Consistent differences in morphological characters for WA Pultenaea clades found in Orthia et al. (2005)
narrow triangular
Australian Systematic Botany
157
158
L. A. Orthia et al.
Australian Systematic Botany
times. This is discussed further under the P. arida
treatment.
Another significant issue is the changed conservation
status of several taxa. Good work has been done in recent
years to locate and conserve populations of WA Pultenaea
species previously identified as rare or threatened, such
as P. pauciflora, P. skinneri and P. pinifolia. This revision
highlights other rare taxa worthy of conservation attention,
such as P. wudjariensis, which is known only from one
collection.
Taxonomic treatment
1. Pultenaea adunca Turcz., Bull. Soc. Nat. Mosc. 26:
279–280 (1853). Typus: ‘Swan River’, 1849, Drummond V
66 (holo-, KW (photo seen); iso-, K (3 sheets)!, BM!, OXF!,
MEL2055281!)
Key to the species of Pultenaea occurring in Western Australia
1. Stipules absent
2. No beard on style; leaves involute; bracteoles scarious.......................................................................................................................................1. P. adunca
2: Beard on style; leaves revolute; bracteoles leaf-like...........................................................................................................................................4. P. barbata
1: Stipules present
3. Two upper calyx lobes much larger than three lower lobes, with the depth of sinus between two upper lobes ≥ depth of sinus between upper and lower
lobes; upper lobes ± obliquely elliptic and widest near middle
4. Leaf margins strongly recurved (particularly clear in young leaves)
5. Average leaf length : width ratio >4 : 1; leaves spiralled.........................................................................................................................17. P. purpurea
5: Average leaf length : width ratio <4 : 1; leaves decussate or whorled
6. Pedicels longer than leaves; branchlets with straight appressed hairs............................................................................................21. P. rotundifolia
6: Pedicels shorter than leaves; branchlets with curly hairs......................................................................................................................9. P. elachista
4: Leaf margins flat or incurved
7. Calyx lobes and leaves aristate...............................................................................................................................................................23. P. spinulosa
7: Calyx lobes and leaves acute or mucronate but not aristate
8. In bud upper calyx lobes are coherent with lower lobes for ≥75% of their length; upper calyx lobe apex acute
9. Prostrate shrub; leaves densely hairy, leaf hairs erect............................................................................................................................8. P. daena
9: Erect shrub; leaves sparsely hairy; leaf hairs usually appressed.............................................................................................................2. P. arida
8: In bud upper calyx lobes are coherent with lower lobes for ≤ 67% of their length; upper calyx lobe apex obtuse
10. Leaves usually whorled; leaf length : width ratio ≥ 4 : 1..................................................................................................................7. P. calycina
10: Leaves decussate; leaf length : width ratio ≤ 2 : 1
11. Leaf and calyx hairs appressed, grooved; leaves elliptic or obovate; keel petals yellow with red spots at apex...............26. P. wudjariensis
11: Leaf and calyx hairs often erect, never grooved; leaves usually obcordate; keel petals red.....................................................18. P. quaerita
3: Two upper calyx lobes ± equal in size to three lower lobes, or if upper lobes larger than lower lobes then upper lobes are fused for almost entire length
and sinus between them < sinus between upper and lower lobes; upper lobes ± triangular or narrowly triangular and widest at base
12. Ovaries and pods have hairs on the inside
13. Leaves without prominent secondary venation
14. Leaf margins flat; keel pale green................................................................................................................................................15. P. pauciflora
14: Leaf margins recurved; keel red to brown
15. Leaves >10 mm long; held ascending to inclined....................................................................................................................16. P. pinifolia
15: Leaves <10 mm long; held divergent to reflexed.....................................................................................................................22. P. skinneri
13: Leaves with prominent reticulate or pinnate venation
16. Leaf venation pinnate; keel pale green or yellow....................................................................................................................3. P. aspalathoides
16: Leaf venation reticulate; keel red to brown
17. Stipules partially enclose stem; leaf apex blunt.......................................................................................................................14. P. ochreata
17: Stipules do not enclose stem; leaf apex acute to aristate.......................................................................................................20. P. reticulata
12: Ovaries and pods glabrous inside
18. Stipules fused to at least one third of their length behind the petiole
19. Leaf margins recurved...............................................................................................................................................................6. P. brachytropis
19: Leaf margins incurved
20. Bracteoles tridentate, with free stipules......................................................................................................................................25. P. vestita
20: Bracteoles ovate, without free stipules
21. Leaves with a sharp point................................................................................................................................................13. P. juniperina
21: Leaves without a sharp point............................................................................................................................................24. P. tenuifolia
18: Stipules not fused behind the petiole
22. Flower subtending bracts leaf-like; stipules minute or cryptic..........................................................................................................1. P. adunca
22: Flower subtending bracts scarious with reduced bract ‘leaf’; stipules prominent
23. All leaves < 4 mm measured in a straight line base to apex (i.e., if recurved, do not ‘straighten’ before measuring)......10. P. empetrifolia
23: Most leaves > 4 mm in length
24. Stipules glabrous
25. Bracteoles broad, surrounding calyx, apex rounded...................................................................................................... 19. P. radiata
25: Bracteoles narrow, not surrounding calyx, apex acuminate.........................................................................11. P. ericifolia complex
24: Stipule margins ciliate, body of stipule hairy or glabrous
26. Leaves mucronate; body of bracts and stipules densely hairy to velutinous, hairs on bracts, stipules and stem white and tightly
curled...................................................................................................................................................................... 5. P. brachyphylla
26: Leaves aristate; body of bracts and stipules glabrous, or if hairy, then hairs are ± straight............................................12. P. indira
Revision of Pultenaea: Western Australian species
Illustrations
Grieve, How to Know Western Australian Wildflowers 2: 421
(1998).
Prostrate or erect shrub, height to 1 m. Lower
stems red beneath metallic-looking bark, glabrous, bark
flaking off; upper stems sparsely to densely hairy just
below inflorescences, hairs appressed. Leaves dark green,
4.5–8.5 × 0.7–0.8 mm, length : width ratio 10 : 1, linear or
narrowly oblong, spiralled, appressed to inclined, slightly
recurved, no prominent venation, lamina smooth or slightly
verruculose, older leaves glabrous, younger leaves densely
hairy, hairs appressed; apex mucronate, base truncate,
margins involute; petioles 0.8–1.0 mm, not decurrent.
Stipules absent or red-brown, 1.0 × 0.1–0.2 mm, not fused
behind the petiole, narrowly triangular, straight, sparsely
to densely hairy, breaking off and fraying with age,
apex acute. Inflorescences axillary, towards branch apices,
indeterminate, 4–10 subsessile flowers, pedicels 0.1 mm;
inflorescence-subtending bracts absent, flower-subtending
bracts leaf-like. Bracteoles greenish brown, 2.4 × 0.2 mm,
linear, inserted at base of calyx, scarious, densely hairy, apex
acuminate. Two upper calyx lobes ± similar in size to lower
three lobes, tube 2 mm, U–U 1.7 mm, U–L 2.8 mm, L–U
2.8 mm, densely hairy, hairs appressed to ascending; upper
lobes 1.0 mm wide, acute, sinuously triangular, curving away
from each other at apex; lower lobes 0.5 mm wide, narrowly
triangular, apex acuminate. Flowers reddish or yellow
suffused purple; standard yellow at margins, orange / red at
centre, with dark radiating lines, 10 × 6.5 mm, claw 5 mm;
wings orange / yellow, 9.5 × 2 mm; keel red, 6 × 1.8 mm.
Anthers 0.4 mm, dorsifixed, filaments 4.5–5.5 mm. Ovary
1.5 mm, cross-section elliptic, velutinous, stipe minuscule
or absent; style 2.5 mm, hooked, glabrous, stigma diameter
slightly larger than style diameter, 0.2 mm. Pods and seeds
not seen.
Phenology
Flowering September–December.
Habitat and ecology
Heath or mallee on sand, silty clay, sandy loam or gravel, on
rises or roadsides. Can be abundant at sites.
Distribution
Australian Systematic Botany
159
Flora Codes. This species has been collected fewer than ten
times in 150 years, and an attempt to locate it again in October
2002 was unsuccessful. This may mean that it is extremely
rare and possibly threatened, although some collectors state
it is abundant at collection sites (J. Chappill, pers. comm.). A
search for existing populations should take place as a matter
of urgency.
Chromosome number
Sands (1975) lists the chromosome number for Pultenaea
adunca as 2n = 14, but the voucher specimen from which
the count was made (Sands 638.17.3) does not belong to
P. adunca, but to P. indira. Keighery (1984) lists it as n = 7,
but his voucher belongs to P. barbata.
Discussion
Morphologically, P. adunca does not closely resemble any
other Pultenaea species, with its small stipules and clustered
inflorescences that are not subtended by inflorescence bracts.
In a molecular study by Orthia et al. (2005), P. adunca fell
outside all Pultenaea s.s. clades, and clustered instead with
Almaleea, which also has minute stipules (Crisp and Weston
1991). Morphological traits it shares with Almaleea
include the growth form where the next season’s growth
appears to come out of the top of the inflorescence of the
previous year, and the shape of the standard (J. Chappill,
pers. comm.)
Notes
Only dried flowers in poor condition were examined for
morphological description in this revision; fresh flowers
might be coloured differently and petals might be slightly
larger.
The species is very poorly known. Table 4 lists the
characters distinguishing P. adunca from taxa commonly
confused with it.
Specimens examined
Gnowangerup, N of Stirling Range, Blackall & Gardner s.n., x.1928
(PERTH); Jerramungup on Albany–Ravensthorpe Road, Stirling Range,
Blackall & Gardner s.n., x.1929 (PERTH); Tributary of Oldfield River,
Crisp 4944, 8.i.1979 (PERTH); W.A., Mueller s.n., n.d. (MEL); ENE
of Esperance, Turner 5551, 12.ix.1974 (PERTH); Pultenaea Reference
Set, Williamson s.n., n.d. (MEL2055281).
Western Australia, botanical districts Roe and Eyre. Found
half way between Ravensthorpe and Esperance, also ∼100 km
ENE of Esperance. Also known from generalised location
descriptions of ‘Gnowangerup’ and ‘Jerramungup’ (Fig. 4a).
2. Pultenaea arida E. Pritz., in Diels, F.L.E. & Pritzel,
E. Bot. Jahrb. Syst. Pflanzengesch. Pflanzengeogr. 35: 258
(1904). Typus: ‘in distr. Coolgardie meridionalis pr. Gilmores
in eucalyptetis lutosis flor. m. Nov. (D. 5275)’ (holo-, B
(probably destroyed); iso-, MEL2057349!)
Conservation
Illustrations
Here recommended for conservation risk code 3K (Briggs
and Leigh 1996) and for Priority Three under CALM Priority
Grieve, How to Know Western Australian Wildflowers 2: 423
(1998).
160
Australian Systematic Botany
L. A. Orthia et al.
Fig. 4. Distribution maps of (a) P. adunca, (b) P. barbata, (c) P. brachytropis, (d) P. juniperina in WA, (e) P. tenuifolia in WA and
( f ) P. vestita in WA.
Multistemmed, much-branched, spreading low shrub,
open to compact, 0.1–0.4(–0.8) × 0.3–0.6(–1) m, sometimes
growing from woody rhizomatous root stock. Bark dark
coloured, thick, often plated, highly textured, fissured and
rough with occasional hairs (rarely soft, smooth and
densely hairy). Leaves grey-green to reddish-purple, vein
and margins sometimes darker, lamina 1.2–5.6(–8.0) ×
0.9–3.1(–4.0) mm, length : width ratio 1 : 1 to 3 : 1, elliptic
to obovate, decussate, sometimes crowded, appressed to
reflexed, midvein prominent on abaxial surface, lamina
Taxon
Leaf margin
curvature
Leaf transverse
posture
Stipules
Bract texture
Bracteoles
Upper calyx lobes
shape
Ovary
Style
indumentum
P. adunca
involute
recurved at tip
recurved at tip
scarious, inserted
at base of calyx
leaf like, herbaceous,
inserted on pedicel
sessile
bearded
P. ericifolia
involute
straight to recurved
prominent
scarious
scarious, inserted at
base of calyx
sessile
glabrous
P. verruculosa
involute
straight to incurved
prominent
scarious
scarious, inserted at
base of calyx
sessile
glabrous
Dillwynia
acerosa
Dillwynia
divaricata
involute
gently slightly
recurved
straight to slightly
incurved
absent
herbaceous,
leaf like
herbaceous,
leaf like
scarious, inserted
on pedicel
scarious, inserted at
base of calyx
apex curved and
pointed outward
narrow triangular,
united into lip
with deep sinus
narrow triangular,
united into lip
with deep sinus
narrow triangular,
united into lip
with deep sinus
deltate, united into
short lip
oblique triangular,
united into lip
with short sinus
glabrous
revolute
herbaceous,
leaf like
herbaceous,
leaf like
sessile
P. barbata
minute, may
seem absent
absent
on long stipe
glabrous
on short stipe
glabrous
involute
absent
Revision of Pultenaea: Western Australian species
Table 4. Characters used to distinguish P. adunca and P. barbata from commonly misidentified species
Table 5. Characters used to distinguish commonly misidentified species with enlarged upper calyx lobes (P. quaerita group)
Leaf shape
P. arida
elliptic to obovate
P. calycina
P. daena
P. quaerita
oblong-linear or
clavate to obovate
elliptic to obovate
obovate
P. wudjariensis
elliptic to obovate
Leaf length :
width ratio
1–3 : 1
4 –11 : 1
2:1
1–2 : 1
2:1
Upper calyx
lobe shape
Upper calyx
lobe apex
% upper lobe
coherent to
lower lobe
Leaf and calyx
hair type
Leaf and calyx
hair orientation
oblique obovate
widely acute
85–95
grooved or round
obtuse
25–50
round
elliptic
elliptic to round
acute
obtuse
75–80
40–67
round
round
elliptic to obovate
obtuse
50–67
grooved
appressed to ascending,
acroscopic
erect, not acroscopic
(rarely appressed)
erect, not acroscopic
erect, not acroscopic
(rarely appressed)
appressed, acroscopic
elliptic to obovate
Bracteole
length
(mm)
1.1–1.9
1.3–3.9
2.5–3.5
1.0–2.2
1.5–2.6
Australian Systematic Botany
Taxon
161
162
L. A. Orthia et al.
Australian Systematic Botany
surface smooth, glabrous or sparsely hairy; hairs grooved,
appressed (rarely ascending), white, thick and denser on
younger leaves to blackened and thin on older leaves;
apex obtuse to rounded or slightly mucronate (occasionally
obcordate), base obtuse to rounded, margins flat to incurved;
petioles 0.4–0.6 mm, not decurrent. Stipules red-brown,
0.5–1.1(–2.7) × 0.5 mm, some stipule tissue remnant behind
petioles but not fused, ovate or triangular (or narrowly
triangular), straight or recurved, glabrous, apex acute;
when older sometimes black, rigid, thickish, frequently
broken and shapeless. Inflorescences axillary, in upper axils,
indeterminate; flowers solitary, pedicels to 2 mm; flower
bracts leaf-like. Bracteoles dark red, 1.1–1.9 × 0.6 mm,
lanceolate, inserted at base of calyx, glabrous, scarious,
apex acuminate. Two upper lobes of calyx much larger
than three lower lobes, tube 1–1.5 mm, U–U 4.0–4.6 mm,
U–L 3.7–4.2 mm, L–U 2.5–3.5 mm, all five lobes coherent for
85–95% of length, glabrous to densely hairy, hairs grooved,
appressed (rarely ascending); upper lobes 1.5–2.2 mm
wide, obliquely obovate, apex widely acute; lower lobes
0.5–0.8 mm wide, narrow triangular, apex acuminate. All
petals vary in colour from orange-yellow through pinkish
to entirely red; standard often has darker smudge near
apex, may have red edges, red on parts of back side and / or
red outline of yellow eye made of irregularly radial lines,
5.0–5.5 × 4.7–5.5 mm, ± round, erect or recurved, claw
1 mm, apex minutely notched; wings sometimes tipped
orange or red, 5.0–5.7 × 1.8–2.0 mm, held over keel,
coherent for length; keel 4.5–5.5 × 2.0–2.5 mm, ± same size
as wings. Anthers 0.4–0.6 mm, dorsifixed, filaments
3.5–4.5 mm. Ovary 2.0–3.5 mm, laterally flattened,
densely hairy, stipe 0.5 mm; style 2.5–3 mm, straight
to curved, glabrous; stigma diameter equal to style
diameter, 0.1 mm. Pod 3 mm, laterally flattened, exterior
sparsely hairy, hairs grooved or erect and terete; interior
glabrous. Seed black with mottled brown markings,
2.1 mm, arillate.
Phenology
Flowering October–December; fruiting December.
Habitat and ecology
Flat plains or gentle slopes, and granite outcrops.
Shrubland, mallee, low woodland and woodland. Includes
remnant patches, roadside verges and land previously
cleared for agriculture. Varied soils—clay, sandy clay,
loam or sand over clay, granite or limestone, can
contain gypsum or gravel. Rare to common, can be
locally abundant.
Distribution
Western Australia, botanical districts Roe and Coolgardie,
with rare collections in Eyre. Extends at least a few kilometres
north of Norseman. Usually east of the 121st meridian, but
occasionally found west of it, including one collection at
South Ironcap, north of Lake King. Generally restricted to
inland, semi-arid zones (Fig. 9a).
Leaf anatomy
Pultenaea arida has stomata, idioblasts (associated with
vasculature) and palisade mesophyll on both sides of the
leaf, and no spongy mesophyll (PG Turner and RPJ de Kok,
unpubl. data).
Typification
Both E. Pritzel, who first described Pultenaea arida, and
F.L.E. Diels, who collected the type specimen, worked from
the Berlin herbarium, most of which was destroyed during
the Second World War. The specimen that the original
description was based on is therefore presumed to have
resided at B, and so probably no longer exists. However, the
Melbourne-based botanist H.B. Williamson, who completed
a revision of Pultenaea during the 1920s, is known to have
taken fragments from type specimens, which he stuck to
his ‘Pultenaea Reference Set’ cards, now housed in MEL.
His Pultenaea arida reference card has such a fragment.
In Williamson’s revision of the species (1922), he states,
‘I have examined a piece of the type which the authorities
of the Berlin Museum of Botany kindly sent along. . .’.
Williamson’s comments about the piece of the type are
concordant with the fragment stuck to his card, and he
makes no mention of having seen any other specimens, so
presumably his fragment is an isotype.
Discussion
This species is variable in leaf shape and size, bracteole
and stipule length and hairiness. The differences do not
follow geographic clines or other clear patterns, and two
forms can be found within a few kilometres of each other.
It might be an interesting project to determine the factors
that govern morphological variation in the species. None
of the collections that have rare character states form
a consistent morphological group, so no infraspecific
divisions are proposed here. It should be noted though that
during a field trip in November 2002, the ‘common’ form
of P. arida, which generally has bracteoles shorter than
1.4 mm, stipules <1.1 mm and a sparsely hairy calyx, was
located three times (including Orthia 77), and in all three
cases was not in flower. On the same trip, two variant forms
of the species, one with longer bracteoles and stipules and
ascending hairs (Orthia 71), and another with completely
glabrous calyces and leaves (Orthia 76), were both in full
flower. It is possible that the different forms flower at different
times within a given season, even though on average their
flowering times overlap. DNA sequence-based phylogenetic
analysis in this study grouped two of the different forms
together, but in a polytomy with P. daena and P. elachista.
This result lends strength to the possibility that the P. arida
Revision of Pultenaea: Western Australian species
populations are not exchanging genetic material, and it
may indicate the presence of cryptic taxa. Pending further
research, all forms are grouped here as P. arida.
Notes
The variation in morphology complicates the diagnosis of the
species, particularly the factors differentiating P. arida from
P. wudjariensis and P. daena, two new species proposed in
this manuscript. Table 5 lists the characters distinguishing
P. arida from morphologically similar species.
Pultenaea arida is most closely related to P. daena and
P. elachista. It differs from the former in having an erect
habit and sparsely hairy leaves, and from the latter in its flat
to incurved leaf margins.
Selected specimens (51 examined)
20–30 km NW of Mt Willgonarinya, Archer 1011903, 10.xi.1990
(CANB, PERTH); 31.5 km NNE of Mt Heywood, Archer 24119013,
24.xi.1990 (MEL); 34 km W of Splinters Rock, Archer 1211952,
12.xi.1995 (AD, CANB, PERTH); 3 km NW of Mt Ney, Archer
1211953, 12.xi.1995 (MEL, NSW, PERTH); 1 km NW of
Peak Charles, Chandler 785, 9.xi.1998 (CANB); Brockway
Timber Reserve, Craig 2455, 18.xi.1992 (PERTH); 32 km ESE
Jyndabinbin Rocks, Keighery 12642, 11.xii.1990 (CANB, 2
dups, PERTH); 41 km West of Boingaring Rocks, Keighery
12667, 4.xii.1990 (CANB, PERTH); 5 km E of Salmon Gums
East Road, Macfarlane 2138, 6.xi.1992 (CANB, PERTH);
Near Mt Ney Rd, Orthia 71, 30.x.2002 (CANB, K, MEL, PERTH);
∼11.1 km along track to Theatre Rock, Orthia 76, 1.xi.2002
(CANB, K, MEL, PERTH); 25.2 km along Norseman–Hyden Rd,
Orthia 77, 1.xi.2002 (CANB, PERTH); Approximately 95 km
E of Norseman, Whibley 3005, 1.x.1969 (AD); Approximately
85 km WNW of Billadonia, Whibley 4528, 28.x.1974 (AD);
82 ml from Esperance, Wrigley WA / 68 5823, 12.xi.1968
(CANB, MEL).
3. Pultenaea aspalathoides Meisn., in Lehmann, J.G.C.,
Pl. Preiss. 1: 73 (1844). Typus: ‘In solo turfoso-subarenoso
sylvae et planitiei prope oppidum Albany (Plantagenet)
m. Oct. et Nov. 1840’, Herb Preiss. No. 838 (lecto-,
LD (photo seen); isolecto-, GOET, K (2 sheets)!,
MO, NY, S, W (2 sheets)) et 1195 (syn-, LD, NY),
here designated
Illustrations
Grieve, How to Know Western Australian Wildflowers 2: 419
(1998).
Erect, compact shrub, little branched, 0.3–1.7 ×
0.4–0.6 m. Bark yellow or red, smooth, lower stems
glabrous, upper stems densely hairy to velutinous, hairs
appressed to stem and grooved or terete. Leaves mustardy
green, 7–27 × 0.8–3.0 mm, length : width ratio 10 : 1,
narrowly elliptic, spiralled, frequently clumped at ends
of short axillary stems, appressed to inclined, slightly
incurved, prominent pinnately netted venation, smooth when
fresh, ridged and rigid when dried, old leaves glabrous,
Australian Systematic Botany
163
young leaves sparsely hairy with erect flat or terete hairs
to velutinous with appressed grooved hairs; apex acute,
aristate, base acute, margins incurved; petioles absent
or to 0.4 mm, decurrent. Stipules pale to mid brown
when young, 2–3(–5.5) × 0.5(–1.2) mm, fused behind the
petiole, obliquely ovate to narrowly triangular, appressed
to stem or recurved, glabrous to sparsely hairy with medial
hairs, may become transparent with thin brown keel with
age, apex acuminate if intact. Inflorescences terminal, in
axillary stems, determinate, with up to 7 sessile flowers
flowering non-synchronously; inflorescence- and flowersubtending bracts brown, 4 × 1.5 mm, leaf much reduced,
densely hairy, scarious. Bracteoles pale brown, 2.7–3.1 ×
1.0–1.2 mm, elliptic, inserted at base of calyx, with medial
and marginal hairs, scarious, apex rounded. Calyx lobes ±
equal in size, tube 2.7 mm, U–U 1.8 mm, U–L 2.2 mm,
L–U 1.7 mm, densely hairy with appressed, grooved
hairs, lobes 2.0 mm wide, deltate to triangular, apices
acute. Standard yellow, occasionally with red markings
around central yellow eye, greenish yellow on back side,
7.0–7.7 × 8.3–9.0 mm,
round,
strongly
incurved,
claw 1.7 mm, apex minutely notched; wings yellow,
7.0–7.7 × 1.2–1.6 mm, not touching each other; keel pale
green or yellow with pink tinge, 7.5–8.0 × 2.3–3.0 mm,
broader than wings, protruding through wings. Anthers
0.5 mm, dorsifixed, filaments 7.5 mm. Ovary 1.4 mm,
cross-section elliptic, velutinous, stipe 0.2 mm; style
6.7 mm, straight to curved, glabrous; stigma diameter
equal to style diameter, 0.1 mm. Pods 4.5 × 3.5 mm,
ovate; sparsely to densely hairy outside, hairs appressed
and grooved; sparsely hairy inside, hairs erect. Seeds
not seen.
Phenology
Flowering September–December.
December.
Fruiting
November–
Habitat and ecology
Flat areas or low slopes, often swampy areas and drainage
lines. Open to dense woodland, shrubland, heath or sedgeland.
Sandy clay, silty sand, peaty sand or peat, usually wet or damp.
Usually rare, sometimes frequent.
Distribution
Western Australia, botanical district Darling, subdistricts
Menzies and Warren. Generally, the area bounded by the
Stirling Ranges, Walpole and Mermaid Point, with one
collection near Cordering in the north-west of the range
(Fig. 8a).
Leaf anatomy
Pultenaea aspalathoides has stomata only on the upper leaf
surface, idioblasts associated with vasculature on both leaf
164
L. A. Orthia et al.
Australian Systematic Botany
surfaces and no palisade mesophyll (PG Turner and RPJ de
Kok, unpubl. data).
Typification
The collection of Preiss specimens at LD are the standard type
specimen set for species described in Plantae Preissianae
P. aspalathoides protologue, Preiss 838 and Preiss 1195,
are represented there. The Preiss 838 specimen has been
chosen as the lectotype of P. aspalathoides because it has
many more duplicates held at other herbaria than does
Preiss 1195.
Discussion
Examination of the species suggested that the morphology
was consistent and that there were no outstanding taxonomic
issues. However, one collection (Jackson s.n.) possessed
some characters setting it apart from most of the other
collections: leaves as large as 27 mm long (the others
rarely reach 20 mm), stipules that were consistently darker
(mid-brown, not becoming transparent with age) and larger
(3–5.5 × 0.8–1.2 mm, rather than the usual 2–3 × 0.5 mm),
young leaves that were densely hairy to velutinous with
appressed grooved hairs rather than sparsely hairy with erect
hairs, and a growth form less clumped than the form most
often collected. These specimens were collected from Bow
River, in the south-western end of the range. It is possible
that they belong to a separate infraspecific taxon from the
rest of the collections. It is possible that some specimens at
PERTH match the Jackson s.n. collections morphologically
but went unnoticed during our stay in Perth for this study.
Therefore, all specimens have been treated here as a
single taxon.
Notes
Leaf venation is the primary character distinguishing
P. aspalathoides from its closest relatives, P. reticulata
and P. ochreata: they both have prominent reticulate venation
and P. aspalathoides has prominent pinnate venation.
Selected specimens (40 examined)
Ficifolia road, Annels 827, 13.ix.1989 (PERTH); 20 km NW of
Denmark, Annels 1972, 18.xi.1991 (PERTH); 13.3 miles from Young’s
Siding, Canning 6276, 12.x.1968 (CANB); 30 m S of Redmond–
Hay River Road, Casson & Kershaw W 175.8, 11.iii.1997 (PERTH);
42 km NE of Albany, Cranfield 1137, 8.xi.1978 (PERTH); Gibbs
Road, Crowley DKN 323, 15.xi.1993 (PERTH); 7 km SSW of
Redmond, Davis 267, 31.x.1995 (PERTH); 50 m S of Ficifolia Road,
Ellery & Godden W 5.15, 11.ii.1997 (PERTH); Albany, Hamilton
810, 18.xii.1902 (NSW); Bow River, Jackson s.n., end.xii.1912
(CANB, NSW, PERTH); 26 km NE of Bakers Junction, Lepschi
& Fuhrer 3695, 26.x.1997 (CANB, PERTH); 7.2 ml NE along
Hassel Highway from Pfeiffer Road, Macfarlane 2097, 29.x.1992
(CANB, PERTH); On Hassell Highway, Orthia 50, 24.x.2002
(PERTH); Walpole–Nornalup National Park, Wheeler 3218, 21.ix.1992
(PERTH); 16 km E of Denmark, Woolcock W2247, 24.ix.1985
(CANB, MEL).
4. Pultenaea barbata C.R.P. Andrews, Journ. W. Austral. Nat.
Hist. Soc. 1: 38–39 (1904). Typus: ‘I found this species in
flower near the Phillips River in Oct. 1903.’ (holo-, PERTH
(photo seen))
Illustrations
Grieve, How to Know Western Australian Wildflowers 2: 419
(1998).
Open, multistemmed, erect or low spreading shrub,
usually sparse but sometimes moderately dense, 0.1–0.4 ×
0.4–0.6 m. Bark brown to orange, fissured on main
stems, lower stems glabrous, upper branchlets velutinous,
hairs appressed or occasionally erect. Leaves adaxially
pale- to light-green, abaxially pale tan, (4.0–)7.0–15.0 ×
1.0–1.2 mm, length : width ratio 7–10 : 1, linear, scattered,
appressed to descending but usually inclined, venation not
prominent, lamina smooth, glabrous, young leaves sparsely
hairy to velutinous, hairs appressed or erect and terete;
apex acute to obtuse, may have small mucro with slight
recurvature, base truncate, margins closely recurved; petiole
0.7–1.3 mm, decurrent. Stipules absent. Inflorescences
terminal, indeterminate, 2–7 flowers per umbel, pedicels
1–3 mm, no inflorescence-subtending bracts, flowersubtending bracts leaf-like. Bracteoles greenish,
4.5 × 0.6 mm, linear, inserted on pedicel 0.5–1.0 mm
below calyx, densely hairy with erect hairs, leaf-like, apex
acuminate. Two upper calyx lobes united into lip, slightly
larger than three lower lobes, tube 1.8 mm at lower lobes,
U–U 2.5 mm, U–L 4.1 mm, L–U 3.9 mm, densely hairy,
hairs erect; upper lobes 1.2 mm wide, obliquely sinuously
triangular, apex acuminate; lower lobes 1.1 mm wide,
narrowly triangular, apex acuminate. Standard yellow with
red margin and red markings flanking central ‘eye’ region,
which may also be red or brown, dark red on back side
with dark veins, 7.0 × 6.0 mm, ovate, recurved slightly, claw
1.2 mm, apex rounded; wings dark red with yellow inner
margin and tips, 6.5 × 1.5 mm, held horizontally over keel,
near coherent for length; keel red to yellow to green, usually
differently coloured patches at base and apex, 6.7 × 2.2 mm,
slightly larger than wings. Stamens not conspicuously joined
to petals; anthers 0.8 mm, basifixed, filaments 5.5 mm.
Ovary 1.6 mm, cross-section elliptic, velutinous, sessile;
style 6.5 mm, tapering from 0.8 mm wide at base to 0.1 mm
at tip, bent through 90 degrees, bearded along inner margin;
stigma diameter equal to style diameter, 0.1 mm. Pods and
seeds not seen.
Phenology
Flowering September–November.
Habitat and ecology
Moderately exposed areas or hill crests in gently undulating
country, including former marine plains. Woodland,
Revision of Pultenaea: Western Australian species
shrubland or heath, dense to very open. Sand or gravelly sand,
well drained. Usually scattered or rare, sometimes frequent
in patches.
Distribution
Western Australia: botanical districts Eyre and Roe. Found
in several locations near to the coast, particularly in
the Fitzgerald River area, but also collected in the
Stirling Range, north of Ravensthorpe and north of
Esperance (Fig. 4b).
Chromosome number
2n = 14. Sands (1975) estimated the diploid chromosome
number
of
the
invalidly
published
species
Pultenaea andrewsii [nomen nudum (Blackall and Grieve
1959)] to be 14, but the specimen on which she based
this count (Sands 638.19.4) is actually Pultenaea barbata.
Similarly, Keighery (1984) counted n = 7 for Pultenaea
adunca, but his voucher, too, is P. barbata.
Leaf anatomy
Pultenaea barbata has stomata on the lower leaf surface
only, idioblasts on both leaf surfaces and palisade mesophyll
on the upper side only (PG Turner and RPJ de Kok,
unpubl. data).
Discussion
Morphologically, this species is quite anomalous
within the study group. For example, it is the only species that
does not have stipules. The completely herbaceous texture
of the bracteoles is unique in the group, as are the basifixed
anthers, bearded style and scattered leaf arrangement. All
of this suggests it belongs in a different genus, not in
Pultenaea. A phylogenetic analysis performed by Orthia
et al. (2005) supports these suspicions, suggesting that the
species is closely related to the group of species currently
circumscribed as genus Phyllota, and should be removed
from Pultenaea to Phyllota. Morphologically, this shift is
justified. This species has several character states in common
with other Phyllota species: the same leaf arrangement
and shape, stipules absent or minute, and similar bracteole
texture and position (Bentham 1864). Some other Phyllota
species have a bearded style (Bentham 1864) and basifixed
anthers. An examination of several Phyllota herbarium
specimens housed at CANB confirmed their similarity to
Pultenaea barbata. Phyllota barbata Benth., in particular,
seems strikingly similar to Pultenaea barbata, and it
may be that the two are very close relatives. Although these
species share the same epithet, the two names were based
on different types. Interestingly, Blackall and Grieve (1959),
who lumped Phyllota into Pultenaea in their census of WA
plants, only mention one species with the epithet barbata,
possibly considering Pultenaea barbata and Phyllota
barbata to be conspecific. Andrews (1904), in describing
Australian Systematic Botany
165
Pultenaea barbata, noted the similarity with other Phyllota
species, but placed more emphasis on two characters
which it holds in common with Pultenaea: stamens free
from the petals and arillate seeds. In light of evidence in
this paper and Orthia et al. (2005), these characters seem
relatively unimportant.
However, pending a decision regarding Crisp and Cook’s
(2003) proposal to lump several genera, including Phyllota,
into genus Pultenaea, no name change is here proposed for
P. barbata. Once a decision is made, either Phyllota barbata
or Pultenaea barbata will require a name change, provided
that the two are not found to be conspecific.
Notes
Table 4 lists the characters distinguishing P. barbata from
taxa with which it is commonly confused.
Selected specimens (25 examined)
Near West Mount Barren, Blackall & Gardner s.n., x.1928 (PERTH);
Location 1110, approximately 30 km NNE of Young River crossing,
Donner 3042, 16.x.1968 (AD, CANB); Hopetoun at base of No Tree
Hill, Fuhrer s.n., ix.1987 (MEL); Base of Mt Toobrunup, Gardner 2196,
11.x.1928 (MEL); approximately 35 km NNW of Young River crossing,
Jackson 1419, 14.x.1968 (AD, CANB); S side of John Forrest Drive,
Jobson 2661, 23.x.1993 (MEL); 47 km S of Ravensthorpe, Keighery
633, 9.x.1974 (PERTH); approximately 2 km WSW of Quoin Head
Track turnoff, Lally 1300 & Lepschi, 6.xi.1996 (CANB, PERTH);
7 km N of Springdale Road, Lally 1309 & Lepschi, 7.xi.1996 (CANB,
PERTH); 3.9 km E along Norwood Road, Macfarlane 2134, 6.xi.1992
(CANB, PERTH); 1 mile N of Hopetoun, Newbey 656, 21.x.1962
(PERTH); 8.5 km NW of Mount Bland, Newbey 11012, 17.x.1985
(CANB, PERTH); locality not available, Newbey 11303, 24.x.1986
(PERTH); 3.9 km along Norwood Rd, Orthia 67, 30.x.2002 (CANB,
PERTH); 33 miles N of Ravensthorpe, Sands 638.19.4, 27.viii.1963
(AD, PERTH, SYD).
5. Pultenaea brachyphylla Turcz., Bull. Soc. Nat. Mosc.
26: 279 (1853). Typus: ‘Nova Hollandia’, n.d., Drummond
V 68 (holo-, KW (photo seen); iso-, BM!, K!, MEL!,
OXF!, W)
Pultenaea verruculosa var. brachyphylla (Turcz.) Benth., Fl. Austral.
2: 129 (1864).
Open shrub 0.2–0.5 × 0.3 m. Bark on lower stems grey,
plated and fissured, on upper stems reddish, smooth;
lower branches glabrous, upper branchlets densely hairy
to velutinous, woolly, hairs curly. Leaves pale green,
2.4–7.0 × 0.8–1.1 mm, length : width ratio 3–7 : 1, narrowly
oblong to oblanceolate, spiralled, appressed to inclined,
straight, venation obscure, lamina smooth, glabrous;
apex mucronate, acute, base truncate, margins involute;
petiole 0.2–0.5 mm, not decurrent. Stipules red-brown,
2.5–4.5 × 0.5–1.0 mm, not fused behind petiole, narrowly
triangular, straight, ciliate, sparsely to densely hairy on
body, going grey and fraying with age, apex acuminate,
indumentum woolly. Inflorescence terminal, indeterminate
heads of up to 19 sessile flowers, 8–10 flowering at any
166
L. A. Orthia et al.
Australian Systematic Botany
time; inflorescence bracts mostly leaf-like with small (≤3 mm
long) to very narrow (0.2 mm wide) bract leaves, bract
stipules to 4.5 mm, free, ciliate, sparsely to densely hairy
on body, indumentum woolly; flower-subtending bracts redbrown, completely scarious with very reduced leaf ± equal
in length to stipules, stipules 3.7–5.2 mm, ciliate, sparsely to
densely hairy on body, indumentum woolly. Bracteoles pale
brown, 3.7–4.0 × 0.9–1.0 mm, elliptic to falcate, inserted at
base of calyx, scarious, ciliate, body densely hairy, hairs
long and erect, apex acuminate. Calyx lobes ± equal in size,
two upper lobes fused slightly higher than other lobes, tube
3.0–3.5 mm, U–U 1.8 mm, U–L 2.7 mm, L–U 2.7 mm; tube
glabrous to sparsely hairy, lobes densely hairy, hairs erect,
terete; upper lobes 0.5 mm wide, deltate, apices acute, lower
lobes 0.6 mm wide, narrowly triangular, apices acuminate.
Standard gold-yellow with central yellow eye surrounded by
red line, back side with a black–dark red patch radiating
shortly to margins, margins and eye yellow, 9.5 × 6.5 mm,
round, slightly recurved, claw 4.2 mm, apex minutely notched
or rounded; wings gold-yellow with red at base, 8 × 1.5 mm,
held vertically over keel, near coherent for length; keel
black-red at apex grading to red at base, 8.0 × 1.8 mm,
± equal to wings in size. Anthers 0.6 mm, dorsifixed,
filaments 6.8–7.8 mm. Ovary 2.1 mm, cross-section elliptic,
velutinous, stipe negligible; style 4.8–5.0 mm, hooked at
apex, glabrous; stigma diameter slightly > style diameter,
0.2 mm. Pod 5.5 × 2.3 mm, ovate, sparsely hairy outside,
hairs curly or straight, glabrous inside. Immature seed seen,
appears to be exarillate.
of these populations are reserved within Fitzgerald River
National Park. The species is poorly known, and should be
further investigated with some urgency.
Chromosome number
Unknown. Sands (1966, 1975) states the chromosome
number of Pultenaea verruculosa var. brachyphylla to be
2n = 14. The voucher specimen listed for this count (Sands
638.16.2) has not been seen by us. However, another
Sands collection, Sands 638.16.12, has the same locality
description as that listed for the voucher specimen: ‘28 ml
E Jerramungup’ (Sands 1966). It is possible that Sands
misquoted the voucher collection number in both her
PhD thesis (1966) and her published article (1975), citing
638.16.2 when she should have cited 638.16.12. Evidence
for this is as follows. A Sands collection of a different
species of Pultenaea, Sands 638.16.7, has the locality 21 ml
E Gnowangerup. Sands’ numbering system for collections
was consecutive for the final digit, and it is unlikely that she
went from 28 miles east of Jerramungup (for collection of
638.16.2) to 21 miles east of Gnowangerup (638.16.7) and
back again to 28 miles east of Jerramungup (638.16.12).
As the 638.16.12 collection does not in fact belong to
Pultenaea brachyphylla, but to Pultenaea empetrifolia,
Sands’ chromosome number count for P. brachyphylla is not
accepted here.
Discussion
Undulating hills. Low heath or open mallee with low heath
understorey. Gravelly clay, sand or sandy loam. Rare to
common.
Pultenaea brachyphylla, originally described by
Turczaninow (1853), was reclassified as a variety of
Pultenaea verruculosa by Bentham (1864). However,
its morphology is quite different from that species in
many aspects of leaves, stipules, bracts and flowers. The
phylogenetic analysis in this study (Fig. 3) and in Orthia
et al. (2005) showed that the two are also genetically distinct,
falling into two different clades. Turczaninow’s classification
is therefore reinstated here.
Distribution
Notes
Western Australia: botanical district Eyre. Restricted to a
small area around the north-eastern part of Fitzgerald River
National Park (Fig. 6a).
Visited by butterflies and native bees (pers. obs.).
Dried flowers stain paper bright purple after boiling (pers.
obs.). The only other taxon in the study group noted to do
this is P. indira subsp. indira.
Pultenaea brachyphylla is closely related to P. indira. The
leaf apices of P. brachyphylla are mucronate while those of
P. indira are aristate. The bracts and stipules of P. brachyphylla
are densely hairy on the body, with tightly curled hairs,
distinguishing it from P. indira, which usually has hairs only
on the margins, or in the case of P. indira subsp. monstrosita,
sometimes has straight hairs on the body. The densely hairy
leaves of P. indira subsp. monstrosita also distinguish that
taxon from the glabrous-leaved P. brachyphylla.
The species is frequently misidentified. Table 6
lists the characters distinguishing P. brachyphylla from
Phenology
Flowering October.
Habitat and ecology
Conservation
Recommended here for conservation risk code 2K (Briggs
and Leigh 1996) and for Priority Two under CALM Priority
Flora Codes. The species has been collected fewer than ten
times in 153 years, judging from the herbarium specimens
seen in this study. Two of those collections were made
from the same location (Orthia 56 and Lepschi 3737), and
both indicated that the plants were rare at that site. All six
collections that specified a location were made at sites less
than 100 km apart, indicating that currently only a maximum
of five populations (possibly fewer) are known. At least two
Taxon
Leaf texture
Hairs on
mature leaves
Leaf transverse
posture
Vestiture on stipules
Bract stipule
apex shape
Bract vestiture
P. brachyphylla
P. empetrifolia
smooth
smooth
absent
absent
smooth or
verruculose
smooth
absent
densely hairy, hairs woolly
glabrous to densely hairy,
hairs woolly
glabrous
acuminate
usually acuminate
P. ericifolia and
P. strobilifera
P. indira subsp.
indira
P. indira subsp.
monstrosita
P. indira subsp.
pudoides
P. sp. Mt Lesueur
straight
straight to strongly
recurved
straight to recurved
absent
straight
ciliate margins, body glabrous
verruculose
straight
verruculose
present, short / fuzzy
and / or long / erect
present, long / erect
recurved at apex
ciliate margins, body glabrous
to densely hairy, hairs straight
ciliate margins, body glabrous
acuminate
verruculose
present, long / erect
straight to incurved
glabrous
acuminate
P. radiata
verruculose
present, long / erect
incurved
glabrous
P. verruculosa
smooth or
verruculose
absent or present,
long / erect
straight to incurved
glabrous
acute or fused to
other stipule
acute
densely hairy, hairs woolly
glabrous to hairy, hairs woolly
or straight
ciliate margins, hairy on body,
hairs straight
margins strikingly ciliate,
body glabrous
ciliate margins, body glabrous
to velutinous, hairs straight
margins strikingly ciliate,
body glabrous
ciliate margins, body densely
hairy, hairs straight
glabrous
may or may not
be acuminate
acuminate
acuminate
Revision of Pultenaea: Western Australian species
Table 6. Characters used to distinguish species with prominent stipules that are not fused behind the petiole (P. ericifolia / P. indira group)
small leaf densely hairy,
body glabrous, margins
may be minutely ciliate
Bract margin texture
Inflorescence
bract number
Bracteole shape
and size
Petal colours
Stigma diameter relative to style
P. brachyphylla
P. empetrifolia
smooth to frayed
smooth to frayed
± equal
± equal
smooth
lanceolate, small
lanceolate, small or rarely
with green leaf lobe
lanceolate, small
yellow and black-red
yellow and red
P. ericifolia and
P. strobilifera
P. indira subsp. indira
P. indira subsp.
monstrosita
P. indira subsp.
pudoides
P. sp. Mt Lesueur
a few
a few to many, do not
form involucre
many, forming involucre
much larger, bulbous
papillose
smooth to frayed
a few
a few
lanceolate, small
lanceolate, small
papillose
a few
lanceolate, small
bright to pale pink,
orange and yellow
yellow and black-red
unknown (probably
yellow and black-red)
yellow and black-red
±equal
smooth to frayed
lanceolate, small
gold-yellow and bright pink
much larger, bulbous
P. radiata
P. verruculosa
smooth to frayed
smooth or frayed
many, forming weak
involucre
a few
a few
round, large
lanceolate, small
vivid orange, pink and yellow
orange, red, and pink
(rarely all yellow)
much larger, bulbous
±equal
±equal
±equal
Australian Systematic Botany
Taxon
167
168
Australian Systematic Botany
morphologically similar taxa, and Fig. 5 compares the bract
morphology of similar taxa.
Specimens examined
2 km NW of boundary of Fitzgerald River National Park, Corrick
8812, 19.x.1983 (CANB); NE of Jerramungup on road to
Ravensthorpe, Lepschi 3737, 27.x.1997 (CANB); Along Highway 1
from Jerramungup, Orthia 56, 26.x.2002 (CANB, PERTH); Pultenaea
Reference Set, Williamson s.n., n.d. (MEL2055285).
6. Pultenaea brachytropis Benth., in Lindl., Edwards’s Bot.
Reg. 27 (1841), Misc. notes No. 76, p 40. Typus: no.loc.,
1841, Hort. Lindley s.n. (holo-, K!)
Pultenaea drummondii Meisn., in Lehmann, J.G.C., Pl. Preiss. 2:
219–220 (1848). Typus: ‘Swan River’, 1844, Drummond II 127 (holo-,
NY (photo seen); iso-, BM!, K (3 sheets)!, OXF!).
Illustrations
Grieve, How to Know Western Australian Wildflowers 2: 420
(1998).
L. A. Orthia et al.
Open to dense, bushy erect shrub 0.25–1.5 ×
0.2–1(–2.5) m. Lower stems glabrous, bark metallic
looking, fissured, flaking off, wood red underneath;
upper stems orange, sometimes with bright green spots,
smooth, sparsely to densely hairy, hairs terete, long and
± appressed. Leaves bright to dark green above, pale green
below, 3.0–18.7 × 0.9–3.0 mm, length : width ratio 5–8 : 1,
oblong or linear, spiralled, inclined to appressed, midvein
prominent, lamina verruculose, upper surface glabrous,
lower surface sparsely hairy; hairs erect and terete; apex
acute and mucronate but recurved so as to appear blunt,
base rounded, obtuse, margins strongly recurved; petiole
1.0 mm, not decurrent. Stipules pale gold to transparent,
1.5–4.0(–6.0) × 0.5–1.0 mm, fused behind the petiole,
± narrowly triangular, straight, strongly keeled, thin and
filmy, glabrous, breaking off and fraying with age, apex
acuminate. Inflorescences terminal, determinate clusters of
3 to >10 ± sessile flowers; inflorescence bracts leaf-like,
flower-subtending bracts golden brown, 4.8 × 2.4 mm, leaf
strongly reduced or absent, stipules enlarged, margins
Fig. 5. Flower-subtending bract morphology for the species with prominent stipules not fused behind the petiole (P. ericifolia / P. indira group),
showing differences in the shape, fusion level and indumentum.
Revision of Pultenaea: Western Australian species
ciliate, body glabrous and shiny, scarious, filmy. Bracteoles
golden brown, 4.0 × 1.5 mm, elliptic to ovate, inserted
at base of calyx, strongly keeled, scarious, filmy; apex
acute, glabrous to sparsely hairy. Two upper lobes of calyx
united into lip and fused for most of length, upper lobes
larger than lower lobes, tube 3.0 mm, U–U 0.8 mm, U–L
3.4 mm, L–U 2.2 mm, calyx sparsely to densely hairy,
hairs golden, long, terete, appressed; upper lobes 1.7 mm
wide, widest point at base, apex rounded; lower lobes
1.1–1.4 mm wide, deltate, apex acute. Standard golden
yellow at margins, eye yellow / red flecked surrounded by red
smudgy radiating lines, back pale yellow, 10.5 × 10.5 mm,
reniform, recurved, claw 2.6 mm, apex notched; wings red
apically, fading at base, 8.2 × 1.5 mm, held lateral to keel,
not touching each other; keel white / cream, 5.2 × 1.8 mm,
much shorter than wings. Anthers 0.4 mm, dorsifixed,
filaments 4.0 mm. Ovary 1.5 mm, cross-section elliptic,
densely hairy, sessile; style 3.1 mm with sharp, 90◦ bend
towards stigma, glabrous; stigma diameter conspicuously
greater than style diameter, 0.3 mm. Pod 5 mm, obovate,
exterior densely hairy with fine gold hairs, interior glabrous.
Seed not seen.
Phenology
Flowering September–November, occasionally July; fruiting
November.
Habitat and ecology
Flat areas or slight slopes, often in damp, swampy areas and
drainage lines, also on ridges. Open woodland, woodland,
open forest, forest, occasionally shrubland, associated with
Pultenaea radiata. Sand, clay, loam and intermediates, often
wet, sometimes humus rich, often with gravel, over laterite
or granite. Occasional to common, sometimes understoreydominant.
Distribution
Western Australia, botanical district Darling, subdistricts
Menzies, Drummond and Warren. Mainly clustered in the
area roughly bounded by Busselton, Nannup and Augusta,
with one dubious outlying population at Ellenbrook, north of
Perth (White s.n.) (Fig. 4c).
Leaf anatomy
Pultenaea brachytropis has stomata only on the lower
leaf surface, idioblasts associated with vasculature
on both sides of the leaf and palisade mesophyll on the upper
side only (PG Turner and RPJ de Kok, unpubl. data).
Discussion
The holotype is the only specimen examined that did not have
scabrous leaves—the leaves of the holotype were soft and
smooth. All other characters matched the other specimens
labelled both P. brachytropis and P. drummondii, so the
Australian Systematic Botany
169
specimen is certainly the same species. This indicates that
the scabrous texture might be environmentally determined,
because the holotype was grown in a greenhouse or garden
in England.
The plant is usually ≤1 m in height, but one specimen
(Royce 3920) describes the height as 6–8 ft (∼1.8–2.4 m).
Notes
This species has no close relatives within Pultenaea or tribe
Mirbelieae, forming a monotypic lineage within the tribe
(Orthia et al. 2005).
Grazed by kangaroos (Alferink 12). Visited by honeybees
(pers. obs.).
Selected specimens (75 examined)
Approximately 500 m N of Gnarawary Road, S of Margaret River,
Alferink 12, 4.xi.1998 (PERTH); 1.6 km N of Warner Glen Road,
Annels 2722, 15.x.1992 (CANB, PERTH); Augusta, Ashby 2374,
12.x.1967 (AD, CANB); Brockman Highway, Corrick 8924, 25.x.1983
(AD, CANB); 5 km S of Margaret River township on Caves Road,
Corrick 8957, 28.x.1983 (MEL, PERTH); 18 km SE of Busselton,
Corrick 9219, 10.x.1984 (CANB, PERTH); Whicher Range (west
end), Treeton–Jindong road near turn-off to ‘Scented Garden’,
Corrick 9736, 4.xi.1985 (MEL); 10 miles E of Alexander Bridge,
Cranfield 955, 2.xi.1978 (CANB, MEL, NSW); Augusta, Western
Australia, Gilbert 248, 1842 (BM, K); On Vasse Highway, 18.6 km
from start of highway in Busselton, Orthia 37, 20.x.2002 (CANB,
K, MEL, PERTH); Kudardup, Royce 3920, 24.x.1952 (CANB);
Brockman highway, Taylor & Ollerenshaw 2039, 21.ix.1983 (AD,
CANB, NSW, PERTH); Between Alexandra Bridge and Steward
Road on Brockman Highway, Whibley 5090, 6.xi.1974 (AD, PERTH);
Ellenbrook, White s.n., n.d. (AD); Whicher Range, Woolcock W2241,
20.ix.1985 (CANB, MEL).
7. Pultenaea calycina (Turcz.) Benth. Fl. Austral. 2: 121
(1864). Euchilus calycinus Turcz., Bull. Soc. Nat. Mosc. 26:
276 (1853). Typus: ‘Swan River’, 1849, Drummond V 75
(holo-, KW (photo seen); iso-, CGE, E, K (3 sheets)!, BM!,
OXF!, NSW!, W)
Illustrations
Grieve, How to Know Western Australian Wildflowers 2: 423
(1998).
Many-branched, compact shrub up to 0.3–1.5 × 0.2–2 m.
Lower stems glabrous to sparsely hairy, bark brown,
fissured and knobbly; upper branchlets yellow,
smooth, velutinous, hairs short, straight, appressed to
ascending. Leaves pale to olive green or bright green,
(3.0–)4.4–12.9 × 0.7–3.3 mm, length : width ratio 4–11 : 1,
oblong to linear or clavate to oblanceolate, usually
in whorls of three but spiralled when foliage is lax
and rarely in whorls of four or decussate, divergent to
ascending, straight to incurved, midvein may or may
not be prominent, secondary veins sometimes visible,
lamina smooth, glabrous to sparsely hairy; hairs erect,
fine; apex obtuse, sometimes minutely notched, base
170
L. A. Orthia et al.
Australian Systematic Botany
truncate, margins incurved to involute but may open up at
apex; petiole 0.5–1.0 mm, not decurrent. Stipules black to
black-red, 0.4–1.5 × 0.2–0.6 mm, not fused but some stipule
material sometimes present behind petiole, triangular
to narrowly triangular, straight to recurved, glabrous to
sparsely hairy, brittle, often chipped and broken or absent
from older nodes, apices acute if intact. Inflorescences
at branch termini or on short axillary stems near
terminus of branchlets, indeterminate racemes of 1 to >20
flowers, pedicels 0.5–3.0 mm, bracts leaf-like; each flower
subtended by stipules, but bract leaves present or absent.
Bracteoles red-brown, 1.5–3.9 × 0.4–0.8 mm, ovate to
lanceolate, inserted at base of calyx, scarious, glabrous.
Two upper calyx lobes much larger than three lower lobes,
tube 0.9–2.2 mm, U–U 4.0–5.5 mm, U–L 3.4–5.5 mm,
L–U 1.0–2.7 mm, upper and lower lobes coherent for
25–50% of upper lobe length, glabrous to sparsely
hairy, hairs long (c. 0.8 mm) and erect or short and
appressed; upper lobes 2.3–3.8 mm wide, elliptic to obovate,
apex obtuse, sometimes with small mucro; lower lobes
0.3–0.6 mm wide, deltate or narrowly triangular, apex
acuminate. Front of standard deep yellow with central
yellow eye surrounded by thin red line, back mostly
yellow but sometimes pale red apex, 5.5–7.4 × 7.4–8.6 mm,
reniform, recurved, claw 1.2–1.7 mm, apex minutely
notched; wings yellow, sometimes red tinges at apex,
5.5–6.3 × 1.6–2.0 mm, held over keel or held apart;
keel pale yellow with pale red apex externally, brighter
with larger red patch internally, 5.5–6.6 × 1.5–2.9 mm,
sometimes protruding slightly through wings. Anthers
0.4 mm, dorsifixed, filaments 4.5–4.8 mm. Ovary
1.5–2.5 mm, laterally flattened, densely hairy, stipe
0.2–0.5 mm; style 2.2–2.8 mm, hooked, glabrous; stigma
diameter ± equal to style diameter, 0.1–0.2 mm. Pod and
seed not seen, but mature ovaries laterally flattened and
sparsely hairy.
Chromosome number
the same species. Some collections of each subspecies show
leaf morphology that tends towards an intermediate form
(e.g. Orthia 62, Wilson 4377).
Notes
Phylogenetic evidence from this study and Orthia et al.
(2005), as well as many aspects of morphology, suggest
P. calycina is closely related to P. quaerita. The two species are
distinguished by the length : width ratio of leaves (P. calycina
has a ratio ≥ 4 : 1; P. quaerita has a ratio ≤ 2 : 1).
Pultenaea calycina is poorly known. Table 5 lists the
characters distinguishing it from morphologically similar
taxa.
Key to subspecies of P. calycina
Leaves oblong or slightly obovate and broadly U-shaped in crosssection.............................................................................7.1. subsp. calycina
Leaves clavate and strongly obovate and involute............ 7.2. subsp. proxena
7.1. Pultenaea calycina (Turcz.) Benth. subsp. calycina
Leaf length : width ratio 4–6 : 1; leaves oblong or slightly
obovate, straight, ±flat to broadly U-shaped in crosssection, midvein prominent, secondary veins sometimes
visible, lamina glabrous, margins incurved; inflorescences
of 3 to >20 flowers; calyx red, sparsely hairy, hairs long
and erect; wings not touching, keel protruding slightly
through wings.
Phenology
Flowering August–October, occasionally March.
Habitat and ecology
Flat plains, hills. Mallee woodland with low shrub
understorey. Gravelly sand or sandy clay. Locally frequent
to abundant.
Distribution
2n = 16 (Sands 1966, 1975). The count is based on both
haploid and diploid counts of P. calycina subsp. calycina
(Sands 1975).
Western Australia, Eyre botanical district. Known from a
few scattered populations stretching from the plains at the
south-east of the Stirling Range across to West Mount Barren
(Fig. 9b).
Discussion
Conservation
Pultenaea calycina subsp. proxena is morphologically very
similar to subspecies calycina, differing primarily in the
shape of the leaves (clavate v. oblong) and the curvature
of the leaf margins (involute v. flat to broadly incurved).
These characters give the two subspecies a very different
appearance at superficial examination. The two subspecies
are also separated geographically, which may account for
the different morphology. However, the striking similarity of
floral characteristics and other vegetative traits suggests that
the two taxa have only recently diverged and still belong to
Pultenaea calycina subsp. calycina has been collected fewer
than ten times since Drummond’s collection in 1849 / 1850.
Briggs and Leigh (1996) list the taxon as conservation risk
code 3KC–, indicating a poorly known taxon with range
>100 km and with at least one population (of unknown
size) in a conservation reserve. The conservation reserve
they refer to is Hassell National Park (Briggs and Leigh
1996). No evidence of populations residing in Hassell
National Park was found in this study, but the taxon has
certainly been collected from the area now contained within
Revision of Pultenaea: Western Australian species
Fitzgerald River National Park. In addition, at least one
population is found just outside the south-east corner of
Stirling Range National Park, and it is possible that there
are populations within the park too. However, this must
be further investigated, as the last collection of P. calycina
subsp. calycina in the Fitzgerald River National Park area
was in 1966. It may be that the species is not currently
reserved at all. In regard to the range of the taxon, known
populations are probably only 100 km apart at most, so
the current conservation risk code might be inappropriate.
The number of known populations is uncertain, since some
given collection localities are vague, but it could be as few
as three. Certainly there is a population near Kojoneerup
(Orthia 51, Davis 2930), there are or were one or two in the
Fitzgerald River area (Gardner s.n., Wilson 4377) and there
is or was one on the Hassell Highway somewhere (collector
indecipherable). The additional twentieth century collections
(Sands 638.14.28, Sands s.n., Lakes s.n.) may or may not
represent these same populations. It is recommended
here that the species remain listed as conservation risk
code 3KC– pending further investigation. It is also
recommended here for Priority Two under CALM Priority
Flora Codes, given the uncertainty about the number of
extant populations.
Specimens examined
Kojoneerup Springs Road, Davis 2930, 19.iii.1997 (PERTH); West
Mount Barren, Gardner s.n., viii.1939 (PERTH); N of Bremer Bay,
Lakes s.n., ix.1958 (PERTH); Kojoneerup Springs Road, Orthia 51,
24.x.2002 (CANB, K, MEL, PERTH); North of Manypeaks, Sands
s.n., 2.ix.1963 (PERTH); Chillinlup to Keerjanup, Sands 638.14.28,
20.viii.1963 (AD, PERTH, SYD); Pultenaea Reference Set, Williamson
s.n., n.d. (MEL 2057344); Along No. 2 Rabbit proof fence, Wilson
4377, 2.x.1966 (CANB); Hassell Highway, collector indecipherable,
19.ix.1964 (PERTH 624683).
7.2. Pultenaea calycina subsp. proxena Orthia &
Chappill, subsp. nov.
A subspecie calycina foliis clavatis et involutis differt. Typus:
[locality abbreviated to protect taxon] E of Ravensthorpe
along road to Esperance, W.A., 5.xi.1992, Macfarlane 2122
(holo-, CANB 508683!; iso-, CANB, PERTH!).
Leaf length : width ratio usually 8–11 : 1; leaves clavate,
oblanceolate, straight to incurved, terete, venation not
prominent, lamina glabrous to sparsely hairy, margins
involute but sometimes opening up at apex; inflorescences
generally 1–5 flowers; calyx red to green, often glabrous,
hairs long / erect or sometimes short / appressed; wing
petals held over keel or held apart, keel may protrude
through wings.
Phenology
Flowering August–November.
Australian Systematic Botany
171
Habitat and ecology
Moderate slopes, including adjacent to creek bed. Mallee
woodland or open woodland, including very young stands,
with low to medium shrub understorey. Sand, clay, sandy
clay or loam, with gravel, over magnesite or clay. Rare to
local understorey-dominant.
Distribution
Western Australia, Eyre botanical district. Known from three
geographically close locations east of Ravensthorpe (Fig. 9c).
Conservation
As an undescribed taxon [Pultenaea sp. Bandalup (G.F.Craig
3625)], P. calycina subsp. proxena is listed on FloraBase
as Priority One under CALM Priority Flora Codes. It is
recommended here for conservation risk code 2K (Briggs
and Leigh 1996). The taxon is now known from several
populations, some of which contain many individuals
(G. Craig, pers. comm.).
Etymology
The name of the new subspecies, P. calycina subsp.
proxena, refers to the fact that on superficial examination,
the herbarium specimens of this taxon look nothing
like P. calycina subsp. calycina (L. prox- = ‘alike’;
L. en = ‘indeed?’). It also stands as a memorial to a
fallen lesbian hero (L. pro- = ‘alas’ or ‘for’; Hollywood
xena = ‘warrior princess’).
Specimens examined
ESE of Ravensthorpe, Craig 3625, 18.ii.1998 (PERTH); Bandalup
Hill, Craig 5744, 28.xi.2002 (PERTH); E of Ravensthorpe, Macfarlane
2122, 5.xi.1992 (CANB, 2 sheets); along South Coast Hwy, Orthia
62, 27.x.2002 (CANB, K, MEL, PERTH); Bandalup Hill, Orthia 63,
27.x.2002 (CANB, K, MEL, PERTH); SE of Ravensthorpe, Williams
157, 16.viii.1999 (PERTH).
8. Pultenaea daena Orthia & Chappill, sp. nov.
A P. arida E.Pritz. habitu prostrato et pilis erectis differt.
Typus: [locality abbreviated to protect taxon] W of
Ravensthorpe W.A., 27.x.1997, Lepschi & Fuhrer 3743
(holo-, PERTH; iso-, CANB!).
Prostrate shrub, sometimes partly covered by soil, to 1 m
across, shooting from rhizomes. Bark fissured on rhizomes;
upper branchlets smooth, sparsely hairy, hairs erect, foliage
sometimes extremely dense. Leaves pale grey-green,
2.4–5.3 × 1.3–2.4 mm, length : width ratio 2 : 1, elliptic,
decussate, ascending to inclined, straight to slightly incurved,
midvein prominent on lower surface, lamina smooth, sparsely
to densely hairy; hairs erect, terete, up to 1.3 mm; apex
acute, base acute, margins flat or slightly incurved; petiole
0.5–0.7 mm, not decurrent. Stipules red-brown, 1.2–2 mm,
not fused but some stipule tissue present behind petiole,
172
L. A. Orthia et al.
Australian Systematic Botany
narrowly triangular, straight, sparsely hairy, may break
off with age, hairs erect, apex acuminate. Inflorescences
axillary, in upper axils, indeterminate, flowers solitary,
pedicels 2.3–3.6 mm, flower bracts leaf-like. Bracteoles redbrown, 2.5–3.5 × 0.2–0.4 mm, lanceolate to linear, inserted
at base of calyx, scarious, sparsely hairy, apex acuminate.
Two upper lobes of calyx much larger than three lower
lobes, tube 0.9–1.8 mm, U–U 3.8 mm, U–L 3.8 mm, L–U
3 mm, upper and lower lobes coherent for 75–80% of upper
lobe length, sparsely to densely hairy, hairs erect, terete
and long (0.5–1.0 mm); upper lobes 1.1–1.3 mm wide,
elliptic, apex acute; lower lobes 0.5–0.8 mm wide, narrowly
triangular, apex acuminate. Petals uniformly orange-yellow
to orange-yellow with dark red apices; standard may have
dark red at edges, dark red on back side and a dark red line
around central yellow eye, 4.5–6.7 × 5.0–6.7 mm, round,
recurved or slightly inwardly folded, claw 1.0–1.5 mm, apex
minutely notched; wings 4.0–6.0 × 1.2–2.1 mm, held over
keel, coherent for length; keel 4.0–6.3 × 1.2–3.1 mm, ± same
size as wings. Anthers 0.4–0.5 mm, dorsifixed, filaments
5.5 mm. Ovary 1.3–2.3 mm, laterally flattened, densely hairy,
stipe 0.2–0.3 mm; style 3–3.9 mm, curved, glabrous; stigma
equal to style diameter, 0.1 mm. Pod 4.3–4.5 × 3–3.1 mm;
seed black, 2.2–2.6 × 1.6–1.7 mm, reniform, exarillate.
Phenology
Flowering October–December, and also in March.
Habitat and ecology
Flat or gently undulating hills. Mallee eucalypt woodland,
with grasses and shrubby regrowth. Growing in very open
areas on edge of gravel road, on edge of salt lake. Very fine,
gravelly sandy-loam or sand. Rare.
Distribution
Western Australia, botanical district Roe. Known from
several localities in the region SE of Lake Cobham (west
of Ravensthorpe), from much further north-east, and from a
locality ∼50 km ENE of Ravensthorpe (Fig. 9d).
Conservation
As an undescribed taxon [Pultenaea sp. Fitzgerald
(B.J.Lepschi & B.A.Fuhrer BJL 3743)], P. daena is listed
on FloraBase as Priority One under CALM Priority Flora
Codes. Recommended here for conservation risk code 3K
(Briggs and Leigh 1996). This species has been collected
fewer than ten times. Where it was collected, it was noted to
be rare. However, it does appear to be reasonably widespread,
with outlying populations ∼100 km apart. A large thriving
roadside population sampled in this study appeared, at a
subsequent visit, to have been severely damaged. A vehicle
had driven up on the embankment over the largest of the plants
and many smaller plants in the adjacent remnant vegetation
had been uprooted, possibly by sheep grazing. The species
has not been collected from conservation reserves, and it
may be a priority to establish some reserves to protect it from
further harm.
Notes
Pultenaea daena is most closely related to P. arida and
P. elachista. It differs from both P. arida and P. elachista
in having densely hairy leaves and a prostrate habit.
Table 5 lists the characters distinguishing P. daena from
morphologically similar species.
Etymology
The species epithet, ‘daena’, is the name of a Zoroastrian
goddess whose name means ‘that which was revealed’, a
suitable name for a new species. Daena was associated
with the first light of dawn, an image that resonates with
the orange and red colours and radiate markings of the
standard petal of Pultenaea daena. Daena’s mother, Armaiti,
is associated with the earth, and is the goddess of fertility
and the dead (who are buried beneath the earth), an
appropriate association for this half-buried shrub. The species
also honours Daena Murray, the first author’s mother, with
the collections Orthia 57 and Orthia 58 being made on
her birthday.
Specimens examined
Lake Magenta, Chappill 6548, 28.xi.2000 (PERTH); Lacum Cronin,
Gardner 15928, 10.xii.1964 (PERTH); W. of Ravensthorpe, Lepschi
3743, 27.x.1997 (CANB); Fitzgerald Road, Orthia 57, 26.x.2002
(CANB, K, MEL, PERTH); West River Road, Orthia 58, 26.x.2002
(CANB, K, MEL, PERTH); Lake Cobham, S. Walsh 88, 15.iii.2000
(MEL, PERTH).
9. Pultenaea elachista (F. Muell.) Crisp
This species was treated in Part III of de Kok and West’s
revision of the genus (2004), so is not given full treatment
here. Their description of P. elachista is largely consistent
with the morphology of the plant in Western Australia.
However, they overlooked some important traits, particularly
the form of the calyx, so some amendments and additions
to their description are made here. Phenology, habitat,
ecology and distribution specific to Western Australia are also
included, based on Western Australian herbarium specimens
to which de Kok and West did not have access for their
revision.
Prostrate, decumbent or erect shrub, 0.2–0.5 × 0.27–1 m.
Leaf length commonly ≥ 5 mm (−7 mm), width up to 4 mm.
Inflorescence often a lax terminal raceme. Two upper calyx
lobes much larger than three lower lobes, tube 1.5–1.7 mm,
U–U 5.6–7.8 mm, U–L 5.2–6.5 mm, L–U 4.2–4.8 mm, upper
and lower lobes coherent for 33–50% of upper lobe length,
densely hairy to velutinous, hairs both long / erect and
short / curly; upper lobes 1.9–3.3 mm wide, elliptic-obovate,
apex obtuse and mucronate; lower lobes 0.7–1.0 mm wide,
Revision of Pultenaea: Western Australian species
narrowly triangular, apex acuminate with long arista. The
three aristate apices of the lower lobes are not adherent
to other parts of the calyx in bud, and protrude from the
bud in a distinctive tridentate arrangement. Standard may
be entirely yellow or yellow with two faint red stripes at
base on front.
Phenology
Flowering July–January, occasionally March.
Habitat and Ecology
Plains to gentle slopes, plateau and coastal scarp.
Shrubland, open woodland or woodland, sometimes disturbed
vegetation. Sand, clay, loam or intermediates, or gypsum, over
limestone [plus, rarely, on granite outcrops (Hopper et al.
1997)]. Rare to abundant.
Distribution
In Western Australia, botanical districts Roe, Coolgardie and
Eucla. Widespread, on calcareous soils, from the Cascade and
Peak Charles areas in the west, across the Nullarbor Plain into
South Australia in the east (Fig. 9e).
Notes
Pultenaea elachista is most closely related to P. arida and
P. daena.
Selected specimens (54 examined)
9 miles SW of Mount Ragged, Aplin 4306, 19.x.1970 (CANB, PERTH);
5.5 km SW of Mount Willgonarinya, Archer 2710906, 27.x.1990
(CANB, PERTH); c. 6 km S of Mt Buraminya, Archer 1709958,
17.ix.1995 (MEL); West end of Great Bight, Carey s.n., 1877 (MEL);
c. 95 km along Parmango Road from Heywood Road, Corrick 11482,
3.xi.1996 (MEL, PERTH); 11.5 km N of Shearer Rd on Parmango
Rd, Craig 3015, 14.xi.1993 (PERTH); 8.5 km NW along Cascade
Road from Beltana Road, Crisp 9328, 1.x.2001 (CANB); 8 km NW
of Mount Ragged, Keighery 1999, 19.xi.1988 (PERTH); Intersection
of Norwood Road and Wittenoom Road, Macfarlane 2135, 6.xi.1992
(CANB, PERTH); 0.4 km N of Kendall Rd on Norwood Rd, Orthia
66, 30.x.2002 (CANB, PERTH); 0.7 miles North of Grass Patch,
Reid 644, 28.ix.1974 (MEL); 111 km S of Balladonia towards Mount
Ragged, Taylor 1537, 8.ix.1983 (CANB, PERTH); Mallee near B.M.
-GJ 24, Cape Arid National Park, Weston 9607, 5.x.1974 (CANB,
PERTH); 4 km S of Caiguna, Wittwer W.1973, 11.xi.1976 (PERTH);
Eyre Highway, 0.5 km from SA / WA Border, Woolcock W2260, 7.x.1985
(CANB, MEL).
10. Pultenaea empetrifolia Meisn., in Lehmann, J.G.C.,
Pl. Preiss. 1: 76 (1844). Typus: ‘In Australia occidentali’,
Herb. Preiss. No. 865 (holo-, LD (photo seen); iso-,
BM!, NY, S)
Pultenaea verticillata Turcz., Bull. Soc. Nat. Mosc. 26: 279 (1853).
Typus: ‘Nova Hollandia’, 1849, Drummond. V 64 (holo-, KW?; iso-,
BM!, K!, NSW!, OXF!)
Low, spreading, prostrate to erect shrub, 0.1–0.7 ×
0.2–0.5 m. Lower stems grey over red wood, glabrous, bark
thin and flaking to coarse and plated; upper stems grey,
Australian Systematic Botany
173
smooth, densely hairy to velutinous, hairs short and curly.
Leaves olive green to bright green, 1.6–4.0 × 0.5–0.9 mm
(up to 6.0 mm long when straightened out), length : width
ratio 3–6 : 1, linear and oblong to oblanceolate and obovate,
spiralled, appressed to divergent, straight to strongly
recurved, venation obscure, lamina smooth, glabrous; apex
blunt to aristate, acute to obtuse, base truncate, margins
involute; petiole negligible, not decurrent. Stipules redbrown, dark brown or grey, 0.7–2.5 × 0.3–0.7 mm, not fused
behind the petiole, ovate, deltate or narrowly triangular,
appressed to slightly recurved, glabrous to densely hairy,
can have ciliate margins and/or hair on the body; hairs
curly; chipping or fraying with age, stipule apex acuminate.
Inflorescences terminal, indeterminate, 1 to >20 sessile
flowers, usually >3, subtended by few to many inflorescence
bracts which vary greatly in size and shape; inflorescence
and flower bracts light to red-brown, up to 6.5 × 3.0 mm,
leaf-like with large, herbaceous central lobe to tridentate
with strongly reduced, scarious central lobe, can be glabrous,
ciliate and/or hairy on body. Bracteoles pale brown to redbrown, 2.3–5.0 × 0.7–1.4 mm, may be entire, completely
fused and scarious or may have herbaceous, green to brown,
large, partially free central lobe; falcate to ovate, inserted at
base of calyx, glabrous, ciliate or with dense medial hairs,
apex acute or acuminate. Calyx lobes ± equal in length,
tube 2.0–3.2 mm, U–U 1.4–1.8 mm, U–L 1.8–2.8 mm,
L–U 1.8–2.7 mm, glabrous to densely hairy, hairs long and
erect; lobes 1.0–1.2 mm wide, narrowly triangular, apex
acuminate. Standard yellow with slightly paler eye area
surrounded by a few short, thin red radiating lines at front,
back yellow at margins with central dark red patch radiating
out along veins and surrounding yellow / white eye, or
yellow-gold with faint red markings, 6.2–9.0 × 5.4–7.5 mm,
reniform, recurved or inwardly folded, claw 2.5–4.0 mm,
apex sometimes minutely notched; wings gold-yellow,
sometimes with reddish basal striations, 5.6–8.0 ×
1.4–2.1 mm, held ± over keel, may be coherent or held
apart; keel dark to bright red, darker at apex, 5.8–7.8 ×
1.8–2.3 mm, ± same size as wings. Anthers 0.3–0.6 mm,
dorsifixed, filaments 4.5–7.0 mm. Ovary 2.0–2.5 mm,
cross-section elliptic, velutinous, stipe negligible; style
4.0–5.0 mm, gently curved or hooked, glabrous; stigma
diameter ± equal to style diameter, 0.15–0.2 mm. Pod
3.5 × 1.8 mm, ovoid, sparsely hairy outside, erect and terete
hairs, glabrous inside. Seeds speckled brown, 2.4 × 2.8 mm,
± circular, arillate.
Phenology
Flowering August–November; fruiting October.
Habitat and ecology
Plains, small hills, valleys or swamps. Heath, mallee
shrubland, mallee open woodland or mallee woodland with
shrub / heath understorey, often open, sometimes disturbed
174
L. A. Orthia et al.
Australian Systematic Botany
roadside vegetation, associated with Pultenaea verruculosa.
Usually sandy gravelly soil, sandy clay, sandy loam or laterite.
Rare to common.
Chromosome number
2n = 14 (Sands 1966, 1975). The count is based on both
haploid and diploid counts (Sands 1975).
Distribution
Western Australia, botanical districts Eyre, Roe, Avon and
Darling (subdistrict Menzies). Widespread over the region
roughly bounded by Ravensthorpe, Kukerin and Frankland,
and reaching the coast in the Bremer Bay / Fitzgerald River
area (Fig. 6b).
Leaf anatomy
Pultenaea empetrifolia has stomata only on the
upper leaf surface and idioblasts only on the lower
side of the leaf (PG Turner and RPJ de Kok,
unpubl. data).
Fig. 6. Distribution maps of (a) P. brachyphylla, (b) P. empetrifolia,
(c) P. indira subsp. indira, (d) P. indira subsp. monstrosita and
(e) P. indira subsp. pudoides.
Revision of Pultenaea: Western Australian species
Typification
The LD specimen is likely to be the holotype of
P. empetrifolia, as for all species described in Plantae
Preissianae (Crisp 1983).
Some type sheets of P. verticillata (e.g. BM) contain
material including both P. empetrifolia and P. neurocalyx
Turcz.
Discussion
Pultenaea empetrifolia is a morphologically diverse species,
varying in aspects of the leaf shape and posture, calyx
hairiness, bract type and bracteole form. Some specimens,
found towards the south-west of the range, resemble the
type specimen in having relatively short, straight, blunt,
appressed leaves, glabrous calyces and bracts with a large,
herbaceous central lobe. Other specimens, found in the
north-east of the range, have long, strongly recurved,
aristate, divergent leaves, hairy calyces and bracts with a
reduced, scarious central lobe. Initially in this study the two
forms were considered to be separate taxa, distinguished
primarily by the bract differences. However, numerous
intermediate forms exist, and this study was unable to find
any characters that reliably differentiate between the forms
in all cases. For example, Macfarlane 2099 and Aplin 3671
both have a glabrous calyx, but their bracts are completely
scarious and the central lobe is strongly reduced. Other
specimens have bracts of mixed types: some bracts have
a scarious / reduced central lobe and others on the same
plant are herbaceous / large. There is also a continuum of
variation in calyx hairiness and in leaf size, shape and
posture, which is not at all correlated with bract form. Other
characters initially thought to be potentially useful—stipule
hairiness, stipule shape and bract hairiness—also showed a
continuum of variation to some extent. Therefore, these very
different forms are here considered to belong to the same
species, P. empetrifolia.
There is one reasonably distinct form found in the
Tarin Rock region, in the far north of the range. It differs
from the other members of the species in the shape of
its bracteoles: its central bracteole lobe is herbaceous,
large and incompletely fused with its stipules, while
other members of P. empetrifolia have bracteoles that
are completely fused and scarious. The Tarin Rock form
also sometimes has a standard petal that is larger than
that of most P. empetrifolia plants, and that is very
bright yellow with fewer red markings than usual. Of all
P. empetrifolia forms, its leaves are the longest and the
most strongly recurved. It was originally described in this
study as a separate species, and then as a subspecies of
P. empetrifolia. However, some specimens that have the
herbaceous bracteole lobe do not have a large, yellow
standard. Others do not have long, strongly recurved leaves.
There have also been collections made at Tarin Rock of
Australian Systematic Botany
175
plants that resemble this form in every way except that
their bracteoles are scarious and completely fused, or their
bracteoles are intermediate between herbaceous / large / free
and scarious / small / fused. Therefore, there appears to be
a continuum of variation which precludes making even
this distinctive form a separate taxon. Genetic evidence
(DNA sequences) from the phylogenetic component of
this study (Fig. 3) suggested only that these three
forms are closely related, and it was unable to detect
significant variation between the four P. empetrifolia
populations sampled. They are therefore considered to be
diverse conspecifics. Future studies of the geographical
patterns of morphological variation in the species may
prove interesting.
Notes
Visited by native bees (pers. obs.).
Pultenaea empetrifolia has no close relatives, but is most
closely related to the P. indira and P. ericifolia groups of taxa.
The most obvious difference between P. empetrifolia and all
relatives is that its leaves are always ≤4 mm in length, when
measured from apex to base without straightening the leaves
first. All other taxa have longer mature leaves.
The species is frequently misidentified. Table 6
lists the characters distinguishing P. empetrifolia from
morphologically similar taxa, and Fig. 5 shows the bract
morphology for the group.
Selected specimens (90 examined)
Corden Inlet NE of Bremer Bay, Aplin 2771, 31.x.1963 (AD, MEL,
NSW); 2 miles from coast, E of Fitzgerald River Inlet, Aplin 3671,
ix.1970 (PERTH); along road towards Toolbrunup, Ashby 556, 5.x.1963
(AD); 4 miles SE of Katanning, Phillips W.A./62 1938, 7.x.1962
(CANB); Reserve No. 25194, Beard 7537, 26.ix.1975 (NSW); 9 km
along Woogenellup Road from Albany–Borden road, Macfarlane 2099,
30.x.1992 (PERTH); 300 m along S Toompup Rd from Salisbury Rd,
Orthia 53, 25.x.2002 (CANB, K, MEL, PERTH); 11.1 km NW of
Boxwood Hill turnoff on South Coast Hwy, Orthia 54, 25.x.2002
(CANB, K, MEL, PERTH); 16.1 km along South Coast Hwy heading
NW from Jerramungup, Orthia 55, 25.x.2002 (CANB, K, MEL,
PERTH); 21.1 km W of Newdegate, Orthia 79, 3.xi.2002 (CANB,
PERTH); 27.5 km W of Lake Grace, Orthia 80, 3.xi.2002 (CANB,
K, MEL, PERTH); 21 miles of E Gnowangerup, Sands 638.16.7,
25.viii.1963 (AD, SYD); Chester Pass, Sands 638.15.1, 22.viii.1963
(SYD); 28 miles E of Jerramungup, Sands 638.16.12, 23.viii.1963
(AD, SYD); 27 miles from Cranbrook, Wrigley s.n., 11.x.1968
(AD, CANB).
11. Pultenaea ericifolia species complex
This group of species presents some challenges for specieslevel taxonomy, with morphology, geography and three sets of
genetic data all telling slightly different stories. Together with
P. radiata, these taxa consistently form a very well supported
monophyletic group in trnL–F, ndhF and ITS trees, but the
internal relationships within the group vary between genomic
regions (Fig. 3).
176
L. A. Orthia et al.
Australian Systematic Botany
An initial assessment of the group based on
morphology led to our recognition of four taxonomic
units: P. ericifolia, P. strobilifera, P. verruculosa and
a new taxon from the Mt Lesueur region [here called
Pultenaea sp. Mt Lesueur (Beard 7827); previously
designated ‘P. quixote Orthia’ on determinavit slips].
Table 7 and Fig. 5 outline some of the morphological
differences between the four taxa. Our investigations
suggested that P. ericifolia and P. strobilifera were
probably two subspecies of the same species, while
P. verruculosa and P. sp. Mt Lesueur (Beard
7827) were two distinct species. The morphological
differences between P. ericifolia and P. strobilifera
are relatively minor: the only consistent character for
distinguishing between them is the level of fusion
between stipules and leaf in the involucral bracts,
with those of P. ericifolia being fused for ≤ two-thirds
of their length and those of P. strobilifera fused for
≥ three-quarters of their length. Although at the extreme
ends of the range the two taxa are relatively easy to tell
apart, with very pronounced bract differences, there are
collections that are almost impossible to identify (e.g.
Ashby 2336, Drummond II 247, Pignatti 852, Woolcock s.n.,
Learmonth ACB44062). These intermediate forms suggested
that the taxa are in fact geographically and possibly
ecologically separated variants of the same species. We
therefore initially proposed to lump the two together under
P. ericifolia, as previously suggested by T. Macfarlane on
determinavit slips.
Pultenaea sp. Mt Lesueur (Beard 7827) and P. verruculosa
are more substantially different from P. ericifolia and
P. strobilifera in morphology, so we initially proposed to
circumscribe each of them as distinct species. However,
given the allopatry of P. sp. Mt Lesueur (Beard 7827) and
P. ericifolia, it is possible that P. sp. Mt Lesueur (Beard
7827) simply comprises a remnant population of P. ericifolia
that has gone through a genetic bottleneck with subsequent
change in degree of some unusual morphological characters.
The fact that many of the differences between P. sp.
Mt Lesueur (Beard 7827) and P. ericifolia are quantitative
rather than qualitative (e.g. hairiness of leaves; relative
lengths of leaves, bracts and calyces; see Table 7)
lends strength to that argument. Should environmental
circumstances bring P. ericifolia back into contact with
P. Mt Lesueur (Beard 7827), the two may recommence
interbreeding. This uncertainty about the taxonomic
distinctiveness of the Mt Lesueur form makes describing it
as a new species risky.
Pultenaea verruculosa, in contrast, possesses some traits
unique in the group that strongly suggest that it is a
separate species, for example its long, slender style and small
stigma (compared with the short thick style and bulbous
stigma of P. ericifolia, P. strobilifera, P. sp. Mt Lesueur
(Beard 7827) and P. radiata). In addition, its sympatry
with P. strobilifera and P. ericifolia, and its coexistence and
synchronous flowering with P. strobilifera at one or more
collection localities (e.g. Orthia 44 and Orthia 45), suggest
that the two are probably not interbreeding and therefore not
the same species.
However, DNA sequence data tell some very different
stories. The trnL–F sequence of P. verruculosa is identical
to one from P. strobilifera (Fig. 3), so if they are different
species, they have probably diverged only recently. Pultenaea
strobilifera itself does not form a monophyletic group, despite
the similarity of the three DNA sequences and its fairly
uniform morphology across its range. The most surprising
result of the trnL–F tree was that the six populations of
P. ericifolia split into two very well supported clades: a
‘northern’ clade, including the populations from the Perth
and Bindoon regions as well as P. sp. Mt Lesueur (Beard
7827), and a ‘southern’ clade, including the populations from
the Whicher Range, Yornup, Denmark and Mt Barker. The
ndhF and ITS trees (Fig. 3) reproduce this northern / southern
split too, although fewer populations and taxa are sampled.
These sequences suggest slightly different relationships,
uniting P. strobilifera and P. verruculosa with part of
P. ericifolia in the southern clade. Distribution maps of
the known populations of P. ericifolia and allies show
that there is a disjunction between these ‘northern’ and
‘southern’ populations of 100–150 km. It seems clear from
this evidence that the northern and southern P. ericifolia
populations have not exchanged genetic material in the
recent past, and may well represent distinct, cryptic taxa.
It is not at all clear where taxonomic boundaries should be
Table 7. Characters used to distinguish taxa in the P. ericifolia species complex
Pultenaea ericifolia s.s. and P. ericifolia ‘southern’ (Orthia 39) are grouped together here because there
are no consistent morphological differences between them
Taxon
P. ericifolia
P. sp. Mt Lesueur
P. strobilifera
P. verruculosa
Tight involucre
as high as
calyx lobes
Leaves obscure
involucre bracts
Level of fusion
between bract
leaf and stipules (%)
Upper calyx
lobes in
distinct lip
yes
no
yes
no
no
yes
no
yes
≤67
c. 67
≥75
u. 90–95
yes
yes
yes
no
Revision of Pultenaea: Western Australian species
drawn: should we split the group into five taxa [P. ericifolia
‘northern’, P. ericifolia ‘southern’ (Orthia 39), P. strobilifera,
P. verruculosa, P. sp. Mt Lesueur (Beard 7827)], maintain
the four morphologically defined units, or perhaps split the
whole group into two species—one northern [P. ericifolia
‘northern’ plus P. sp. Mt Lesueur (Beard 7827)] and one
southern [P. ericifolia ‘southern’ (Orthia 39), P. strobilifera
and P. verruculosa]?
Despite the northern / southern division being
the strongest genetic division in the group, detailed
morphological work failed to find any characters that could
consistently be used to distinguish between the northern
and southern populations of P. ericifolia. All collections
from the Perth region have massive, spectacularly showy
inflorescences and long leaves not seen elsewhere, and were
it not for the single collection from the Bindoon region,
these traits could be used to differentiate between the two
geographic forms. But the Bindoon collection has smaller
inflorescences and leaves similar to the southern populations,
rendering clade differences cryptic.
In the absence of consistent diagnostic characters, we were
hesitant to describe a new species based only on genetic
and geographic evidence. In addition, the close genetic
relationship between P. sp. Mt Lesueur (Beard 7827) and
the northern P. ericifolia populations added further doubt to
the taxonomic distinctiveness of P. sp. Mt Lesueur (Beard
7827), so we were hesitant to describe it formally under any
taxonomic rank. The genetic data throw doubt even on the
taxonomic status of P. verruculosa, given the similarity of
its DNA sequences to those of P. strobilifera and southern
P. ericifolia. It is conceivable that the chance development
of P. verruculosa’s different style / stigma morphology in an
ancestral P. verruculosa plant prevented cross-pollination
with P. ericifolia / P. strobilifera, leading to speciation.
One solution would be to describe P. verruculosa as a
species separate from the rest of the P. ericifolia complex.
This solution would leave a diverse paraphyletic residual
P. ericifolia complex, so is not very satisfactory. In addition,
some aspects of P. verruculosa’s morphology are variable
across its range (e.g. inflorescence and leaf size, flower colour,
bract shape), yet only one individual was included in our
genetic study. It is possible that it, too, is paraphyletic, and it
would be preferable to be confident of its monophyly before
treating it as a species.
Rather than proposing a falsely confident new formal
classification for these plants, the solution we have chosen
is to unite the five morphological / geographic / genetic
taxonomic units that we have identified under the P. ericifolia
‘species complex’ banner. This approach maintains the status
quo of names and ranks pending more decisive research,
while allowing the identification of collections at least to the
level of morphotype, and also indicating to future workers
where we believe that significant taxonomic boundaries lie
in the group.
Australian Systematic Botany
177
Future studies should also include P. radiata given its close
relationship to the P. ericifolia complex, and may consider the
following issues:
1. A single collection of P. strobilifera from the ‘northern’
region was made in 1902 (Andrews 198)—it might be
informative to find this population and add its DNA
sequences to phylogenetic analyses.
2. Given that only one ‘northern’ P. ericifolia collection, from
north of Bindoon, resembles the ‘southern’ morphotype
(Crisp 8451), it would be informative to find this
population or any others in the ‘northern’ region outside
of the Perth general area and to compare its morphology
to ‘southern’ plants.
3. Pultenaea verruculosa has a widespread distribution
almost identical to the combined distribution
of P. strobilifera and ‘southern’ P. ericifolia. A
comprehensive phylogeographic survey of these three
taxa is needed to determine whether they are distinct
lineages. Such a study should use more variable markers
such as AFLP to improve resolution.
4. Hybridisation experiments may also be useful, for example
to find out whether P. sp. Mt Lesueur (Beard 7827)
is in fact reproductively compatible with ‘northern’
P. ericifolia, and to test the compatibility of the ‘northern’
and ‘southern’ P. ericifolia populations.
5. Pultenaea verruculosa and P. strobilifera / P. ericifolia
‘southern’ (Orthia 39) might be two forms of the same
species, maintaining their distinctness by diversifying
selection acting at a very local level, or by very tight
linkage disequilibrium. Breeding experiments could also
test this.
6. An examination of the level of genetic variation present
within P. sp. Mt Lesueur (Beard 7827) may shed light
on the question of whether or not this taxon underwent a
bottleneck in recent history, leading to the fixation of rare
characters.
7. This group of taxa could contribute to understanding the
biogeographic relationship between broad areas within
south-west WA. There are several disjunctions present
in the current distributions, some of which (e.g. the
disjunction between northern and southern P. ericifolia)
are perhaps more ancient than others (e.g. the disjunctions
between Mt Lesueur, Bindoon and Perth, or the disjunction
between the Stirling Range / Bremer Bay area and the
Esperance / Cape Arid area).
Key to taxa in the P. ericifolia species complex
1. Inflorescence bracts form a tight, multi-layered involucre that is not
obscured by leaves
2. Inflorescence bracts fused for ≤ two-thirds of length
3. Growing north of 32◦ 30′ S............11.1. P. ericifolia s.s. (‘northern’)
3: Growing south of 33◦ 30′ S ...................................
............................................11.2. P. ericifolia ‘southern’ (Orthia 39)
178
L. A. Orthia et al.
Australian Systematic Botany
2: Inflorescence bracts fused for ≥ three quarters of length...................
.................................................................................11.3. P. strobilifera
1: Inflorescence bracts form a lax involucre or simple whorl that is partly
obscured by leaves
4. Growing south of 33◦ S parallel, style >4 mm long and tapering into
stigma ± same diameter as style............................11.4. P. verruculosa
4: Growing north of 32◦ S parallel, style <4 mm long bending sharply
into bulbous stigma much larger in diameter than style........................
....................................................11.5. P. sp. Mt Lesueur (Beard 7827)
11.1. Pultenaea ericifolia s.s.
Pultenaea ericifolia Benth., in Lindl., Edwards’s Bot. Reg.,
Appendix to Vols 1–23, A Sketch of the Vegetation of the
Swan River Colony: xiii (1839). Typus: ‘Swan River’, 1839,
Drummond I (lecto-, K (photo seen); isolecto-, BM!) here
designated.
Illustrations
Grieve, How to Know Western Australian Wildflowers 2: 420
(1998).
Erect, low shrub, 0.2–0.4 × 0.3–0.5 m. Bark grey or
reddish, thin and cracking or flaking on lower stems,
smooth on upper stems, lower stems glabrous, upper
stems sparsely hairy to velutinous, hairs erect and curly or
straight. Leaves olive to bright green, 2.1–9.5 × 0.5–0.9 mm,
length : width ratio 5–12:1, linear, spiralled, ascending
to divergent, straight to slightly recurved, venation
obscure, weakly to strongly verruculose, mature leaves
glabrous, rarely a few mature leaves on a specimen
sparsely hairy, hairs both short and long; apex acute,
blunt, base truncate, margins involute; petioles 0.3 mm,
not decurrent. Stipules red-brown, 1.5–3.5 × 0.2–0.4 mm,
not fused behind the petiole, lanceolate, straight to
recurved or twisted, glabrous, sometimes frayed, broken or
dried with age, apices acuminate. Inflorescences terminal,
indeterminate, sessile flowers a few to >30, sometimes
multiple clusters per inflorescence, inflorescence subtended
by involucre of 10 to >20 bracts; bracts red-brown, up to
9.5 × 4.5 mm, as long as calyx lobes, densely hairy, scarious;
bract leaf strongly reduced; bract stipule apices acuminate,
margins ciliate, body of bracts glabrous to densely hairy;
flower-subtending bracts similar to inflorescence-subtending
bracts but sometimes longer, narrower and more shallowly
fused. Bracteoles red-brown, 4.5–6.5 × 0.3–0.9 mm, falcate
or lanceolate, inserted at base of calyx, sparsely to densely
hairy, scarious, apices acuminate. Calyx lobes ± equal in
length, two upper lobes united into lip, tube 3.0–3.5 mm, U–U
2.0–2.5 mm, U–L 4.0–4.5 mm, L–U 3.8–4.5 mm, sparsely to
densely hairy; hairs erect, terete; lobes 0.7–1.0 mm, narrowly
triangular, apices acuminate. Standard usually orange with
dark red markings and yellow eye, 10.0–11.0 × 8.5–9.0 mm,
reniform, sometimes recurved or inwardly folded, claw
5.5–6.0 mm, apex minutely notched; wings pink or dark
red, 7.5–10.0 × 1.5 mm, held over keel, overlapping at
apex; keel usually dark red, sometimes paler pink at base,
6.5–8.0 × 1.0 mm, much smaller than wings. Anthers
0.4–0.5 mm, dorsifixed, filaments 4.5–5.0 mm. Ovary
1.4–1.7 mm, cross-section elliptic, velutinous, stipe
negligible; style 2.8–3.7 mm, straight for length with sharp
90◦ bend just below stigma, glabrous; stigma bulbous,
diameter much wider than style diameter, 0.25–0.4 mm. Pod
4.0–5.0 × 1.9–2.5 mm, elongate or broadly ovoid, sparsely
to densely hairy outside, hairs erect, terete; glabrous inside.
Seed not seen.
Phenology
Flowering September–early November; fruiting March.
Habitat and ecology
Undulating, gentle slopes to steep slopes, granite outcrops.
Diverse vegetation—open forest, woodland, open woodland,
open to closed heathlands, disturbed roadsides. Often
gravelly soils, gravel quartzite or laterite, clay loam, loam
or sandy clay. On granite outcrops (Hopper et al. 1997).
Scattered to locally common.
Distribution
Western Australia, botanical district Darling, subdistrict
Drummond. From the outer Perth region and north of Perth
(Fig. 7a).
Conservation
This taxon has been collected often from the outer Perth
region, but only once from a locality outside this area—
the Crisp 8451 collection from 27 km north of Bindoon, so
may qualify as a rare plant based on range size. No changes
to its conservation status are proposed here, but existing
populations should be monitored.
Chromosome number
Unknown. Sands (1966, 1975) counted the number at 2n = 14
for both P. ericifolia and P. strobilifera, but the vouchers she
cites for the counts belong to other taxa. Sands 638.16.19,
Sands 638.19.5 and Sands 638.19.7 are P. indira and Sands
638.14.9 is P. verruculosa.
Typification
Bentham (1839) cites no specimens in his original
description of the species. However, in his 1864 revision, he
cites Drummond I as a specimen he has seen. The Drummond
I collection is one of two mentioned in the revision that
was collected before 1840 and there is a specimen at K,
where Bentham was based. The other pre-1840 collection
is Brown 5046 (a BM collection), but geographically and
morphologically it belongs to P. ericifolia ‘southern’ (Orthia
39). The Drummond I K specimen fits better with the
P. ericifolia protologue, and is therefore designated the
lectotype, with a single isolectotype housed at BM.
Revision of Pultenaea: Western Australian species
Australian Systematic Botany
179
Fig. 7. Distribution maps of (a) P. ericifolia s.s., (b) P. ericifolia ‘southern’ (Orthia 39), (c) P. strobilifera, (d) P. verruculosa, (e) P. sp.
Mt Lesueur (Beard 7827) and ( f ) P. radiata.
Discussion
This taxon has a reasonably consistent morphology
in the outer Perth region, with leaves at the longer
end of the range and inflorescences on the larger
side. The single collection from north of Bindoon,
however, has quite a different appearance and is more
similar in gross morphology to P. ericifolia ‘southern’
(Orthia 39).
Notes
Common name: Mop pea (Hamilton 27).
A proposal to lump the taxa P. ericifolia and P. strobilifera
as subspecies of P. ericifolia has been informally put forward
180
L. A. Orthia et al.
Australian Systematic Botany
at least twice. Wheeler et al. (2002) named a taxon in their
study group P. ericifolia subsp. ericifolia, but did not state
the names of any other subspecies within P. ericifolia that
would make the autonym subsp. ericifolia. Similarly, the
Western Australian botanist T. Macfarlane, who has worked
on Pultenaea for many years, designated several specimens
of P. ericifolia and P. strobilifera as subspecies of P. ericifolia
on determinavit slips during the 1990s, but did not formally
publish the designation or his justification for lumping the
two taxa.
The commonly used spelling of the name of this species,
P. ericifolia, is here retained against the original spelling,
P. ericaefolia (International Code of Botanical Nomenclature
2000, Article 60.8, example 13).
Selected specimens (29 examined)
Ellis Brook Valley Reserve, Bowler 351, 30.ix.1996 (PERTH); Great
Northern Highway, 27 km N of Bindoon, Crisp 8451, 21.ix.1993
(CANB, PERTH); 19 km. east of Perth, edge of escarpment, Hamilton
27, 19.ix.1984 (MEL); ad fluvi Cygnorum collegit Mylne, Hance
8401, 1887 (CANB); Talbot Road Reserve, Midland, Keighery
12146, 21.ix.1991 (CANB, PERTH); To W of Ridge Hill Rd,
Gooseberry Hill, Markey 947, 18.ix.1996 (PERTH); Ellis Brook
Valley, Markey 1257, 9.x.1996 (PERTH); Perth, Kalamunda, Orthia
88, 6.xi.2002 (CANB, K, MEL, PERTH); Darling Range, Pritzel
792, x.1901 (AD-U); Lesmurdie Open Space, Purdie 3741, 23.x.1990
(CANB); Darlington, Roark s.n., 18.ix.1949 (AD-U); Kalamunda,
19 km E of Perth, Ross 2921, 8.x.1985 (MEL, PERTH); Near
Ridge Hill Road, Seabrook 606, x.1978 (PERTH); W.A., Tepper
s.n., ?1892 (NSW 517686); Hills at Darlington, White 5243,
1.xi.1927 (BRI).
11.2. Pultenaea ericifolia ‘southern’ (Orthia 39)
As for P. ericifolia s.s. except:
Open, sparse to low dense erect shrub, 0.1–1.2 ×
0.3–1.0 m. Leaf lengths and number of flowers per
inflorescence generally at the lower end of the range. Standard
petal apricot, yellow, cream-pink or salmon-pink, wing petals
magenta or salmon pink, keel petals red, black–red or
dark salmon.
Habitat and ecology
Flat areas, gentle slopes, swampy areas, granite outcrops.
Diverse vegetation—open forest, woodland, open woodland,
low woodland, tall shrubland, sedgeland, disturbed roadsides,
associated with Pultenaea radiata. Clay, loam, sand or
intermediates, often over granite, ironstone or clay, also peat
or peaty sand. On granite outcrops (Hopper et al. 1997).
Occasional to very common.
Distribution
Western Australia, botanical district Darling, subdistricts
Dale, Warren and Menzies. Collected from two slightly
disjunct areas: (1) the area south of Busselton and in the
vicinity of Manjimup, and (2) the area surrounding Denmark,
Mount Barker and Albany (Fig. 7b).
Conservation
This taxon has been collected only sporadically, often
from populations with few individuals. No changes to
its conservation status are proposed here, but existing
populations should be monitored.
Chromosome number
Unknown (see P. ericifolia s.s. discussion).
Discussion
This taxon is defined as distinct from P. ericifolia s.s.
primarily by its southern distribution and different DNA
sequences, although its general habit and the reported colours
of the petals may also often be different.
Specimens examined
Pt 5100, South Coast Highway, Annels 1815, 24.x.1991 (PERTH);
Donelly Plain, west of Manjimup, Ashby 502, 28.ix.1963 (AD); King
Georges Sound, Brown 5046, xii.1801 (BM); 870 m into car park on
Mount Barker Road, Day & Casson W166.5, 7.iii.1997 (PERTH);
Bank of the Kent River, Hammersley 2082, 31.x.1998 (PERTH);
Scott River Plain, Keighery 4067, 7.x.1981 (CANB); Smith Road,
Whicher Range, Keighery 13330, 10.ix.1992 (PERTH); Fish Road
Nature Reserve, Macfarlane 2076, 21.x.1992 (PERTH); Hithergreen,
McCutcheon 2356, 4.x.1991 (PERTH); On Sue’s Road, c. 500 m
north of Sabina East Road, Orthia 39, 20.x.2002 (CANB, K, MEL,
PERTH); South Western Highway, 7.1 km from Yornup, Orthia 41,
21.x.2002 (CANB, K, MEL, PERTH); On Mt Barker, 70 m E of car
park in scrub, Orthia 46, 23.x.2002 (CANB, K, MEL, PERTH); On
South Coast Highway, 16.6 km E of Denmark town centre, Orthia
48, 23.x.2002 (PERTH); 1.76 km along Gibsons track, Papenfus 208,
18.x.1995 (PERTH).
11.3. Pultenaea strobilifera Meisn., in Lehmann, J.G.C.,
Pl. Preiss. 1: 75 (1844). Typus: ‘In regionibus interioribus
Australiae meridionali-occidentalis’, m. Oct. Herb. Preiss.
No. 1185. (lecto-, LD (photo seen); isolecto-, NY) et in
districtu Minto, m. Nov. 1840. No. 1190. (syn-, LD), here
designated
Pultenaea pteronioides Turcz., Bull. Soc. Nat. Mosc. 26: 280–281
(1853). Typus: ‘Nova Hollandia’, n.d., Drummond V 67. (holo-, KW
(photo seen); iso-, BM!, CGE, K!, OXF!, W).
As for P. ericifolia s.s. except:
Open to dense, sprawling, domed or spindly erect shrub,
0.1–1.0 × 0.3–1.0 m. Hairs on stems erect and straight;
inflorescences up to approximately 10 flowers, rarely multiple
clusters per inflorescence; inflorescence-subtending bracts
are up to 7.5 × 3.5 mm, appressed, bract lobes fused for >75%
of length; bracteoles 5.5–6.5 × 0.5–0.9 mm, falcate; standard
apricot, yellow, salmon pink, bright pink or vermillion.
Phenology
Flowering September–early November; fruiting October.
Revision of Pultenaea: Western Australian species
Habitat and ecology
Flat plains to hilltops, including granite outcrops, coastal
hills, stable dunes and swampy areas. Woodland, low open
woodland, open heath, often tangled amongst other shrubs,
sometimes in disturbed habitats such as old gravel pits
or roadsides, associated with Pultenaea verruculosa. Sand,
sandy loam or sandy clay, often gravelly or rocky, white, grey
or brown, also in gravel and in peaty sand. Rare to locally
common, usually only a few plants.
Distribution
Western Australia, botanical districts Avon, Eyre and Darling
(subdistricts Dale and Menzies). Known from three disjunct
areas: (1) east of Esperance, in the Cape Le Grand and Cape
Arid regions, (2) in the vicinity of the Stirling Ranges to
north-west of Kojonup, and (3) a population in the vicinity
of Mogumber, approximately 100 km north of Perth, known
from a single collection (Andrews 198) (Fig. 7c).
Chromosome number
Unknown (see P. ericifolia s.s. discussion).
Typification
The LD type specimens of P. strobilifera are likely to have
been used by Meisner in describing P. strobilifera (Crisp
1983). Preiss 1185 is here designated the lectotype, because it
has a duplicate at NY. Some fragments of Drummond II 247,
one of the collections mentioned in the original description,
appear to belong to P. ericifolia, so it is not listed here as a
syntype of P. strobilifera.
Selected specimens (44 examined)
Mogumber—60 m N of Perth, Andrews 1st coll. 198, ix.1902 (BM);
c. 4.5 km SE of Mt Merivale, Archer 2409951, 24.ix.1995 (MEL);
Between South Stirling and Many Peaks, Ashby 629, 21.x.1963 (AD);
Salt River Road, adjacent Stirling Range, Ashby 2336, 28.ix.1967 (AD);
Cape Arid National Park along Tagon Rd, Corrick 9544, 25.ix.1985
(MEL); Salt River Rd 17 km west of its junction with Chester Pass
road, Corrick 9676, 17.x.1985 (CANB); Swan R., Drummond 247,
n.d. (K); Kululinup Nature Reserve, Keighery 2563, 26.x.1997 (CANB,
PERTH); Tenterden, Stirling Range, Morrison s.n., 26.ix.1902 (BM);
Patch of bush E side of Yeriminup Road, Orthia 44, 22.x.2002 (CANB,
K, MEL, PERTH); 17.0 km along Salt River Road, Orthia 52, 25.x.2002
(PERTH); Cape Le Grand NP, Gravel pit overlooking Lucky Bay, Orthia
75, 31.x.2002 (CANB, MEL, PERTH); District N. West Plantagenet,
Pritzel 687, ix.1901 (AD, BM); 116 km E of Esperance towards Cape
Arid, Taylor 2331, 28.ix.1983 (AD, CANB); Cape Le Grand National
Park, opposite Lucky Bay, Turley 1/1097, 12.x.1997 (PERTH).
Australian Systematic Botany
181
Spreading to erect, dense to open shrub, single- or
multi-stemmed, 0.2–1 × 0.2–0.5 m. Bark grey over red
wood, plated, fissured; lower stems glabrous; upper stems
yellow, smooth, velutinous, hairs erect, terete. Leaves
mid- to grey-green, (4.0–)6.3–9.7(–16.2) × 0.3–0.7 mm,
length:width ratio (6–)8–12(–26) : 1, linear, spiralled,
inclined through to reclined and occasionally ascending
or descending, usually incurved but sometimes straight
to slightly recurved, venation obscure, lamina smooth to
strongly verruculose, glabrous to sparsely hairy; hairs
long (≥1 mm), erect; apex acute, often mucronate, rarely
aristate, base truncate, margins involute; petioles 0.5 mm,
not decurrent. Stipules red-brown, 1.0–1.5 × 0.5 mm,
not fused behind petiole, lanceolate, often recurved
and twisted like dried grass, glabrous, age either by
becoming grey and dry or by splitting off, apices acuminate.
Inflorescences terminal, indeterminate, in clusters of
7–20 (occasionally >30) sessile flowers; inflorescence
bracts leaf-like or like flower bracts; flower bracts dark
red-brown, 4.0 × 2.0 mm, invagination between stipule
and leaf lobes shallow with fusion almost complete,
scarious, glabrous on body, central lobe (bract leaf) strongly
reduced and densely hairy, margins sometimes minutely
ciliate. Bracteoles pale red-brown, 3.0–3.5 × 0.6 mm,
lanceolate to falcate, inserted at base of calyx, scarious,
sparsely to densely hairy, apices acute to acuminate.
Calyx lobes ± equal in size, tube 3.0–3.5 mm, U–U
1.3–1.7 mm, U–L 2.2–2.6 mm, L–U 2.4–3.2 mm, glabrous
to sparsely hairy; hairs long, erect; lobes 0.9–1.0 mm
wide, narrowly triangular, apex acuminate. Petals rarely all
yellow; standard colour usually gold-orange at front with
red radiating veins surrounding yellow eye, back vermillion
at margins with central dark maroon or brown patch
radiating along veins to edge, 10.0–12.2 × 6.5–8.5 mm,
reniform, inwardly folded, claw 5.5–6.2 mm, apex notched;
wings usually red-pink, 8.5–10.2 × 1.5–2.0 mm, held
over keel, overlapping apically; keel dark red to purple,
7.5–9.5 × 2.0 mm, much smaller than wings. Anthers
0.6 mm, dorsifixed, filaments 5.0–5.5 mm. Ovary
1.0–1.2 mm, cross-section elliptic, velutinous, stipe
negligible; style 4.7–6.5 mm, curved, glabrous; stigma
diameter usually ± equal to style diameter, 0.15(−0.3) mm.
Pods 3.6–4.5 × 2.1–3.0 mm, ovoid, sparsely hairy outside,
hairs erect, terete; glabrous inside. Seeds dark brown, spotted
black, 2.1 × 1.4 mm, ±ovoid, arillate.
Phenology
Flowering August–October; fruiting September–January.
11.4. Pultenaea verruculosa Turcz., Bull. Soc. Nat. Mosc.
26: 278–9 (1853). Typus: ‘Swan River’, 1849, Drummond V
65 (holo-, KW (photo seen); iso-, BM!, CGE, E, K!, MEL!,
NSW!, OXF!, W)
Pultenaea verruculosa var. pilosa Benth., Fl. Austral. 2: 129 (1864),
syn.nov. Typus: ‘Cheynes Beach’, n.d., Oldfield s.n. (holo-, K!).
Habitat and ecology
From flats to gentle hills, valleys, saddles or steep mountain
slopes, granite outcrops, often in open areas which have
been disturbed such as roadsides and drains. Variable
vegetation, from low coastal heath or open scattered montane
182
Australian Systematic Botany
heath through tall shrubland to forest, associated with
Pultenaea empetrifolia and P. strobilifera. Usually gravelly
sandy soil types, including sand, sandy clay, loamy sand
and clayey sand. Also rocky areas, or areas with a lot
of bare patches. Over various parent materials, including
granite, laterite, quartzite, spongelite and soapstone. Usually
well drained, occasionally in swampy peaty soils. Rare to
locally abundant.
Distribution
Western Australia, botanical districts Eyre and Darling
(subdistricts Menzies and Dale), with outlying populations
in Roe and subdistricts Drummond and Warren. Widespread
(Fig. 7d).
Chromosome number
2n = 14 (Sands 1966, 1975). This count was based on both
haploid and diploid counts (Sands 1975).
Discussion
Bentham (1864) recognised three varieties of Turczaninow’s
species, Pultenaea verruculosa. One of those varieties,
P. verruculosa var. brachyphylla, is here reinstated at species
level as P. brachyphylla. Pultenaea verruculosa var. recurva
is treated as a nom. dub. because no type material could
be located, but it is thought to be a likely synonym of
P. empetrifolia (see Nomen Dubium section, below). The
remaining varieties, Pultenaea verruculosa var. verruculosa
and P. verruculosa var. pilosa are here merged into a
single species. The characters Bentham used to define var.
pilosa—‘sprinkled with spreading hairs’ and ‘Bracts rather
longer and more numerous’—are part of a continuum of
variation in the species, rather than representing a distinct
form, as shown by many collections made subsequent to
Bentham’s work.
The taxon is reasonably variable across its large range. In
the far north-west of the range, in the vicinity of Busselton and
Donnybrook, specimens that have very large inflorescence
heads containing over 30 flowers have been collected. These
specimens tend to have narrower, longer leaves too. A
continuum of variation in many characters prevents further
subdivision of the taxon at this stage.
Of all five taxa in the P. ericifolia species complex,
P. verruculosa is the most clearly differentiated
from a morphological perspective, and is probably
a distinct species. It is only its placement in the
phylogenetic trees in this study that makes its taxonomic
status questionable.
Selected specimens (111 examined)
Miles east of Rocky Gully on Mt Barker Road, Ashby 2009, 14.x.1966
(AD); Track to Magog, Beard 7608, 30.ix.1975 (NSW); 3 km SE
of Ironstone Gully Falls, Corrick 10693, 2.x.1991 (CANB, PERTH);
Stirling Range, saddle 3 km ESE of Donnelly Peak, Crisp 8476,
L. A. Orthia et al.
23.ix.1993 (CANB, PERTH); Branson Road, off Chester Pass Road,
Croxford 6097, 9.x.1988 (PERTH); Corner of Boyup Brook–Kojonup
Road & Orchid Valley Road, Flowers 347, 17.ix.1997 (CANB, PERTH);
On Borden Road, Macfarlane 1823, 11.x.1988 (PERTH); Patch of
bush E side of Yeriminup Road, Orthia 45, 22.x.2002 (CANB,
PERTH); Rd to Cape Riche Sands 638.4.9, 20.viii.1963, (SYD); Old
Saltbush Rd, Sands 638.15.6, 22.viii.1963 (AD, SYD); Rd to Cape
Riche, Sands 638.14.8, 20.viii.1963 (AD, SYD); Rd to Cheyne Bay,
Sands 638.14.2, 20.viii.1963 (AD, SYD); Albany to Denmark, Sands
638.12.14, 17.viii.1963 (SYD); Beckon to Chillinlup, Sands 638.14.21,
20.viii.1963 (AD, SYD); Off Sandlewood Road, Scarth-Johnson 975,
10.x.1971 (BRI).
11.5. Pultenaea sp. Mt Lesueur (Beard 7827)
Erect shrub, 0.25–0.4 m high. Lower stems glabrous, bark
grey over dark red wood, thin, fissured; upper stems red,
smooth, densely hairy, hairs short and erect. Leaves olive
green, 5.9–10.0 × 0.5–0.7 mm, length : width ratio 10–20 : 1,
linear, spiralled, appressed to divergent, straight to slightly
incurved, venation obscure, lamina verruculose, glabrous
to sparsely hairy; hairs both short and long (> 1 mm);
apex acute, mucronate, base truncate, margins involute;
petiole 0.3–0.5 mm, not decurrent. Stipules red-brown,
1.0–1.5 × 0.1–0.2 mm, not fused behind the petiole,
lanceolate, straight to slightly recurved or twisted, glabrous,
dried and frayed when old, apices acuminate. Inflorescences
terminal, indeterminate, partially obscured by leaves
appressed to inflorescence, 4–12 sessile flowers, subtended
by involucre of >10 bracts; bracts red-brown, up to
5.0 × 2.2 mm, longer than calyx tube but usually shorter
than calyx lobes, fused for approximately 67% of
length, scarious, margins ciliate, body sparsely hairy;
central lobe strongly reduced, densely hairy; bract stipule
apices acuminate; flower-subtending bracts similar to
inflorescence-subtending bracts but sometimes longer and
wider. Bracteoles light brown, 4.5 × 0.7 mm, lanceolate,
inserted at base of calyx, densely hairy, scarious, apex
acuminate. Calyx lobes ± equal in length, two upper
lobes united into lip, tube 4.0 mm, U–U 2.0 mm,
U–L 4.5 mm, L–U 4.5 mm, densely hairy, hairs long and
erect; lobes 0.6–0.7 mm wide, narrowly triangular, apices
acuminate. Standard orange-gold with central yellow eye
surrounded by broad red area with darker radiating lines
at front, back the same but eye red and yellow flecked,
12.0 × 11.0 mm, reniform, recurved or undulating, claw
6.0 mm, apex notched; wings bright pink, 9.0 × 1.8 mm,
held above keel, overlapping at apex; keel bright pink,
8.0 × 1.5 mm, much smaller than wings. Anthers 0.5 mm,
dorsifixed, filaments 4.0 mm. Ovary 2.0 mm, cross-section
elliptic, velutinous, stipe 0.2 mm; style 3.0 mm, straight for
length with sharp 90◦ bend just below stigma, glabrous;
stigma bulbous, diameter much wider than style diameter,
0.35 mm. Pods 5 × 2.5 mm, ovoid, sparsely hairy outside,
hairs erect, terete, glabrous inside. Immature seeds seen:
grey-green, ± obovate.
Revision of Pultenaea: Western Australian species
Phenology
Flowering September–November; fruiting November.
Habitat and ecology
Near crest of hill among rocks, on steep slope and at base of
slope. Species-rich low open heath or closed heath to 1.2 m
with occasional shrubs to 2.5 m. Gravelly sand, sometimes
humus rich. Locally common.
Distribution
Western Australia, botanical district Darling, subdistrict
Drummond. The vicinity of Mount Lesueur (Fig. 7e).
Conservation
The taxon has been collected on only four occasions, all
from approximately the same locality. It appears to be
very restricted in range, but the population is conserved
within Lesueur National Park. Pultenaea sp. Mt Lesueur
(Beard 7827) is recommended here for conservation risk
code 2K (Briggs and Leigh 1996) and for Priority Two
under CALM Priority Flora Codes, should it be found
to be a species or an evolutionarily significant unit by
future workers.
Discussion
Excluding the unconfirmed record of Pultenaea juniperina
at Geraldton, P. sp. Mt Lesueur (Beard 7827) distinguishes
itself by growing further north than any other Western
Australian Pultenaea.
Notes
This species has been informally identified as ‘Pultenaea
quixote Orthia’ on determinavit slips.
Specimens examined
Mount Le Sueur, Beard 7827, 16.ix.1976 (NSW); Mt Lesueur, Corrick
10573, 20.x.1989 (CANB); S of Mt Leseur, Griffin 2105, 3.ix.1979
(CANB); Mount Lesueur, Orthia 86, 5.xi.2002 (CANB, K, MEL,
PERTH); Mt Lesueur, Orthia 87, 5.xi.2002 (CANB, PERTH).
12. Pultenaea indira Orthia & Crisp, sp. nov.
A P. brachyphylla Turcz. foliis aristatis, et bracteis et stipulis
glabris vel bracteis et stipulis pilis rectis differt. Typus: Cape
Arid National Park, along Tagon Road. 3 km south of its
junction with Merivale Rd., 25.ix.1985, Corrick 9538 (holo-,
PERTH; iso-, CANB!, MEL).
Procumbent or erect, sparse or bushy shrub,
0.1–0.6 × 0.2–0.45 m. Lower stems glabrous, bark plated
or fissured and flaking off revealing red underbark;
upper stems densely hairy, hairs curly. Leaves mid-green,
(3.0–)5.3–14.5 × 0.6–1.1 mm, length : width ratio 6–15 : 1,
linear to oblanceolate, spiralled, reclined to ascending,
slightly incurved to recurved, venation obscure, lamina
smooth or verruculose, glabrous or sparsely-densely hairy;
Australian Systematic Botany
183
hairs variable; apex acute, aristate, base truncate, margins
involute; petiole 0.2–0.5 mm, not decurrent. Stipules pale
brown to red-brown, 4.0–5.3 × 0.6–1.0 mm, not fused behind
the petiole, narrowly triangular, usually straight but may
recurve at apex and twist, margins ciliate, body glabrous or
rarely sparsely to densely hairy, becoming white or grey and
fraying or chipping with age; apex acuminate. Inflorescences
terminal, indeterminate clusters of 6–12(–20) sessile
flowers, subtended by inflorescence bracts; inflorescence
bracts leaf-like or sometimes like flower-subtending bracts;
flower-subtending bracts mid-brown to deep red-brown,
4.0–6.0 × 1.5–3.7 mm, leaf strongly reduced, stipules
enlarged, scarious, stipule margins ciliate and minutely
papillose, apices frequently recurved, body of bract glabrous
(rarely sparsely hairy to velutinous), leaf glabrous to densely
hairy, bracts of inner flowers may have some medial hairs,
bracts usually hairy on adaxial side, so hairs visible when
apices recurve. Bracteoles pale brown with darker midrib,
3.2–4.5 × 0.7–1.4 mm, ovate to lanceolate-falcate, fusion
between stipules and leaf might be incomplete but leaf
material indistinguishable from stipule material, inserted at
base of calyx, sparsely to densely hairy, scarious, hairs erect
and terete, apex acute to acuminate. Calyx lobes ± even
in size, tube 2.5–4.0 mm, U–U 1.5–2.0 mm, U–L
2.7–3.0 mm, L–U 2.6–3.2 mm, tube sparsely hairy,
lobes densely hairy, hairs erect and long; lobes 0.5–1.0 mm
wide, triangular to narrowly triangular, apex acuminate.
Standard orange or yellow at front sometimes with some
black-red or maroon markings, back has black-red or
maroon patch radiating shortly to margin, eye pale yellow
front and back, 7.3–10.2 × 5.0–7.0 mm, reniform, minutely
notched, claw 3.0–5.0 mm; wings orange or yellow,
6.6–8.5 × 1.3–2.0 mm; keel black-red or maroon,
6.3–9.5 × 1.7–2.1 mm, slightly larger than wings; petal
arrangement in three dimensions unknown. Anthers
0.4–0.6 mm, dorsifixed, filaments 5.7–7.5 mm. Ovary
1.4–1.8 mm, cross-section elliptic, velutinous, sessile or
stipe 0.2 mm; style 4.5–6.4 mm, gently curving at apex,
glabrous; stigma ± equal to or slightly larger than style
diameter, 0.2 mm. Pods 4.0–6.0 × 1.5–3.0 mm, ovate,
glabrous to densely hairy outside, hairs erect, terete;
glabrous inside. Seeds light brown, speckled dark brown,
2.1 × 1.4 mm, obliquely obovate, arillate.
Phenology
Flowering August–November. Fruiting October–February.
Chromosome number
2n = 14. Sands (1966, 1975) collected P. indira subsp. indira
several times and P. indira subsp. monstrosita twice, each
time misidentifying it as some other Pultenaea species. She
used four of these collections to count chromosome numbers:
Sands 638.17.3 was used to count the chromosome number of
184
L. A. Orthia et al.
Australian Systematic Botany
Pultenaea adunca, 638.19.7 for P. strobilifera (both actually
P. indira subsp. indira), and 638.16.19 and 638.19.5 for
P. ericifolia (both actually P. indira subsp. monstrosita)
(Sands 1975). All had 2n = 14 chromosomes.
Leaf anatomy
Pultenaea indira has stomata only on the upper leaf surface,
idioblasts only on the lower side and palisade mesophyll both
sides, according to PG Turner and RPJ de Kok (unpubl. data).
Turner and de Kok obtained these data using the specimen
Cranfield 814, which they identified as P. ericifolia, but which
is actually P. indira subsp. indira.
Discussion
Pultenaea indira subsp. indira is variable in form across
its range. In particular, the far north-west populations,
in the vicinity of Newdegate and Tarin Rock, tend to
distinguish themselves from the more common southern
forms in several ways: their foliage is more lax, the leaves
are more consistently long (>7 mm), wide (>1 mm) and
divergent, the ciliate hairs on the stipules and bracts are
longer and more feathery, stipules are more often recurved
or twisted, and the stipule material forms very large plates
covering much of the stem. Some north-western plants have
inflorescences in bunches, giving the appearance of huge
inflorescences numbering 50 flowers. These north-western
forms most closely resemble subsp. pudoides, although their
leaves are glabrous and smooth as in the rest of the range of
subsp. indira.
Pultenaea indira subsp. monstrosita resembles some
populations of P. indira subsp. indira in general form, but is
easily distinguished by its leaves which are verruculose and
hairy with two types of hairs—long, erect hairs ≥0.8 mm,
and a fuzz of short hairs 0.1–0.2 mm. Sometimes the body
of its stipules and bracts are densely hairy to velutinous with
straight hairs.
Pultenaea indira subsp. pudoides is distinguished from
subsp. indira and from subsp. monstrosita primarily by its
verruculose leaves with sparsely distributed, long (≥1.0 mm),
erect leaf hairs (not short fuzzy hairs). Its geography is also
distinctive: it is currently known only from populations west
of the 118th meridian, whereas subsp. indira and subsp.
monstrosita are only known from localities east of that
meridian. Some of the westernmost populations of subsp.
indira have a few leaf hairs, indicating that there might be a
continuum of variation between the two subspecies that has
been poorly sampled, or that land clearing for agriculture has
destroyed some intermediate populations. As far as current
knowledge extends, subsp. indira and subsp. pudoides are
different enough to be classified as separate taxa. There are
no intermediate forms between subsp. monstrosita and the
other two subspecies but its gross morphology suggests it is
a conspecific localised variant.
Notes
Dried flowers stain paper bright purple after boiling (pers.
obs.). The only other taxon in the study group noted to do
this is P. brachyphylla.
Pultenaea indira is closely related to P. brachyphylla.
It differs from it primarily by having aristate rather than
mucronate leaves, and by bract and stipule indumentum:
P. indira’s stipules and bracts usually have ciliate margins
but are glabrous on the body (subsp. monstrosita sometimes
has a densely hairy to velutinous body, with straight
hairs), whereas P. brachyphylla’s stipules and bracts are
densely hairy on the body with tightly curled woolly
hairs. Pultenaea brachyphylla, like P. indira subsp. indira,
always has smooth, glabrous leaves, whereas P. indira
subsp. monstrosita and subsp. pudoides have verruculose,
hairy leaves. Pultenaea indira’s leaves are generally longer
than those of P. brachyphylla and narrower relative to
their length.
Table 6 lists some of the characters distinguishing both
subspecies of P. indira from morphologically similar taxa,
and Fig. 5 shows the bract morphology for the group.
Etymology
The name of this species honours the Australian weed
ecologist and environmental activist, Indira Narayan.
Key to subspecies of P. indira
Leaves smooth and glabrous..............................................12.1. subsp. indira
Leaves verruculose and densely hairy, some hairs short and fuzzy..................
..................................................................................12.2. subsp. monstrosita
Leaves verruculose and sparsely hairy, hairs long and erect...........................
......................................................................................12.3. subsp. pudoides
12.1. Pultenaea indira Orthia & Crisp subsp. indira
Bark plated, stems sometimes obscured by panels of
stipule material marked in yellow and red. Leaves
(3.0–)5.3–8.1(–12.0) mm
long,
length : width
ratio
6–10 : 1, slightly incurved to slightly recurved, usually
straight (rarely strongly recurved), smooth, glabrous; stipules
and bracts red-brown, glabrous on body; inflorescences
with 6–12(–20) flowers; apical shoot not observed to
continue growing while flowering; pods glabrous to sparsely
hairy on outside.
Habitat and ecology
Flat to hilly country, swale to crest; often in disturbed areas
such as roadsides, gravel pits, plantations, rehabilitated bush;
usually in open areas or clearings; found in rocky coastal areas
and on margin of salt lake. Low heath, shrubland, mallee open
woodland or mallee woodland, often dense shrub understorey.
Clay, sand, loam and intermediates, often with gravel. Moist
to well drained. Very rare to abundant; 500+ plants observed
at one locality (Mt Burdett, Craig 2234).
Revision of Pultenaea: Western Australian species
Australian Systematic Botany
185
Distribution
Distribution
Western Australia, botanical districts Eyre, Roe and
Avon. Pultenaea indira subsp. indira is widespread,
from Mount Ragged in the Cape Arid region across
to Ravensthorpe and Lake King, with more isolated
populations in the region of Hyden, Newdegate and
Tarin Rock (Fig. 6c).
Western Australia, botanical districts Eyre and Roe. In the
area roughly bounded by Ravensthorpe, Fitzgerald and Lake
King, plus one population north of Newdegate and another
north of Lake King (Fig. 6d).
Selected specimens (57 examined)
Ravensthorpe–Hopetoun Road, Albrecht 4521, 17.ix.1990 (MEL);
Bodallin, 21 km along Hockling Road, Chandler 857, 12.ix.1999
(CANB, PERTH); Mt Burdett Nature reserve, Craig 2234, 25.ix.1992
(PERTH); 60 miles W of Norseman, Cranfield 814, 23.xi.1978
(MEL, NSW, CANB); 1.6 km W of Tarin Rock Road, Crisp 9178,
14.ix.2000 (CANB); Ca. 20 km by road SSE of Ravensthorpe,
Haegi 1026, 17.ix.1976 (AD); 18.4 km S of Ravensthorpe towards
Hopetoun, Macfarlane 2115, 5.xi.1992 (CANB, PERTH); Mylies
Beach, Newbey 11146, 9.ix.1986 (PERTH); On road from Ravensthorpe
to Hopetoun, Orthia 59, 27.x.2002 (CANB, K, MEL, PERTH);
Corner of Norwood and Wittenoom Roads, Orthia 70, 30.x.2002
(PERTH); Ravensthorpe to Hopetoun, Sands 638.17.3, 25.viii.1963
(AD); 10 ml W Newdegate, Sands 638.19.7, 27.viii.1963 (AD,
SYD); 26 km W of Newdegate, Taylor 2273, 26.ix.1983 (CANB,
NSW); Mt Ragged, ca. 160 km east of Esperance, Wilson 2920,
10.ix.1964 (AD); ca. 48 km north of Esperance, Wilson 3023,
12.ix.1964 (AD).
12.2. Pultenaea indira subsp. monstrosita Orthia,
subsp. nov.
A subspecie indira et subspecie pudoide foliis pilis densis
differt. Typus: [locality abbreviated to protect taxon] W of
Lake King on hwy, measured from turnoff to Ravensthorpe,
2.xi.2002, Orthia 78 (holo-, PERTH!; iso-, CANB!,
MEL!, K!).
Bark fissured, sand may get trapped amongst leaf
hairs giving the plant a dirty appearance. Leaves
(4.0–)5.5–9.2 × 0.7–1.0 mm, length : width ratio 9–10 : 1,
slightly incurved to slightly recurved, verruculose,
(sparsely–) densely hairy, especially abaxially; hairs
erect, two types always present: long (average ≥ 0.8 mm)
and short (0.1–0.2 mm), forming a fuzz; stipules and bracts
mid-brown, glabrous to velutinous on body, hairs straight;
in bracteoles fusion between stipules and leaf might be
incomplete but leaf material indistinguishable from stipule
material; inflorescences with a few flowers; apical shoot
often continuing to grow while flowering; pods densely hairy
on outside.
Habitat and ecology
Gentle slopes, flat or undulating plains, sometimes well
exposed, sometimes adjacent to salt lake. Open mallee
woodland or shrubland over low heath. Sand, sandy clay or
loamy sand, with gravel, moderately to well drained. Usually
rare, sometimes common.
Conservation
Recommended here for conservation risk code 3K (Briggs
and Leigh 1996) and for Priority Three under CALM Priority
Flora Codes. The taxon has been collected at least ten
times, from at least eight different localities, but is often
rare at the collection sites. In 2002, an attempt was made
to find P. indira subsp. monstrosita at one of the previous
collection sites, and the site was found to have been cleared
in a long broad line, presumably for fire control. The
subspecies was not found there. Further investigation is
required to judge the health and size of other populations,
and to accurately assess whether or not the taxon is rare
or threatened.
Discussion
There is limited morphological variation within this
subspecies, but one character that does vary somewhat is
the vestiture of stipules and bracts. Some specimens have
no hairs at all on the body of stipules and bracts, where
others have completely velutinous bracts and stipules of
variable vestiture. There are also some collections (Sands
638.19.5, Sands 638.16.19) that show an intermediate
character state, with bract vestiture varying on a single plant.
In general, bracts and stipules are hairiest in the northernmost populations.
Notes
This subspecies has been informally identified as ‘Pultenaea
monstrosita Orthia’ on determinavit slips.
Etymology
The name of this subspecies refers to its monstrous
appearance: it is covered in grey hairs, appearing as fuzz.
In Latin, the word monstrosus means strange or wonderful.
Specimens examined
15 km from Ravensthorpe along road to Lake King, Barnsley 474,
10.i.1979 (CANB); ENE of Fitzgerald, Beauglehole 49302, 28.viii.1974
(CANB); Dragon Rocks Nature Reserve, Bright & Harding DS 2.29,
18.vi.1998 (PERTH); Dragon Rocks Nature Reserve, Coates 3163,
10.x.1991 (PERTH); no location given, Newbey 5060, 17.viii.1977
(PERTH); W of Lake King on hwy, Orthia 78, 2.xi.2002 (CANB,
PERTH); Jackson Rocks Nature Reserve, Papenfus 727, 30.x.1997
(PERTH); 55 ml E Jerramungup, Sands 638.16.19, 23.viii.1963 (SYD);
34 ml N Ravensthorpe, Sands 638.19.5, 27.viii.1963 (AD, SYD); W
of Ravensthorpe, Woolcock W658, 14.viii.1982 (CANB); 7 miles from
Lake King, Wrigley s.n., 6.xi.1968 (CANB).
186
L. A. Orthia et al.
Australian Systematic Botany
12.3. Pultenaea indira subsp. pudoides Orthia,
subsp. nov.
A subspecie indira et subspecie monstrosita foliis pilis sparsis
differt. Typus: [locality abbreviated to protect taxon] NE of
Arthur River townsite WA, 15.ix.2001, Obbens 26 / 01 (holo-,
PERTH!).
Bark plated, stems sometimes obscured by panels
of stipule material marked in yellow and red. Leaves
(5.0–)10.0–14.5 mm long, length : width ratio 10–15 : 1,
recurved to straight, verruculose, sparsely hairy, hairs
≥ 1.0 mm, erect; stipules and bracts pale to mid-brown,
glabrous on body; inflorescences with c. 8 flowers; apical
shoot may continue to grow while flowering.
Habitat and ecology
Upper slope of ridge. Tall heath or low scrub. Coarse loam
or sandy clay with lateritic gravel. Localised.
Distribution
Western Australia, botanical district Avon. Known from two
populations north-east of Narrogin and one population northwest of Corrigin (Fig. 6e).
Conservation
Pultenaea indira subsp. pudoides is recommended here for
conservation risk code 2K (Briggs and Leigh 1996), and
for Priority Two under CALM Priority Flora Codes. The
taxon is known from only three populations (or possibly four,
since the collection Warren 2 has an ambiguous location
description), but all of them are reserved. The subspecies
should be investigated further.
Etymology
The subspecies epithet of P. indira subsp. pudoides refers to
the hairy nature of the plant, which is like Indira Narayan’s
cat, Draupadi (Pud for short).
Specimens examined
Boolanelling Nature Reserve, Bennett BO 6.17, 3.ix.1998 (PERTH);
Boolanelling Nature Reserve, Bennett BO 6.34, 3.ix.1998 (PERTH);
Reserve A21064, Obbens 26 / 01, 15.ix.2001 (PERTH); Yilliminning
Rock, Piggott & Roe s.n., 14.ix.1992 (PERTH); 75 km N of Yilliminning
Rock, Warren 2, 8.ix.1998 (PERTH).
13. Pultenaea juniperina Labill
This species, more common in the eastern states than in
Western Australia, was treated in Part II of de Kok and West’s
(2003) revision of the genus, so is not given full treatment
here. Morphologically, the WA specimen resembles forms of
the species that grow in the Grampians region.
Pultenaea juniperina has been apparently collected from
Western Australia only once, from a location in the vicinity
of Geraldton. In the eastern states, the Grampians in Victoria
mark the western limit of the species’ distribution, and it
is restricted to latitudes south of the 35th parallel (de Kok
and West 2003). The disjunction between the Grampians
and Geraldton is great in terms of distance and climate;
it is much greater than that of P. tenuifolia (and possibly
P. vestita), which clearly traversed the region now known as
the Nullarbor Plain at some stage. A natural jump from the
Grampians to Geraldton seems exceedingly unlikely. Despite
being unlikely, it is possible that a chance dispersal event
carried the seed of the plant from Victoria to Geraldton
at some point. For this reason the species is included in
the census of Western Australian Pultenaea in this study.
However, a more likely explanation is that the specimen
was mislabelled. It is unlikely that the collection was of a
garden escapee, brought to WA by humans in the twentieth
century, since the plant is not widely cultivated (e.g. it is not
mentioned in Wrigley and Fagg 1998). The matter should
be further investigated, with the population located again if
it still exists.
Phenology
Flowering in Western Australia in August.
Distribution
In Western Australia, known from a single collection in the
vicinity of Geraldton, botanical district Irwin (Fig. 4d).
Specimen examined
Vicinity of Geraldton, west coastal Western Australia, Stewart
Aust-1024, 15.viii.1959 (AD).
14. Pultenaea ochreata Meisn., in Lehmann, J.G.C., Pl.
Preiss. 1: 75 (1844). Typus: ‘In districtu Wellington’,
5.xii.1839, Preiss. 1038 (holo-, LD (photo seen);
iso-, NY)
Illustrations
Grieve, How to Know Western Australian Wildflowers 2: 420
(1998).
Erect, slender or rounded shrub, 0.3–1.8 × 0.35–1.5 m.
Lower stems orange-red, sometimes with green spots,
weakly ribbed, glabrous; upper stems orange, smooth,
glabrous to velutinous, hairs appressed, straight. Leaves
grey-green or yellowish, veins yellow when dry,
3.0–13.5(–29) × 1.0–4.0(–7.1) mm, length : width ratio
2–5 : 1, obovate–elliptic, spiralled, inclined to ascending,
straight, reticulate venation prominent, lamina smooth when
fresh, ridged when dried, glabrous; apex acute or obtuse,
rounded or cordate, base acute, margins flat to incurved
(rarely undulate); petiole 0.8 mm, decurrent. Stipules mostly
transparent with small brown keel, 2.0–2.8 × 0.5–0.8 mm,
fused behind the petiole, ovate, semi-ochreate, glabrous,
fraying or thinning with age, apices rounded. Inflorescences
terminal, determinate, 1 or 2 ± sessile flowers; inflorescence-
Revision of Pultenaea: Western Australian species
and flower-subtending bracts brown-red going transparent,
2.5–3.0 × 1.1–2.0 mm, elliptic, glabrous to densely hairy
along medial line, scarious, apex either round or cordate.
Bracteoles like bracts but 2.0–2.5 × 0.8–1.1 mm, sometimes
ovate, inserted at base of calyx, apex acute. Calyx
lobes ± equal in size, tube 1.9–2.2 mm, U–U 1.7–2.2 mm,
U–L 2.2–3.1 mm, L–U 2.5–3.1 mm, densely hairy, hairs
appressed to ascending, straight; lobes 1.0–1.2 mm wide,
deltate, apex acute-acuminate. Standard gold at front with
small dark red-brown ring of colour around yellow eye, back
dark red brown with gold at margins and in central stripe
(rarely standard all yellow), 8.5–9.0 × 9.0–9.5 mm, round,
claw 1.5 mm, apex notched, margin slightly incurved; wings
all gold or gold with red basally, 7.0 × 2.0 mm, not touching;
keel dark red brown (rarely all yellow), 7.5 × 2.5 mm,
broader than wings, protruding through wings. Anthers
0.6 mm, dorsifixed, filaments 6.3 mm. Ovary 1.7 mm, crosssection elliptic, velutinous, sessile; style 7.0 mm, straight
to gently curved, glabrous; stigma diameter ± equal to
style diameter, 0.1 mm. Pod 6.5–10.0 × 4.5–6.5 mm, ovoid,
glabrous to densely hairy on outside, hairs erect, terete;
inside sparsely hairy, hairs erect. Seeds black, 4.0 × 2.3 mm,
ovate, arillate.
Phenology
Flowering July–October, occasionally December; fruiting
October–November.
Habitat and ecology
Plains and gentle slopes, often swampy or winter wet areas,
moist depressions, disturbed areas. Open heath, shrubland,
low open woodland, open woodland, forest. Sandy soils,
including sand, peaty sand, sandy clay, sandy loam, over
laterite. Occasional to locally abundant, sometimes local
understorey-dominant.
Distribution
Western Australia, botanical district Darling, subdistricts
Drummond, Menzies, Dale and Warren. Widespread west of
the line stretching from Mount Frankland in the south all the
way north to the Perth region (Fig. 8b).
Notes
This species is closely related to P. reticulata. It is
distinguished from P. reticulata by its blunt leaves
(P. reticulata has aristate leaves) and stipules that
encircle the stem (P. reticulata’s stipules do not encircle
the stem).
Jewel beetles observed eating petals and crawling all over
shrub in large numbers (pers. obs.). Visited by native bees,
which land on wings and push the keel down a long way
(pers. obs.).
Australian Systematic Botany
187
Selected specimens (60 examined)
Corner of Corballup and Seaton Ross Roads, Annels 3658, 15.ix.1993
(PERTH); West of Yarloop, Cranfield 470, 22.viii.1978 (CANB, MEL,
NSW); 10 km along Northern road from turnoff on Perup road, Crisp
8469, 23.ix.1993 (CANB, PERTH); 4 km NE of Cookernup, Davis
1381, 19.vi.1996 (PERTH); 19 km N along Cup Road / Northern Road
from Muirs Highway, Davis 6018, 18.ii.1998 (PERTH); Fremantle
Rd (from Forrestdale), Demarz D6872, 8.ix.1978 (CANB, PERTH);
Remnant bushland near corner of Bartram and Boronia Roads, Banjup,
Hislop 1154, 4.x.1998 (PERTH); Kirup, Koch 2055, x.1910 (MEL,
PERTH); Palgarup, N of Manjimup, Koch 2672, xi.1922 (MEL, PERTH,
MEL); Yornup State Forest, S of Yornup, Orthia 42, 21.x.2002 (CANB,
K, MEL, PERTH); Unicup Nature Reserve, Orthia 43, 22.x.2002
(CANB, K, MEL, PERTH); Manjimup, Royce 2728, 28.ix.1948 (CANB,
PERTH); Spoil dump slopes at Muja, Collie Basin, Saffrey 1780,
1.xi.1979 (CANB, PERTH); 40 km N of Walpole, Wilson 6341, 3.x.1967
(CANB, PERTH); 7 km N of Manjimup, Woolcock W2231, 2.ix.1985
(CANB, MEL).
15. Pultenaea pauciflora M.B.Scott., Kew Bull.: 378
(1914). Typus: ‘Narrogin Experiment Farm’, 31.iii. 1914,
Stoward 64 (holo-, K!; ?iso-, MEL2055266!)
Illustrations
Grieve, How to Know Western Australian Wildflowers
2: 419 (1998).
Dense, much-branched, rounded shrub 0.5–1 × 1–2 m.
Bark red, fissured, smooth or ribbed, glabrous on lower stems,
upper branchlets densely hairy with long erect hairs. Leaves
pale green, 7.7–25 × 0.6–2.0(–3) mm, length : width ratio
7–22 : 1, linear or narrowly elliptic, spiralled, divergent to
ascending, strongly incurved to straight, midvein prominent
on older leaves, lamina smooth, densely hairy when young
to glabrous when older, hairs long and erect; apex acuminate
to acute and aristate, base narrowly cuneate, tapering into
petiole, margins flat; petiole 0.6–1.0 mm, decurrent. Stipules
red-brown, 3.5–4.5 × (0.3–)0.8–1.0 mm, fused behind the
petiole, narrowly triangular, recurved, twisted, prominently
keeled, glabrous, sometimes frayed, broken, blackened
and / or not fused when older; apex acute. Inflorescences
terminal, determinate, solitary or paired flowers sessile
or on short (1 mm) pedicels, bracts leaf-like with
reduced leaf and enlarged stipules. Bracteoles red-brown,
2.0–4.5 × 0.5–1.2 mm, elliptic, inserted at base of calyx,
stand completely free from flower, sparsely hairy with long
appressed hairs, scarious, apex acute. Calyx lobes ± even,
tube 3.0–4.0 mm, U–U 2.0–2.5 mm, U–L 2.4–2.5 mm,
L–U 2.2–2.4 mm, densely hairy with long erect hairs; all
lobes 2.0 mm wide, deltate, apex acute. Standard yellow-gold
at front, pale yellow at back with reddish smudge sometimes
present at apex, occasional faint lines surround central yellow
eye, 9.0–14.2 × 8.5–14.2 mm, round, inwardly folded, claw
2–3 mm, apex notched; wings yellow-gold occasionally
smudged with pale red, 8.2–12.0 × 2.5–3.8 mm, held over
keel, do not touch; keel pale green, 8.0–11.2 × 3.0–4.8 mm,
188
L. A. Orthia et al.
Australian Systematic Botany
Fig. 8.
Distribution maps of (a) P. aspalathoides, (b) P. ochreata, (c) P. pauciflora, (d) P. pinifolia, (e) P. reticulata and ( f ) P. skinneri.
broader than wings, protrudes through wings. Anthers
0.8–1.2 mm, dorsifixed, filaments 7.2 mm. Ovary
2.3 mm, cross-section elliptic, velutinous, sessile; style
5.0–7.7 mm, curved, glabrous; stigma diameter ± equal
to style diameter, 0.1 mm. Pod pale green towards base
and reddish apically when immature, 7.5 × 5.7 mm,
obliquely ovate, slightly laterally compressed, exterior
sparsely hairy with long, erect hairs, interior sparsely
hairy with hairs erect. Seeds dark brown, 3.3 × 2.1 mm,
arillate.
Phenology
Flowering
December.
October–December;
fruiting
November–
Revision of Pultenaea: Western Australian species
Australian Systematic Botany
189
Habitat and ecology
Notes
Generally upper slopes and plateaus of undulating country,
sometimes in depression in upper drainage line. Shrubland
with few trees, open woodland, woodland, or open forest.
Clay, sandy clay, clay loam, loamy sand or loam over
laterite or granite. Rare to locally abundant, sometimes local
understorey-dominant.
Common name: Narrogin Pea (Grieve 1998).
Pultenaea pauciflora is closely related to P. pinifolia and
P. skinneri, differing from those species by its flat leaf margins
(P. pinifolia and P. skinneri have recurved leaf margins)
and green keel (P. pinifolia and P. skinneri have a brown /
orange keel).
Flowers visited by butterflies and native bees, leaves eaten
by grasshoppers (pers. obs.).
One sheet examined (Macfarlane 1628, sheet 4) contained
a specimen with a highly unusual growth form. The specimen
has the appearance of being a young plant (it has what looks
like a root but is only a few centimetres high), densely covered
in small leaves. A close examination under the microscope
revealed the dense plant to be made up of an extremely
broad (4 cm across) flattened ‘stem’ with many rows of
leaves crossing it and growing out of its apex. The stem is
(or was) apparently obtriangular or turbinate, and possibly
hollow. One explanation for this that springs to mind is
that the plant was infected by a tumour-causing organism
such as Agrobacterium. The phenomenon might be worthy
of further investigation either for management purposes
or general interest.
Plants from one location that were dissected in this study
had developmental mutations: flowers had six calyx lobes,
three keel petals and 12 stamens (Orthia 84), which might be
a sign of inbreeding depression.
Distribution
Western Australia, botanical districts Avon and Darling
(subdistrict Dale). Known from three localities near Narrogin,
Quindanning and Wandering (Fig. 8c).
Conservation
Listed as Conservation Risk Code 2E (endangered).
The only Pultenaea species to be gazetted as rare
flora on the Wildlife Conservation (Rare Flora) Notice
1999. Listed as vulnerable on the federal Environment
Protection and Biodiversity Conservation Act 1999. This
species was presumed extinct from 1914, when the second
collection was made (the first being the type), until
1984, when WA botanist T. Macfarlane recollected it
in the Narrogin area. Since then, two other populations
have been located. One population is protected within a
conservation reserve.
Typification
The sheet MEL2055266 contains a fragment that Williamson
took from one of the two 1914 K specimens. It is possible that
the fragment came from the holotype, but it is impossible to
be sure. Williamson does not specify the month of collection
of his fragment, which is the only way to distinguish
between the holotype and the other K 1914 collection.
In his revision (Williamson 1920), Williamson does not
state whether he is even aware that two K collections
exist, so his ‘kleptotype’ fragment is listed here as an
uncertain isotype.
Discussion
May hybridise with P. pinifolia (see P. pinifolia discussion).
Managers of this species have wondered whether the
different populations of P. pauciflora are taxonomically
distinct (T. Macfarlane, pers. comm.). No significant
morphological differences between populations were
observed in this study. There were trends towards different
leaf sizes in each, sometimes strikingly so, but it is unknown
how much of this variation is environmentally determined
rather than genetic. The trnL–F region of the chloroplast
chromosome was sequenced for each of the three known
populations of P. pauciflora and the sequences were almost
identical. It seems clear that there are no infraspecific
distinctions to be made for the species. Despite this, it is
possible that gene flow between populations is now limited
or extinguished because of habitat fragmentation.
Selected specimens (51 examined)
Atkins s.n., 27.x.1986 (PERTH); Cochrane 3207, 30.xii.1998 (PERTH);
Durrell 52, 3.xi.1993 (PERTH); Hort 507, 18.vi.1999 (PERTH);
Macfarlane 1628, 28.xi.1984 (PERTH); Ochtman 3856, xi.1997
(PERTH); Orthia 83, 4.xi.2002 (CANB, PERTH); Orthia 84, 4.xi.2002
(CANB, K, MEL, PERTH); Orthia 85, 4.xi.2002 (CANB, K, MEL,
PERTH); Patrick 303, 26.viii.1986 (PERTH); Robson s.n., 10.xi.1994
(PERTH); Robson s.n., 22.v.1997 (PERTH); Narrogin Experiment
Farm, Stoward 64, 31.iii.1914 (K); Narrogin Experiment Farm:
W. Australia, Stoward 64, xii.1914 (K); Pultenaea Reference Set,
Williamson s.n., n.d. (MEL 2055266).
16. Pultenaea pinifolia Meisn., in Lehmann, J.G.C.
Pl. Preiss. 2: 220 (1848). Typus: ‘Swan River’,
Drummond II 109. (holo-, LD (photo seen); iso-,
BM!, K (2 sheets)!, OXF!, W; ?iso-, K H/278/98 280!,
K H/0278/98 178!)
Illustrations
Grieve, How to Know Western Australian Wildflowers
2: 420 (1998).
Tall erect slender or much branched open or dense shrub
(0.4–)1–3 × 0.8–3 m. Bark orange-red, longitudinally ribbed,
lower stems sparsely hairy, upper branchlets velutinous,
hairs long, appressed. Leaves dark green on upper surface,
pale green below, 12.5–40 × 1.1–2.0 mm, length : width ratio
usually 20 : 1, leaves linear, alternate, ascending to inclined,
190
L. A. Orthia et al.
Australian Systematic Botany
midvein prominent and adaxially yellow, lamina smooth,
glabrous or with sparsely distributed appressed hairs on
abaxial surface; apex acute, slightly recurved, mucronate to
aristate, base obtuse, tapering into petiole, margins revolute;
petiole 0.6–0.8 mm, weakly decurrent. Stipules brown,
2.5–4.0 × 0.8 mm, fused when young but grow apart slightly
with age, obliquely ovate to linear, sometimes recurved
strongly, prominently keeled, sparsely hairy, breaking off
with age, apices acute to rounded. Inflorescences terminal,
determinate subumbellate clusters of (1–)6 to >10 flowers,
pedicels 3.5–8.0 mm; inflorescence-subtending bracts
leaf-like, bract leaf reduced and stipules enlarged; flower
bracts red-brown, 5.5 × 3 mm, clavate, scarious, possibly
caducous, sparsely hairy with grooved hairs. Bracteoles
yellow-brown, 5.5–6.5 × 1.1–1.5 mm, clavate, inserted
on petiole approximately 2.5 mm below calyx, scarious
but filmy not rigid, caducous, sparsely hairy, apex acute
to round. Calyx lobes ± evenly sized, tube 2.5–3.0 mm,
U–U 1.5–1.8 mm, U–L 2.3–2.7 mm, L–U 2.3–2.5 mm,
sparsely hairy, hairs appressed and appear to exude oily
substance; upper lobes 1.9 mm wide, deltate, apices acute;
lower lobes 1.6 mm wide, deltate, apices acute. Standard
gold-yellow with pale yellow eye surrounded by red
smudgy line at front, same at back except red smudge
radiates out into strong dark orange / red lines along
veins, 8.2 × 8.0 mm, round, margins slightly incurved,
claw 1.5 mm, apex notched; wings gold-yellow,
7.7–8.0 × 2.5–2.8 mm, meet at apex beyond keel; keel
brown / orange apically, paler basally, 8.2 × 3.0 mm,
protruding through wings, broader than wings at widest
point. Anthers 0.6 mm, dorsifixed, filaments 5 mm. Ovary
1.8 mm, cross-section elliptic, velutinous, sessile; style
7.5 mm, J-curved, glabrous; stigma diameter ± equal to
style diameter, 0.1 mm. Pods 9.5 × 5.8 mm, ovate to almost
orbicular, sparsely to densely hairy outside, hairs long and
appressed, sparsely hairy inside, hairs erect. Seeds shiny
black, 4.0 × 2.3 mm, ovate, arillate.
Phenology
Flowering September–November; fruiting December, March,
May.
Habitat and ecology
Permanently or seasonally wet areas: creek lines, riverbanks,
swamps. Low to tall open woodland, woodland or forest,
over dense sedge and shrub understorey, associated with
Pultenaea reticulata. Rich soils, including loam, sandy loam,
clay loam, rich in organic matter or litter, less commonly
found on sand with gravel, over laterite. Locally frequent to
locally abundant.
Distribution
Western Australia, botanical district Darling, subdistricts
Menzies, Warren and Drummond. Mainly concentrated in
the Whicher Range, also found in the Pemberton area and
north of Lake Yeagarup (Fig. 8d).
Conservation
Currently listed as Conservation Risk Code 2KC– (reserved
in D’Entrecasteaux National Park) and Priority Three under
CALM Priority Flora Codes (Briggs and Leigh 1996).
Leaf anatomy
Pultenaea pinifolia has stomata only on the lower leaf surface,
idioblasts on both sides of the leaf and palisade mesophyll on
the upper side only (PG Turner and RPJ de Kok, unpubl. data).
Typification
The LD specimen of P. pinifolia is likely to be the holotype
(Crisp 1983). The two questionable isotypes from K are
Drummond collections with no collection number or year
specified, so it is impossible to be certain if they are type
material or not.
Discussion
The species is reasonably uniform throughout its range with
the exception of one examined specimen. McCutcheon 918
has leaves that are sparsely hairy with long, erect hairs, leaf
margins that are flatter than is usual for P. pinifolia, leaf
apices that are strongly aristate with a long, acicular arista,
and only one flower per inflorescence. These characteristics
are reminiscent of P. pauciflora, and it is possible that
the collection represents a hybrid population. According to
Orthia et al. (2005), the two are sister species. Evidence
against the hybrid hypothesis is that the collection locality
is not close to the known populations of P. pauciflora, being
in the south end of P. pinifolia’s range, but it might be
worth investigating.
Alternatively, the collection could represent a subspecies
of P. pinifolia. In the absence of more collections it is treated
here simply as an unusual form of P. pinifolia.
Notes
Pultenaea pinifolia is closely related to P. pauciflora, differing
from that species by its recurved leaf margins (P. pauciflora
has flat leaf margins) and brown / orange keel (P. pauciflora
has a green keel), and P. skinneri, differing from that species
by its ascending to inclined leaves >10 mm long (P. skinneri’s
leaves are divergent to reflexed and <10 mm long).
Distinctive sweet scent (Macfarlane 2078). Visited by
beetles, which appeared to be eating the petals (pers. obs.).
Selected specimens (35 examined)
Annels 2956, 9.xi.1992 (PERTH); Annels 4669, 20.x.1994 (PERTH);
Bates 24287, 20.x.1990 (AD); Bright & Annels SC 169.5, 9.i.1997
(PERTH); Casson & Annels SC 29.23, 3.xii.1996 (PERTH); Corrick
10624, 29.x.1989 (CANB); Western Australia, Gilbert s.n., 1842
(BM); Hislop 707, 9.iii.1997 (PERTH); Horan 10, 30.i.1997 (PERTH);
Revision of Pultenaea: Western Australian species
Keighery 7954, 20.xi.1985 (PERTH); Macfarlane 2078, 21.x.1992
(CANB, PERTH); McCutcheon 514, 11.x.1974 (PERTH); McCutcheon
918, 25.v.1978 (PERTH); Orthia 40, 20.x.2002 (CANB, K, MEL,
PERTH); Woolcock F43, 19.ix.1985 (AD).
17. Pultenaea purpurea (Turcz.) Crisp & Orthia, comb.
nov.
Basionym: Euchilus purpureus Turcz., Bull. Soc. Nat. Mosc. 26: 276
(1853). Typus: ‘Nova Hollandia’, n.d., Drummond V 70 (holo-, KW
(photo seen); iso-, BM!, K (2 sheets)!, MEL!, NSW!, W).
Pultenaea conferta Benth. in Fl. Austral. 2: 118–119 (1864). Typus:
‘Nova Hollandia’, n.d., Drummond V 70 (holo-, K!; iso-, BM!, K!, KW,
MEL!, NSW!, W).
Prostrate shrub (rarely low spreading shrub),
0.05–0.2(–0.4) × 0.3–1.5 m, sometimes half buried in soil.
Lower stems reddish brown, glabrous, bark plated or
fissured; upper stems reddish brown, smooth, sparsely hairy
to velutinous, hairs short, erect and straight. Leaves pale grey
to mid green, 2.9–10.5 × 0.5–1.2 mm, length : width ratio
(3–)5–9(–20) : 1, oblong, arrangement spiralled but often
obscure, divergent to ascending, straight to recurved at
apex, venation not prominent, lamina smooth (rarely weakly
verruculose), glabrous to sparsely hairy, young leaves
(flower-subtending bracts) densely hairy; hairs straight,
appressed; apex mucronate or aristate, base truncate,
tapering slightly, margins revolute; petiole 0.6–1.7 mm, not
decurrent. Stipules red-brown, 2.0–5.0 × 0.2–0.6 mm, not
fused but some stipule tissue present behind the petiole,
narrowly triangular, straight, glabrous to sparsely hairy
on margins, becoming black and brittle with age, apex
acuminate. Inflorescences lax terminal indeterminate
racemes of 1–4 flowers, often on short axillary stems,
flowers solitary, axillary, pedicels 0.8–4.5(–7.8) mm, bracts
leaf-like. Bracteoles red-brown, 2.0–4.5 × 0.3–0.7 mm,
narrowly triangular or linear, inserted at base of calyx,
scarious, glabrous to sparsely hairy, apex acuminate. Two
upper calyx lobes much larger than three lower lobes, calyx
tube 1.6–1.8 mm, U–U 2.9–4.8(–6.7) mm, U–L 2.9–4.2
(–6.5) mm, L–U 2.3–2.7(–6.5) mm, upper and lower lobes
coherent for 90–100% of upper lobe length, calyx densely
hairy, hairs appressed, of medium length (∼0.5 mm);
upper lobes 1.2–1.9 mm wide, oblique obovate, apex
acute (rarely with long arista); lower lobes 0.6 mm wide,
narrow triangular, apex acuminate (rarely with very long
arista). Standard colour variable, even on one plant, all
gold-yellow front and back with red smudges lateral to eye
at front, or all red front and back with some gold-yellow
in middle of petal and towards base, or any intermediate
coloration, 6.0 × 7.4 mm, round, recurved, claw 1.5 mm,
apex minutely notched; wings all gold-yellow or yellow at
base and red at tip, 6.2 × 2.0 mm, held over keel, coherent
for length; keel all gold-yellow or yellow at base and
red at tip, 6.0 × 2.5 mm, ± same size as wings. Anthers
0.5–0.6 mm, dorsifixed, filaments 4.5 mm. Ovary 1.8 mm,
laterally flattened, densely hairy, stipe 0.4–0.5 mm; style
Australian Systematic Botany
191
4.0 mm, hooked, glabrous; stigma diameter ± equal to style
diameter, 0.1 mm. Pods 4.5–4.7 × 2.8–3.2 mm, broadly and
obliquely ovate or elliptic, laterally flattened, sparsely hairy
outside with short, appressed hairs, glabrous inside. Seed
not seen.
Phenology
Flowering August–January;
November.
fruiting
in
August
and
Habitat and ecology
Flat areas, slight depressions and gentle slopes in flat or
undulating areas. Dense young mallee woodland or open
mallee woodland with scrub / heath understorey, often in
open or disturbed areas such as drains and road edges, seen
regenerating after fire, associated with Pultenaea spinulosa,
P. neurocalyx and P. elachista. Diverse soils, on sand,
clay, loam, sandy loam, clay loam or sandy clay, often
with gravel, sometimes with limestone or laterite. Rare
to abundant.
Distribution
Western Australia, botanical districts Roe, Eyre and
Coolgardie. Widespread, in the area roughly bounded by
Newdegate, Norseman and Cape Arid (Fig. 9f ).
Leaf anatomy
Pultenaea purpurea has stomata only on the lower leaf
surface, idioblasts on both sides of the leaf and palisade
mesophyll only on the upper leaf surface (PG Turner and
RPJ de Kok, unpubl. data).
Typification
The description of the species Euchilus purpureus published
by Turczaninow (1853) does not conform to the type
specimen nominated by him in several respects: the
description says phyllotaxis is opposite (it is spiralled), leaves
are involute (they are revolute) and all petals are purple
(they are yellow or red). Despite this, the type is specified
as Drummond V no. 70, and because Turczaninow was
working at KW, it is presumed that the KW duplicate of this
collection is the holotype. A photograph of this presumed
holotype clearly indicates that the specimen matches other
duplicates of Drummond V no. 70, which have a spirally
arranged phyllotaxis, revolute leaves and yellow or red petals.
Despite Turczaninow’s erroneous description, it is clear that
his Euchilus purpureus is the same taxon as Bentham’s
Pultenaea conferta. The fact that Turczaninow classified the
species in the genus Euchilus suggests that his assignment of
the epithet purpurea to the type Drummond V no. 70 was not
an error. Examination of the Drummond collection clearly
indicates its affiliations with the other species Turczaninow
classified as members of Euchilus. All have the distinctive
trait of enlarged upper calyx lobes, a trait mentioned in
192
L. A. Orthia et al.
Australian Systematic Botany
Fig. 9. Distribution maps of (a) P. arida, (b) P. calycina subsp. calycina, (c) P. calycina subsp. proxena, (d) P. daena, (e) P. elachista in WA and
( f ) P. purpurea.
Turczaninow’s description: ‘labio superiore parum longiore
et multo latiore. . .’ (upper lip slightly longer and much
broader [than lower lip]). Turczaninow also describes stipules
of the species as ‘stipulis lineari – setaceis petiolum
superantibus’ (stipules linear to bristle-like overtopping
the petiole), a trait characteristic of the Drummond V
no. 70 specimens. Therefore, we here reinstate Turczaninow’s
species epithet.
Discussion
Pultenaea purpurea is quite consistent in morphology across
its range. One of the rare forms has aristate leaves and
Revision of Pultenaea: Western Australian species
calyx lobes, sometimes with very long filiform stipules
and bracteoles: this form has been collected from the
Mount Buraminya area in the far east of the range and
also in the Coolgardie district in the far north of the
range. These collections also tend to have longer leaves
than average. However, some plants well within the main
part of the range (e.g. Orthia 69) also have aristate leaves
and calyx lobes, and others have relatively long leaves
(Corrick 8798, Macfarlane 2121). The form is therefore
considered to be within the normal range of variation of
the species, and has not been given separate taxonomic
status, even though it has been recognised by some workers
(on determinavit slips) as a separate taxon (‘aff. conferta’).
Some collections (Bruhn 2 / 221193, Lepschi 3827) show
plants with very small leaves (at the 3–5 mm length end of
the scale), and these have also been recognised by some
workers (on determinavit slips) as possibly separate taxa
(also called ‘aff. conferta’). However, there are plants that
show an intermediate leaf length (e.g. Wrigley WA / 68 5088,
Wrigley WA / 68 5332), so again, this form is considered
to be a localised variant of the species rather than a
separate taxon.
The erect habit is unusual for the species, and seems to
have been observed only in the Mount Heywood area.
Notes
Pultenaea purpurea is related to P. arida, P. calycina,
P. daena, P. elachista, P. quaerita, P. rotundifolia, P. spinulosa
and P. wudjariensis. It is distinguished from all of
them by its spirally arranged leaves and tightly revolute
leaf margins.
Selected specimens (82 examined)
10 km N of Mt Heywood, Archer 1211954, 12.xi.1995 (CANB,
PERTH); Kau Rock Road, 14 km S of Mt Ney, Archer 3112952,
31.xii.1995 (MEL, PERTH); 29 km ESE of Mt Heywood, Archer
1711905, 17.ix.1990 (MEL); Oldfield 1343, Bruhn 2/221193,
22.xi.1993 (PERTH); 20 km E of Ravensthorpe on Highway 1, Corrick
8798, 18.x.1983 (CANB, PERTH); 2 km SE of Mt Buraminya, Corrick
11485, 3.xi.1996 (MEL, PERTH); Mount Ney Road, Hislop 1249,
20.xi.1998 (PERTH); 3.7 km NNW of Rollands Road on Fields Road,
Lepschi & Fuhrer 3827, 30.x.1997 (CANB, PERTH); On South Coast
Hwy, 19.4 km E of Ravensthorpe, Orthia 60, 27.x.2002 (CANB, K,
MEL, PERTH); 2.7 km N of Truslove Rd on Esperance–Norseman Hwy,
Orthia 65, 30.x.2002 (CANB, K, MEL, PERTH); On Norwood Rd,
3.5 km W of Wittenoom Road, Orthia 69, 30.x.2002 (CANB, PERTH);
c. 82 km E of Norseman, Pullen 9600, 23.xi.1974 (CANB, NSW); 20 km
E of Ravensthorpe, Woolcock W750, 19.ix.1982 (MEL, PERTH); Near
Mt. Desmond, Wrigley WA / 68 5088, 27.x.1968 (CANB);
75.2 km from Esperance, Wrigley WA / 68 5332, 2.xi.1968
(CANB, PERTH).
18. Pultenaea quaerita Orthia, nom. nov.
Pultenaea obcordata (R.Br.) Benth., Fl. Austral. 2: 120–121
(1864), nom. illeg. non Pultenaea obcordata Andrews, Bot.
Repos. 9: 574 (1809). Euchilus obcordatus R.Br., Hort. Kew. 2:
17 (1811). Typus: ‘King George’s Sound’, 1801, Brown 4089
Australian Systematic Botany
193
(lecto-, BM!; isolecto-, CANB!, DBN, E (2 sheets), K (2 sheets)!, MEL)
here designated.
Pultenaea heterochila F.Muell., Fragm. 4: 21 (1863). nom. inval.
Illustrations
Grieve, How to Know Western Australian Wildflowers 2: 422
(1998).
Erect, decumbent or prostrate shrub, dense, domed or
slender, 0.2–1.8 × 0.2–2 m. Lower stems glabrous, bark
grey, knobbly and sometimes fissured; upper stems grey,
smooth, velutinous, hairs short and erect (forms north
of Ravensthorpe sparsely hairy, hairs tightly curled).
Leaves grey-green to mid-green, concolorous or brighter
on adaxial surface, 2.8–11.0 × 1.8–8.2 mm, length : width
ratio 1–2 : 1, obovate, decussate, divergent to ascending,
mid-vein prominent, often reticulate veins faintly visible,
lamina smooth, glabrous to densely hairy, sometimes rather
succulent; hairs appressed to ascending, terete; apex obtuse,
emarginate or rounded or flat, base obtuse to acute, margins
flat or slightly recurved or incurved; petiole 0.5–1.5 mm, not
decurrent. Stipules dark red-brown, 0.7–1.9 × 0.3–0.6 mm,
not fused but some stipule material present behind the
petiole, triangular, narrowly triangular or lanceolate, straight
to recurved, glabrous to densely hairy, going black, thick
and rigid or breaking off with age, apices acute if intact.
Inflorescences in upper leaf axils or in terminal racemes,
indeterminate, solitary, pedicels 0.5–4.8 mm, inflorescencesubtending bract leaves often reduced or apparently
missing, flower bracts leaf-like. Bracteoles red-brown,
1.0–2.2 × 0.5–1.3 mm, narrowly triangular, lanceolate or
ovate, inserted at base of calyx, sometimes appressed or
divergent, glabrous to densely hairy, scarious, apices acute
to acuminate. Two upper calyx lobes much larger than
three lower lobes, tube 1.0–2.0 mm, U–U 3.0–5.2 mm,
U–L 3.2–5.2 mm, L–U 1.6–3.2 mm, upper and lower lobes
coherent for 40–67% of upper lobe length, glabrous to
densely hairy, hairs terete, erect / long or appressed / short;
upper lobes 2.4–4.4 mm wide, elliptic, sometimes almost
circular, apex obtuse with weak mucro; lower lobes
0.4–0.7 mm wide, narrow triangular, apex acuminate.
Standard usually yellow-gold at front with basal radiating
red patch surrounding yellow eye, may have red margin
and red patches lateral to eye, back yellow-gold with
pale to bright red patches either side of central yellowgold area, 7.0–8.5 × 8.7–9.7 mm, reniform, slightly inwardly
folded, claw 2.0–2.2 mm, apex notched; wings yellowgold grading to pale red at base, may have red margins
at apex, 6.3–7.0 × 1.8–2.5 mm, not touching; keel red or
brown-red, 6.3–7.5 × 2.0–2.5 mm, ± same size as wings,
protruding through wings. Anthers 0.3–0.5 mm, dorsifixed,
filaments 4.4–5.2 mm. Ovary 1.8–2.0 mm, laterally flattened,
velutinous or glabrous on body with row of hairs along top
and bottom, stipe 0.5–0.6 mm; style 3.6–3.8 mm, curved or
194
L. A. Orthia et al.
Australian Systematic Botany
hooked, glabrous; stigma diameter ± equal to style diameter,
0.1–0.2 mm. Pods 5.0–6.6 × 2.0–3.7 mm, obliquely ovate,
flat, glabrous to densely hairy outside, may have a row of
hairs along top and bottom with body glabrous, hairs erect,
terete; glabrous inside. Seeds pale brown or ochre brown
sometimes with black spots, 1.4–2.3 × 1.2–1.4, round or
ovate, arillate.
Phenology
Flowering May–November, occasionally February; fruiting
September–December.
Habitat and ecology
Varying landforms including plains, dunes, cliffs and
roadsides, often on limestone outcrops or on granite outcrops
(Hopper et al. 1997), in exposed or sheltered conditions.
Open heath, shrubland, mallee open woodland or mallee
woodland. Usually shallow sand over limestone, often rocky
or gravelly, rarely clay, clayey sand, sandy loam or over
granite. Rare to abundant, often locally common.
leaves (average length ∼3.5 mm). These smaller leafed plants
also tend to have smaller and less hairy calyces than the
western populations. However, numerous collections have
been made which indicate that leaf size, calyx size and
calyx hairiness vary more or less continuously (Orthia 73,
Davis 9201, Sands 638.18.4). Populations around Mt Ragged
tend to have small leaves that are narrow (length : width ratio
2 : 1) and somewhat succulent, but again, the variation is
within the range of the species generally. Populations east of
Esperance frequently have narrow leaves, and plants in the far
east of the range also have leaves tending towards succulence.
This wide range of variation precludes any infraspecific
subdivision, and simply indicates that it is a variable
species adapted to diverse habitats. trnL–F sequences from
both the large-leafed coastal form of P. quaerita and
the narrow-leafed Cape Arid form were compared in the
phylogenetic component of this study, and were found
to be very similar to each other and to both subspecies
of P. calycina.
Notes
Distribution
Western Australia, botanical districts Eyre, Roe and Darling
(subdistricts Menzies and Warren), as well as near-coastal
areas of the South Western Interzone (Coolgardie district) and
on the western coastal edge of the Nullarbor region (Eucla
district). Mainly coastal, with scattered inland populations in
the Cape Arid region and north of Ravensthorpe. Also found
on Middle Island and Observatory Island in the Recherche
Archipelago (Fig. 10a).
Chromosome number
2n = 18 (Sands 1966, 1975). The count was based on both
haploid and diploid chromosome counts (Sands 1975).
Typification
Brown does not mention any collections in the protologue,
simply stating that the species is a ‘Nat. of the South-west
coast of New Holland’, so a lectotype should be selected for
his name. There are two collections that are good candidates:
(1) the 1801 collection made by Brown himself from King
George’s Sound, and (2) collections made from plants grown
from seed at Kew Gardens, seed which Brown supplied. The
1801 collection is chosen as the type because it is certain that
Brown saw it, with the BM specimen chosen as lectotype.
Discussion
This species is variable in aspects of leaf, ovary and hair
morphology as well as habit. The most strikingly variable
element is leaf size. In the west of the range (Eyre, Menzies
and Warren botanical districts), the leaves are relatively large
(average length ∼8 mm), but in the north and east of the
range (Roe, Coolgardie and Helms), both inland and along
the coast, there are populations which have much smaller
Flowers have a sweet smell (MacMahon 19).
Genetic evidence (Fig. 3), as well as many aspects
of morphology, suggest P. quaerita is closely related to
P. calycina. It differs from P. calycina in leaf arrangement
(decussate; P. calycina’s leaves are usually whorled) and leaf
length : width ratio (≤2 mm; P. calycina’s ratio is ≥4 mm).
Table 5 lists the characters distinguishing P. quaerita from
morphologically similar taxa.
Etymology
The new species epithet, quaerita (based on
L. quaeritare = to explore), refers to the adventurous
and ‘curious’ nature of the species: it has a wide distribution,
spanning not only several phytogeographic regions, but
also a stretch of sea, having taken up a life on at least two
of the islands in the Recherche Archipelago.
Selected specimens (133 examined)
1.6 km SW of Pink Lake—11 mile beach road junction, Chinnock
3305, 9.x.1976 (AD, PERTH); Cape Arid National Park, along
Tagon Rd, Corrick 9537, 25.ix.1985 (CANB); 3.5 km S of Lake
Gore, Cranfield 10511, 6.x.1995 (MEL, PERTH); 20.6 km S of
Cocklebiddy, Davis 9201, 3.vi.2000 (PERTH); 41 Whalebay Drive,
MacMahon 19, 30.viii.2000 (PERTH); 8 miles (12 km) north of
Israelite Bay, Nelson ANU16492, 13.ix.1972 (CANB, PERTH);
Coastal sand dunes, ca 14 km east of the mouth of the Oldfield
river, Orchard 1481, 12.x.1968 (AD, CANB, PERTH); Track to
Cave Point Lighthouse, Orthia 49, 23.x.2002 (CANB, K, MEL,
PERTH); Parmango Rd, Orthia 73, 30.x.2002 (CANB, K, MEL,
PERTH); 1 km north of Hopetoun, Ross 3022, 16.x.1985
(CANB, PERTH); Frank Hann National Park, Royce 10236,
10.xii.1971 (PERTH); Ravensthorpe to Hopetown, Sands 638.17.6,
25.viii.1963 (PERTH, SYD); Ravensthorpe to Esperance, Sands
638.18.4, 25.viii.1963 (AD, PERTH, SYD); Cape Arid Nat. Park,
7 km from Mt Ragged towards Esperance, Taylor 1569, 9.ix.1983
Revision of Pultenaea: Western Australian species
Fig. 10.
Australian Systematic Botany
195
Distribution maps of (a) P. quaerita, (b) P. rotundifolia, (c) P. spinulosa and (d ) P. wudjariensis.
(AD, CANB); Middle Island, Recherche Archipelago, Weston 8769,
17.xi.1973 (PERTH).
19. Pultenaea radiata H.B.Will., Proc. Roy. Soc. Victoria
33: 137 (1921). Typus: ‘In National Herbarium, Vic., from
Busselton, W.A., 1870, A. and E. Pries, among specimens of
P. verruculosa, var. pilosa.’ (lecto-, MEL35273!; isolecto-,
BM!, MEL (4 sheets)!, NSW!, K!; ?isolecto-, K H 278/98
88!), here designated
Illustrations
Grieve, How to Know Western Australian Wildflowers 2: 421
(1998).
Erect, multi-branched, dense, open or wiry shrub,
0.15–0.5 × 0.15–0.4 m. Bark flaking or plated, epidermis
with a metallic sheen above a reddish surface; glabrous on
lower stems, upper stems sparsely hairy with short erect
hairs. Leaves light to dark green, 6–11(–16) × 0.5–1.0 mm,
length : width ratio 15–20 : 1, linear, spiralled, ascending
through to descending, incurved, venation obscure, lamina
verruculose, sparsely hairy, hairs long and erect; apex
acute, base truncate, margins involute; petiole 0.3–0.5 mm,
not decurrent. Stipules red-brown, 1.2–1.3 × 0.2 mm, not
fused behind the petiole, lanceolate, recurved, glabrous,
becoming transparent / white, frayed and thinned like dried
grass with age, apices acuminate. Inflorescences terminal,
indeterminate, 2–11 sessile flowers per cluster, rarely
inflorescences of only a few flowers appear in leaf axils
further down the stem; inflorescence bracts leaf-like; flowersubtending bracts red-brown, 2.6 × 1.5 mm, central lobe
reduced to short linear protrusion, stipules fused for most
of length, hairs on bract leaf and at base, scarious. Bracteoles
red-brown, 4 × 2.5 mm, enclosing entire tube of calyx and
part of lobes, central lobe much reduced, obovate, hairs
on margins, midline and at base, scarious, apex rounded.
Calyx lobes ± equal in length, tube 2.5 mm, U–U 1.5 mm,
U–L 2.7 mm, L–U 2.4 mm, long hairs on lobe margins,
short sparse hairs elsewhere on calyx; lobes 0.8 mm wide,
triangular to narrowly triangular, apices acuminate. Petals
brightly coloured or gaudy, standard deep yellow to orange
apically grading to pink or purple at base, with central
196
L. A. Orthia et al.
Australian Systematic Botany
yellow eye flanked by lateral pale pink patches at front, back
has dark red striations radiating outward from flecked redyellow centre, 10.5–12.3 × 8.0–9.0 mm, narrow reniform,
slightly inwardly folded, claw 5.0 mm, apex notched; wings
pink to purple, 9.0 × 1.9 mm, held over keel, overlapping
apically; keel red at apex grading to pink or purple at
base, 5.7 × 1.5 mm, much smaller than wings. Anthers
0.4 mm, dorsifixed, filaments 4.5 mm. Ovary 0.8–1.2 mm,
cross-section elliptic, velutinous, stipe negligible; style
2.8 mm, straight with sudden bend at stigma, glabrous;
stigma much larger in diameter than style, 0.4 mm. Pods
5.5 × 2.2 mm, ovoid, sparsely hairy outside, hairs erect,
terete; glabrous inside. Seeds light- to dark-brown, may
have black spotty markings, 2.0 × 1.0–1.3 mm, roughly
elliptic, arillate.
fragment on Williamson’s ‘Pultenaea Reference Set’, so all
are considered to be from the same collection, as are all sheets
in other herbaria bearing the same or similar description
label. One specimen housed at K, which has no similar
label, is considered here to be a probable part of the same
collection pending evidence to the contrary, because it is
noted that the specimen was sent to K from MEL, possibly
by Williamson. Of the five MEL sheets, Williamson did not
specify a holotype, so a lectotype should be selected. The
sheet MEL35273 is here selected as the lectotype, because it
has the words ‘new species’ written across it in Williamson’s
handwriting in big letters.
Notes
Flowering September–October; fruiting January.
Pultenaea radiata is closely related to the P. ericifolia species
complex (Fig. 3). Table 6 lists the characters distinguishing
P. radiata from related and morphologically similar taxa, and
Fig. 5 shows the bract morphology for the group.
Habitat and ecology
Selected specimens (42 examined)
Flat plains to gentle slopes, ridges. Forest, open woodland,
low woodland or shrubland with diverse shrub and herb
understorey, often in open areas, including at edge of stream,
associated with Pultenaea brachytropis and P. ericifolia.
Found growing in vegetation regenerating after fire. Gravelly
sand, loam or clay soil over laterite, including regenerating
gravel pit and bare or littered areas. Occasional to
locally abundant, can be a major component of the local
sub-shrub layer.
18 km south-east of Busselton, Corrick 9218, 10.x.1984 (CANB);
Whicher Range, Corrick 10629, 29.x.1989 (CANB); 18 km SW of
Busselton, Cranfield 8338, 26.ix.1992 (PERTH); Whicher Range,
Davis 234, 21.ix.1995 (PERTH); Near Busselton, Fairall 2550,
14.ix.1969 (PERTH); Sabina Road, Keighery 5335, 6.x.1982 (PERTH);
Whicher Nature Reserve, McCutcheon 2617, 22.x.1992 (PERTH); On
Vasse Highway, Orthia 38, 20.x.2002 (CANB, K, MEL, PERTH);
Jarrah forest ∼10 km SE of Busselton, Paijmans 3615, 13.ix.1980
(CANB); 2 km W of Evans road and Claymore road intersection,
Papenfus 210, 18.x.1995 (AD, PERTH); WSW of Busselton, Papenfus
220, 20.x.1995 (CANB, PERTH); SW of Busselton, Papenfus 222,
20.x.1995 (AD, PERTH); 1.5 km S along Sabina road, Papenfus 400,
8.x.1995 (MEL, PERTH); 22.6 km south-east of Busselton, Ross 2972,
13.x.1985 (CANB); Sabina Rd., Whicher Range, Woolcock W2235,
7.ix.1985 (CANB, MEL).
Phenology
Distribution
Western Australia, botanical district Darling, subdistrict
Drummond. Confined to the Whicher Range area south of
Busselton (Fig. 7f ).
Conservation
Listed as conservation risk code 2K (Briggs and Leigh 1996);
Briggs and Leigh state it is listed as Priority One under
CALM Priority Flora Codes, but it was recently removed
from the priority list after survey work showed it wasn’t in
need of special protection (J. Chappill, pers. comm.). The
species has a very restricted range but has been collected from
large populations.
Leaf anatomy
Pultenaea radiata has stomata only on the upper leaf surface,
idioblasts on both sides of the leaf and palisade mesophyll
on the lower side only (PG Turner and RPJ de Kok,
unpubl. data).
Typification
H.B. Williamson, who first described this species, worked at
the MEL herbarium. There are four sheets in MEL containing
specimens collected by A. & E. Pries in Busselton, plus a
20. Pultenaea reticulata (Sm.) Benth., Fl. Austral. 2:
119–120 (1864). Daviesia reticulata Sm., Trans. Linn. Soc.
9: 256 (1808). Jacksonia reticulata (Sm.) DC., Prod. 2: 107
(1825). Typus: ‘King George’s Sound, New Holland’, 1792,
Menzies s.n. (holo-, LINN (photo seen); iso-, BM!)
Pultenaea aciphylla Benth., in de Hügel, C.L.B., Enum. Pl.
Hueg.: 35 (1837). Typus: ‘King Georges Sound’, Hügel (holo-, W
(photo seen)).
Pultenaea aciphylla var. latifolia Meisn., in Lehmann, J.G.C., Pl.
Preiss. 1: 74 (1844). Typus: ‘In solo sublimoso distr. Sussex’, m. Dec.
1839. Herb. Preiss. No. 1044. (holo-, LD (photo seen)).
Illustrations
Grieve, How to Know Western Australian Wildflowers 2: 419
(1998).
Dense, erect, rounded or spreading shrub, much branched,
0.3–3(–5) × 0.5–2 m. Bark orange, sometimes with green
spots, ribbed; lower stems glabrous, upper stems densely
hairy to velutinous, hairs appressed. Leaves grey-green
to yellowish green, veins yellow when dried,
Revision of Pultenaea: Western Australian species
6.0–33.5 × 1.2–10.4 mm, length : width ratio 2–10 : 1,
usually elliptic, sometimes obovate or ovate, spiralled,
appressed to divergent, straight (rarely slightly recurved),
reticulate venation prominent, lamina smooth when fresh,
ridged when dried, glabrous to sparsely hairy; hairs long,
erect; apex acute to acuminate, aristate, base acute, rounded,
margins flat to incurved or undulate; petiole 1.0–2.0 mm,
decurrent. Stipules red-brown, 0.7–1.5 × 0.3–1.0 mm,
fused behind the petiole, deltate to narrowly triangular,
straight to recurved, glabrous, blackening with age; apex
acuminate. Inflorescences terminal, determinate, 1–6(–12)
sessile flowers, flower-subtending bracts red-brown,
2.5–6.0 × 2.0–4.5 mm, elliptic, glabrous to densely hairy,
usually with hairs only on medial line, scarious; apex
rounded, sometimes aristate. Bracteoles red-brown,
1.2–1.5(–5.0) × 0.8–1.5 mm, ovate, inserted at base of calyx,
medially hairy, scarious, apex rounded to acute. Calyx
lobes ± equal in size, tube 2.5–3.0 mm, U–U 1.9–5.0 mm,
U–L 3.1–7.0 mm, L–U 2.9–6.7 mm, sparsely hairy to
velutinous, hairs appressed, short and grooved or long
and terete; lobes 1.2–2.0 mm wide, narrowly triangular,
apices acuminate. Standard yellow at front with pale
reddish smudgy line around central yellow eye, back
yellow in central stripe and at margins, with dark red
smudge radiating along veins either side of central stripe,
10.5–12.0 × 10.5–11.0 mm, round, margin slightly incurved,
claw 3.0–3.2 mm, apex rounded; wings yellow grading
to red stripe at base, 9.3–10.0 × 2.3–2.4 mm, do not meet
each other; keel dark red, 9.3–10.0 × 3.5–3.7 mm, broader
than wing, protruding through wings. Anthers 0.8–1.0 mm,
dorsifixed, filaments 9.0 mm. Ovary 1.8 mm, cross-section
elliptic, sessile, velutinous; style 7.5–8.0 mm, curved,
glabrous; stigma diameter ± equal to style diameter, 0.1 mm.
Pod 8.0–11.0 × 5.0–7.0 mm, ovoid, sparsely to densely hairy
outside, hairs appressed or erect, sparsely hairy inside, hairs
erect. Seeds dark brown to black, 3.4–4.3 × 2.0–2.2 mm,
obovate, arillate.
Australian Systematic Botany
197
Distribution
Western Australia, botanical districts Darling (all
subdistricts), and possibly Irwin. From Albany to Manjimup
extending south to the coast, and north of Manjimup almost
to the Mt Lesueur area. Collected more often than any other
WA Pultenaea species (Fig. 8e).
Chromosome number
2n = 8 (Sands 1966, 1975). Sands’ count was based
on haploid and diploid chromosome counts from seven
different collections (Sands 1975). One of her specimens
(Sands 638.13.5), housed at K, is annotated with the
words ‘2n = 16’. Sands (1966, 1975) cites neither the
specimen nor the 2n = 16 chromosome count though,
without explanation.
Leaf anatomy
Pultenaea reticulata has stomata on both leaf surfaces, a
large number of idioblasts associated with vasculature on both
sides of the leaf and no palisade mesophyll (PG Turner and
RPJ de Kok, unpubl. data).
Discussion
This species is quite variable in aspects of leaf size and
length : width ratio and the size of flowers and flower organs.
In general, plants with broader, undulate leaves tend to
have much bigger flowers than plants with slender, straight
leaves, but there is certainly a continuum of variation in
these traits that prevents any infraspecific taxa from being
proposed here.
Notes
Phenology
Pultenaea reticulata is closely related to P. ochreata
and P. aspalathoides. It is differentiated from the
former by its aristate (v. blunt) leaves and from the
latter by its prominent reticulate (v. prominent pinnate)
leaf venation.
Flowering August–December; fruiting August–December,
occasionally February.
Selected specimens (233 examined)
Habitat and ecology
Often found in swamps, permanent drainage lines, seasonally
wet areas, dune swales, but also found on moderate to steep
slopes, ridges and dunes. Diverse vegetation types, including
forest, open forest, woodland, open woodland, shrubland,
heath, shrub, heath or sedge understorey, vegetation dense or
open, area sometimes disturbed; associated with Pultenaea
pinifolia. Diverse soils, usually sandy or peaty, including
sand, peat, sand-loam-clay intermediates, often gravelly, over
granite, limestone or laterite. Rare to abundant.
Denmark, Beard 7745, 20.x.1975 (NSW); 0.5 km along old forestry
track on N side of road, c. 15 km along Redmond West Road, Chandler
953, 21.ix.1999 (CANB); Lake Sepping, Albany, Cranfield 4942,
30.ix.1984 (CANB); 5 km along Quaranup road from Frenchman
Bay road, Jobson 2556, 20.x.1993 (MEL); On South Coast Highway,
c. 30 km W of Denmark, Orthia 47, 23.x.2002 (CANB, K, MEL,
PERTH); Near Whaling Station, Reid 1075, 25.x.1974 (MEL); ca 5.8 km
NNW of West Cape Howe, Rodd 4976, 14.xi.1985 (CANB, MEL,
PERTH); Dunsborough, Sands 638.11.1, 16.viii.1963 (AD, PERTH,
SYD); Yallingup, S. of siding road, Sands 638.11.3, 16.viii.1963
(SYD); S Witchcliffe turn off, Busselton to Augusta, Sands 638.11.6,
16.viii.1963 (AD, PERTH, SYD); Alexandra Bridge, Sands 638.12.5,
17.viii.1963 (SYD); Manjimup (opp. cemetary), Sands 638.12.7,
17.viii.1963 (AD, PERTH, SYD); N Walpole, Sands 638.12.8,
198
L. A. Orthia et al.
Australian Systematic Botany
17.viii.1963 (PERTH, SYD); Albany, Emu Point, Sands 638.12.12,
17.viii.1963 (AD, PERTH, SYD); Before Porongorups, Sands 638.13.5,
19.viii.1963 (AD, PERTH, SYD).
21. Pultenaea rotundifolia (Turcz.) Benth., Fl. Austral.
2: 121 (1864). Euchilus rotundifolius Turcz., Bull. Soc.
Nat. Mosc. 26: 277 (1853). Typus: ‘Swan River’, 1849,
Drummond V 78 (holo-, KW (photo seen); iso-, CGE, K
(4 sheets)!, BM!, W)
Euchilus crinipodus F. Muell., Fragm. 1: 145 (Jun. 1859). Typus: ‘In
montibus East Mount Barren, Fitzgerald et Phillips Ranges’. Maxwell.
(syn- K, BM!)
Bossiaea strigillosa Benth., Fl. Austral. 2: 157 (1864). Typus:
‘Western Australia’, Drummond V? 81 (lecto-, K). (Ross 2001)
Procumbent, straggling or erect shrub, dense and domed
to spreading and open, 0.15–1 × 0.25–1.2 m. Lower stems
grey, bark fissured or plated to almost smooth, glabrous;
upper branchlets grey to red, smooth, glabrous to sparsely
hairy, hairs short and erect. Leaves pale to dark green,
1.1–4.0(–5.6) × 1.0–3.9 mm, length : width ratio usually 1 : 1
but can be 0.67–2 : 1, obovate to widely depressed obovate or
almost circular, decussate, divergent to appressed, midvein
prominent, secondary venation sometimes visible, lamina
smooth to weakly verruculose or wrinkled, leathery, adaxial
surface glabrous to sparsely hairy, abaxial surface sparsely to
densely hairy; hairs short, appressed; apex obtuse, mucronate
but recurved so may appear obcordate (rarely acute and
aristate), base obtuse to rounded, margins recurved; petioles
0.3–0.7 mm, not decurrent. Stipules red-brown to black,
0.9–1.5(–4.0) × 0.1–0.3 mm, not fused but some stipule
tissue behind the petiole, narrowly triangular to linear,
straight (strongly recurved or twisted), sometimes sparsely
hairy, sometimes broken and shorter when older, apices
acuminate. Inflorescences axillary, indeterminate, flowers
solitary; pedicels (2–)5–11.5 mm, held erect or horizontal
with flowers erect to drooping, pedicels sometimes twisted
or tortuous, bracts leaf-like. Bracteoles brown-red, 0.8–1.5
(–2.7) × 0.1–0.2 mm, lanceolate, inserted at base of calyx
during flowering but when fruiting 1 mm below calyx on
pedicel, sometimes sparsely hairy, scarious, apex acuminate.
Two upper calyx lobes much larger than three lower
lobes, tube 0.6–1.2 mm, U–U 2.5–3.8 mm, U–L 2.0–3.2 mm,
L–U 1.2–2.5 mm, upper and lower lobes coherent for
70–95% of upper lobe length, glabrous to sparsely hairy,
hairs appressed, short, sometimes grooved; upper lobes
1.7–2.0 mm wide, broadly, obliquely elliptic, apex obtuse,
sometimes with small mucro (rarely aristate); lower lobes
0.4–0.5 mm wide, narrowly triangular, apex acuminate.
Standard yellow front and back with central yellow eye
at front, flanked by small red flares radiating shortly
along veins, 3.3–3.8 × 3.9–4.7 mm, reniform, recurved, claw
1.0–1.3 mm, apex notched; wings yellow with basal red
smudge, 3.0–3.6 × 1.2–1.8 mm, splayed away from keel and
from each other; keel outside pale yellow, inside has dark red
apical patches, 2.8–3.8 × 1.2–1.5 mm, ± same size as wings.
Anthers 0.2 mm, dorsifixed, filaments 2.5–3.2 mm. Ovary
1.2–1.8 mm, laterally flattened, densely hairy to velutinous,
stipe 0.2–0.3 mm; style 1.0–1.5 mm, hooked, glabrous;
stigma diameter ± equal to style diameter, 0.1 mm. Pod
4.5–5.5 × 3.0–3.5 mm, obliquely ovate, laterally flattened,
exterior sparsely hairy, hairs appressed and grooved, interior
glabrous. Seeds brown-black, 2.4 × 1.5 mm, discoid, arillate.
Phenology
Flowering September–November, occasionally January;
fruiting October–November, occasionally January.
Habitat and ecology
Flat plains, depressions and crests in undulating country and
summit of larger hills. Open to dense mallee woodland with
shrub understorey, low scrub or heath, sometimes old bush
not burnt for many years. Usually clay but also loam, sandy
loam or sandy clay, often gravelly, sometimes associated with
limestone. Rare to locally abundant.
Distribution
Western Australia, botanical district Eyre, mainly in the
region around and east of Ravensthorpe, but also from the
area north of the Stirling Range. One collection from east
of Esperance (Wilson 8064). Also one collection from a
locality described as the ‘Phillip Ranges’ (Maxwell s.n.),
which probably refers to the hills near the Phillips River,
south of Ravensthorpe (Fig. 10b).
Leaf anatomy
Pultenaea rotundifolia has stomata only on the lower
leaf surface, idioblasts on both sides of the leaf and
palisade mesophyll on the upper side only (PG Turner and
RPJ de Kok, unpubl. data).
Discussion
This species is relatively constant, varying occasionally in the
size and / or length : width ratio of the leaves. One collection,
from the Hamersley River area (George 7226) has long
aristate projections on stipules, bracteoles, calyx lobes and
leaves, but is otherwise clearly P. rotundifolia.
Notes
Pultenaea rotundifolia is very closely related to
P. wudjariensis, and most obviously differs from that
species in leaf margin curvature (recurved; P. wudjariensis
has incurved leaf margins).
Selected specimens (47 examined)
Mid Mt Barren, Bates 24916, 24.x.1990 (AD); Collett’s Track,
Fitzgerald River National Park, Croxford 3156, 30.x.1983 (PERTH);
Pallinup River, along telephone line, Croxford 5876, 28.x.1987
(PERTH); 40 km north of mouth of Oldfield river, Eichler 20357,
21.x.1968 (AD, CANB, PERTH); N of Hamersley R. estuary, George
Revision of Pultenaea: Western Australian species
7226, 1.xi.1965 (PERTH); Phillips Ranges, Maxwell s.n., n.d. (PERTH
637971); Doubtful Island Bay Sandy Flats, South-West Australia,
Maxwell 242, 1861 (BM); FitzGerald Ranges, South-West Australia,
Maxwell 280, 1861 (BM); 3 miles S of Chillinup Pool, Newbey 2399,
16.i.1966 (PERTH); 12 km E of Ongerup, Newbey 4277, 8.viii.1974
(PERTH); 8.2 km NW of Kybulup Pool, West River, Newbey 11332,
29.x.1986 (CANB, PERTH); 30.1 km E of Ravensthorpe, Orthia 61,
27.x.2002 (CANB, K, MEL, PERTH); In Jerdacutup Pastoral Co.
paddock, Tink 431, 15.ix.1999 (PERTH); ca 67 km E of Esperance,
Wilson 8064, 30.ix.1968 (CANB, PERTH); 20 km E of Ravensthorpe,
Woolcock 2351, 2.x.1985 (CANB, MEL).
22. Pultenaea skinneri F.Muell., Fragm. 8: 166 (1874).
Typus: ‘Ad flumen Blackwood River’, 1873, McHard s.n.
(lecto-, MEL625090!; isolecto-, CANB!, MEL625091!, NY,
PERTH!, USA (2 sheets)), here designated
Illustrations
Grieve, How to Know Western Australian Wildflowers
2: 418 (1998).
Bushy or open spreading shrub, 0.7–2 m × 1.5 m. Bark
orange-brown with green blotches, stems ribbed, sparsely
to densely hairy, hairs erect and long. Leaves bright green
on upper surface, paler below, 4.5–11.0 × 0.8–3.5 mm,
length : width ratio 3–6 : 1, triangular or elongate cordiform,
spiralled, divergent to depressed, straight to slightly
recurved, midvein prominent, wrinkled, glabrous; apex
aristate, acuminate, base cordate, margins revolute; appear
sessile. Stipules brown, 0.5–1.5 × 0.4–0.5 mm, fused behind
the petiole, triangular, ascending, recurved, prominently
keeled, glabrous, breaking off and splitting apart or
shrivelling when older, apex acuminate. Inflorescences
terminal, determinate clusters of 1 to >15 flowers, pedicels
4.0–4.5 mm; inflorescence-subtending bracts leaf-like but
leaf reduced and stipules enlarged; flower bracts redbrown, 5.5–7.5 × 1.5–3.5 mm, lanceolate or rhombic, a
few blackened glandular hairs at base, scarious. Bracteoles
yellow-brown, 9.5 × 1.0 mm, lanceolate, inserted on pedicel
approximately 2 mm below base of flower, sparsely hairy
with blackened glandular hairs, scarious, apices acute.
Calyx lobes ± equal in size, tube 5.0 mm, U–U 3.5 mm,
U–L 3.7 mm, L–U 3.5 mm, densely hairy, hairs blackened
and glandular, exuding oily substance; upper lobes 2.5 mm
wide, deltate, apex acute; lower lobes 2.0 mm wide,
deltate, apex acuminate. Standard gold-orange at front
with yellow eye surrounded by red smudgy line, red with
darker veins on back, 13.5–15 × 14–16 mm, ± round,
claw 4 mm, apex notched, margin slightly incurved; wings
gold-orange apically to reddish basally, 12 × 4.7 mm, meet
at apex beyond keel; keel red, 11 × 4.5 mm, broader at
widest part than wings, protruding through wings. Anthers
0.9–1.0 mm, dorsifixed, filaments 14 mm. Ovary 2 mm,
cross-section elliptic, velutinous, sessile; style 9–10.5 mm,
curved, glabrous; stigma diameter ± equal to style
Australian Systematic Botany
199
diameter, 0.1 mm. Pods 6.5–11.0 × 4.0–7.5 mm, ovoid,
sparsely to densely hairy outside with erect long hairs,
sparsely hairy inside with erect hairs. Seeds black,
5 × 2.5 mm, reniform, arillate.
Phenology
Flowering July–October, occasionally December–January;
fruiting August–October.
Habitat and ecology
Poorly drained flat areas which might be seasonally
wet, including old river courses, swamps and low-lying
depressions. Shrubland, open woodland, woodland, open
forest, forest. Usually sand, also clayey sand, sandy clay
and clay, sometimes gravelly, over laterite. Rare to locally
abundant.
Distribution
Western Australia, botanical district Darling, subdistricts
Drummond, Dale, Menzies and possibly the western
tip of Warren. Scattered throughout the area bounded
approximately by Perth, Collie, Nannup and Augusta
(Fig. 8f ).
Conservation
Listed as Conservation Risk Code 2R and Priority Four under
CALM Priority Flora Codes (Briggs and Leigh 1996).
Leaf anatomy
Pultenaea skinneri has stomata only on the lower leaf surface,
idioblasts on both sides of the leaf and palisade mesophyll on
the upper side only (PG Turner and RPJ de Kok, unpubl. data).
Typification
There are two specimens at MEL labelled with the type
locality and collector. Only one also has the date, the name
of the taxon and Mueller’s initials: this specimen is chosen as
the lectotype.
Notes
Pultenaea skinneri is closely related to P. pinifolia and
P. pauciflora. It differs from both species in having reflexed
leaves that are <10 mm long (P. pinifolia and P. pauciflora
have divergent to inclined leaves that are usually >10 mm
long).
Selected specimens (39 examined)
Collie district, Armstrong s.n., 20.vii.1974 (PERTH); Cook s.n.,
2.xii.1976 (AD); Corrick 10694, 2.x.1991 (CANB, PERTH); Cranfield
17343, 27.ix.2001 (PERTH); Bunbury, Irwin s.n., 30.ix.1963 (CANB);
Jones 15192, 30.viii.1997 (CANB); Keighery 1163, 28.ix.1993 (CANB,
PERTH); Keighery 12808, 9.ix.1992 (PERTH); Mollemans 4611,
8.x.1993 (PERTH); Nannup Shire, Mollemans 4675, 18.xi.1993
(PERTH); Orthia 36, 19.x.2002 (CANB, K, MEL, PERTH); In Darling
Range, Pritzel 188, i.1901 (AD); School Teacher 3, ix.1949 (PERTH);
200
L. A. Orthia et al.
Australian Systematic Botany
Woolcock W2229, 29.viii.1985 (CANB, MEL); Woolcock W2230,
1.ix.1985 (CANB, MEL).
23. Pultenaea spinulosa (Turcz.) Benth., Fl. Austral. 2:
121–122 (1864). Euchilus spinulosus Turcz., Bull. Soc. Nat.
Mosc. 26: 275 (1853). Typus: ‘Nova Hollandia’, 1849,
Drummond V 71 (holo-, KW (photo seen); iso-, BM!, CGE, E,
K (3 sheets)!, W)
4.2 mm, gently curved, glabrous; stigma diameter ± equal to
style diameter, 0.1 mm. Pod 4.5–6.1 × 2.5–3.2 mm, obliquely
ovate, laterally compressed, exterior sparsely hairy, hairs
appressed and acroscopic, interior glabrous. Seeds dark
ochre-brown, 1.3 × 1 mm, arillate.
Phenology
Flowering September and October.
Illustrations
Habitat and ecology
Grieve, How to Know Western Australian Wildflowers 2: 423
(1998).
Flat areas or undulating hills, one collection from rock
crevices in low outcrop, also on disturbed roadside verge.
Heath, mallee open woodland or mallee woodland, low to
high shrub understorey associated with Pultenaea purpurea
and P. neurocalyx. Sand, sandy clay, sandy clay loam or clay,
often gravelly. Rare to locally common.
Prostrate, decumbent or erect shrub, open, spreading or
dense, 0.15–0.7 × 0.5–0.8 m. Bark dark red-grey, knobbly,
fissured or plated and glabrous on lower stems, upper
branchlets smooth and sparsely to densely hairy, with hairs
erect and wriggly or curly. Leaves light grey-green or
bright green, 3.7–12.0(–15.5) × 0.8–2.1 mm, length : width
ratio usually 3.5–4.5 : 1, narrowly elliptic, decussate, often
crowded, appressed through to inclined, incurved, midvein
prominent, cross veins faintly visible, lamina smooth,
glabrous to sparsely hairy; hairs long / erect or short / curly;
apex acuminate and mucronate–aristate, base acute, rounded,
margins flat; petioles 0.4–0.5 mm, not decurrent. Stipules redbrown, 3–5 × 0.7–1.5 mm, not fused but some stipule tissue
present behind the petiole when young, ovate to lanceolate,
recurved, sometimes sparsely hairy, prominently keeled,
colour dulls with age and older stipules are thin, brittle and
broken; apex aristate. Inflorescences axillary, indeterminate,
flowers solitary, pedicels (sessile-) 1.0–5.0 mm, bracts leaflike. Bracteoles red-brown, 4–6 × 0.9–1.1 mm, lanceolate,
inserted at base of calyx, often strongly recurved, scarious,
sparsely hairy especially along midline, tapering into fine
acuminate point. Two upper calyx lobes much larger than
three lower lobes, tube 1.0–1.4 mm, including arista U–U
4.0–5.5 mm, U–L 4.3–5.6 mm, L–U 3.5–4.3 mm, excluding
arista U–U 3.2–3.5 mm, U–L 3.5–4.0 mm, L–U 1.2–1.5 mm;
upper and lower lobes coherent for 40–50% of upper lobe
length, calyx sparsely to densely hairy, hairs long and
erect; upper lobes 1.6–1.8 mm wide, elliptic, acute apex
extending into arista ∼1.5 mm long; lower lobes 0.4–0.5 mm
wide, narrowly triangular, acuminate apex extending into
long arista ∼3.0–3.5 mm long, aristae not coherent with
upper lobes. Standard yellow back and front with faint
orange lines radiating laterally to eye region, eye indistinct,
7.1 × 5 mm, round to reniform, compressed forwards over
wings, claw 1 mm, apex minutely notched; wings yellow
with some faint orange marks basally, 5.4 × 2.1 mm, held
over keel; keel red at apex grading to yellow basally, with
brightest red colour on the inside of the keel, 5.2 × 2 mm,
±equal in size to wings, may protrude very slightly through
wings. Anthers 0.5 mm, dorsifixed, filaments 3.7 mm. Ovary
1.6 mm, laterally flattened, velutinous, stipe 0.5 mm; style
Distribution
Western Australia, botanical districts Roe and Eyre. Mainly
restricted to the area north of Esperance and east to Israelite
Bay, but one population known from the Fitzgerald River
region (Fig. 10c).
Conservation
Listed as conservation risk code 3KCi (Briggs and Leigh
1996), indicating a poorly known taxon with range >100 km
and with at least one population (of <1000 plants) in a
conservation reserve. However, the reserve referred to is
Fitzgerald River National Park (Briggs and Leigh 1996),
which harbours only one known population. That population
is known from two collections—Newbey 4599 (not seen by
us) and Newbey 5145—made in 1974 and 1978 respectively.
Newbey did not indicate the number of plants in the
population on specimen labels, so the estimate of plant
numbers in Briggs and Leigh might be wrong. This
population should be sought, and if it cannot be found, then
the conservation status of the species should be reassessed,
and possibly amended to conservation risk code 3K
(ie, not reserved). Recommended here for Priority Three
under CALM Priority Flora Codes.
Notes
Pultenaea spinulosa is closely related to P. calycina,
P. quaerita, P. rotundifolia and P. wudjariensis. It is
differentiated from all of them primarily by the characteristic
long aristae on its leaves and calyx lobes.
Specimens examined
Archer 20099510, 20.ix.1995 (MEL); Craig 2234, 10.x.1992
(PERTH); Craig 2236, 25.ix.1992 (PERTH); Craig 2315, 10.x.1992
(PERTH); Cranfield 10432, 3.x.1995 (PERTH); Near Cape
Arid, Maxwell s.n., 1875 (PERTH); Newbey 5145, 13.x.1978
(PERTH); Orthia 68, 30.x.2002 (CANB, K, MEL, PERTH);
Orthia 74, 30.x.2002 (PERTH); Taylor 1585, 9.ix.1983 (CANB);
Turley 19 / 995, 25.ix.1995 (PERTH); Vincent 13F, 22.x.1987
Revision of Pultenaea: Western Australian species
(PERTH); Woolcock 2252, 3.x.1985 (CANB, MEL); Woolcock 2255,
4.x.1985 (CANB, MEL).
24. Pultenaea tenuifolia R.Br. & Sims
This species, more common in the eastern states than in
Western Australia, was treated in Part II of de Kok and
West’s (2003) revision of the genus, so is not given full
treatment here. The species is morphologically uniform
throughout WA, resembling the Mornington Peninsula forms
from Victoria, with slight differences in habit being the only
significantly variable trait: often prostrate but sometimes
domed shrub, 0.05–0.27 × 0.12–0.71 m. The de Kok and
West treatment did not present the full distribution details
for P. tenuifolia within WA, so they are presented here for
completeness.
Phenology
In Western Australia, flowering September–November,
occasionally
March;
fruiting
October–November,
occasionally March.
Habitat and ecology
In Western Australia, exposed areas, including cliff rims,
cliff slopes, hill tops, roadsides, also dune crests and swales.
Heath, open shrubland or open woodland, associated with
Pultenaea quaerita. Sand (rarely loamy sand) over limestone.
Scattered to abundant.
Distribution
In Western Australia, botanical districts Eyre and Darling
(subdistrict Warren). In scattered coastal localities, from west
of Walpole to east of Esperance (Fig. 4e).
Conservation
Pultenaea tenuifolia is common and widespread in eastern
Australia, but has only been collected from a few populations
in Western Australia. It should not be considered rare or
threatened from a national perspective, but might be relatively
rare within Western Australia. No CALM Priority Flora Code
is recommended at this stage.
Notes
The phylogenetic work completed by Orthia et al. (2005)
showed a genetic affinity between eastern (Victor Harbour,
SA) and western (Esperance) populations of this species.
DNA sequences of those populations from the trnL–F and
ndhF regions of the chloroplast chromosome showed the
populations strongly nested within an eastern Australian
clade with other morphologically similar species. This
suggests that the species originated in the east and crossed
into Western Australia at some stage. It is possible that
the species had a broader distribution at some time in
the past, either within the Nullarbor Plain area, at a
latitude lower than it when that area was less arid, or at a
Australian Systematic Botany
201
latitude higher, perhaps during the Pleistocene, when rising
and falling sea levels repeatedly exposed the continental
shelf, creating an ephemeral dune environment that was
potentially hospitable for this coastal species (Hopper 1979;
Nelson 1981; White 1994). Alternatively, the species may
have arrived in Western Australia by a random dispersal
event. The fact that P. vestita may share a similar disjunct
distribution between Streaky Bay and Esperance suggests
a vicariance explanation for the disjunction rather than a
dispersal event, but P. vestita’s status in WA is uncertain
(see P. vestita discussion).
Conversely, P. tenuifolia was strongly paraphyletic in
trees (Orthia et al. 2005), indicating either a significant
time since the east and west populations exchanged genetic
material or cryptic taxa (cf P. ericifolia). Alternatively, the
broad species concept of de Kok and West (2003) may be
too broad.
Specimens examined
Near Bannitup Lake, 4.5 km SW of Mt Merivale, Archer 810952,
8.x.1995 (MEL); Esperance, west of town near 11 mile beach, Corrick
9506, 23.ix.1985 (CANB); 3 km S of Lake Gore, Cranfield 10513,
6.x.1995 (PERTH); Denmark Shire. Bibbulum Track, Hammersley
1827, 31.x.1997 (PERTH); Tourist Loop Road, Macfarlane 2126,
5.xi.1992 (PERTH); 3 km NE of Haul Off Rock, Newbey 4661,
16.iii.1975 (PERTH); 6.5 km SSW of Bremer Bay, Newbey 11813,
7.xi.1987 (PERTH); On road from Esperance to 11 mile beach,
Orthia 64, 28.x.2002 (PERTH); c. 20 km W of Esperance on
Great Ocean Drive, Turley 6 / 995, 23.ix.1995 (PERTH); Bremer
Bay, Wellstead s.n., 1901 (PERTH); Walpole–Nornalup National
Park, Conspicuous Cliffs, Wheeler 2683, 28.ix.1991 (PERTH);
Walpole–Nornalup National Park, Conspicuous Cliffs, Wheeler 3320,
25.ix.1992 (PERTH).
25. Pultenaea vestita R.Br.
This species, more common in the eastern states than in
Western Australia, was treated in Part II of de Kok and
West’s (2003) revision of the genus, so is not given full
treatment here.
It is possible that P. vestita has never grown naturally
in Western Australia at all. The only evidence that it may
grow in WA is provided by three specimens from the
herbarium of Ferdinand von Mueller, believed to have been
collected in Western Australia. The localities of two of the
specimens (MEL 35403 and MEL 568478)—Point Sherratt
and Sherratts Brook—are obscure, not found by us on any
map nor in any place name database or gazetteer. The clue
linking these localities to Western Australia is that each
locality is followed by the words, ‘Esp. bay’, presumably
Esperance Bay. Bentham (1864) cites the ‘Point Sherratt’
collection as coming from Esperance Bay, and its collector
as Maxwell, who certainly worked and collected in southwestern W.A. (Olde and Marriott 1994). However, on this
specimen, the word ‘Esp’ is followed by a letter which
looks like an ‘s’, indicating that the ‘Esp. bay’ may not
signify ‘Esperance Bay’ at all. An AD specimen with a
202
L. A. Orthia et al.
Australian Systematic Botany
Mueller herbarium label (i.e. an ex-MEL specimen) is more
convincing: it clearly has the collection locality ‘Esperance
Bay, WA’. But it has no collector, no date, and the label is
not written in Mueller’s handwriting, so it is possible that the
locality description is a later worker’s misinterpretation of the
MEL specimen labels.
It is not inconceivable that P. vestita does or did grow in
Western Australia. In South Australia, P. vestita’s distribution
extends as far west as the Streaky Bay area (de Kok and
West 2003). In other words, it grows about as far west as
P. tenuifolia, which has been collected many times in WA
in recent years. It is possible that the two species once
shared a greater distribution across the region now known
as the Nullarbor Plain, and both subsequently retreated (see
discussion in P. tenuifolia description).
In any case, P. vestita has not been collected in Western
Australia since the time of Mueller, or possibly before the
time of Mueller. It is likely that it is rare, threatened or
extinct in that state. The infrequent collection of this species
in WA was not detected earlier because specimens identified
as P. vestita at PERTH were found to be P. indira. A search
for the species in the Esperance area should be undertaken to
determine whether or not it is extant in WA.
Distribution
age, hairs grooved, apex acuminate. Inflorescences axillary,
indeterminate, flowers solitary; pedicels 1.0–3.0 mm long,
frequently as long as or longer than bracts, sparsely hairy
with grooved hairs, flowers erect through to reflexed; no
inflorescence bracts, flower-bracts leaf-like. Bracteoles
dark red-brown, 1.5–2.6 × 0.3–0.4 mm, lanceolate to
slightly falcate, inserted at base of calyx, sparsely hairy
with grooved hairs, scarious, apex acuminate. Two upper
lobes of calyx much larger than three lower lobes, tube
1.4 mm, U–U 3.9 mm, U–L 3.5 mm, L–U 2.2 mm, upper
and lower lobes coherent for 50–67% of upper lobe length,
sparsely covered in grooved appressed hairs; upper lobes
2.3 mm wide at widest point, elliptic to obovate, apex
obtuse, mucronate; lower lobes 0.7 mm wide, narrowly
triangular, apex acuminate. Standard yellow, sometimes
with faint red line around central eye, 7.3 × 8.5 mm,
round-reniform, claw 1.5 mm, apex notched; wings yellow,
darkening at base, 6.5 × 2.5 mm; keel red blotches at apex,
yellow at base, 6.0 × 2.5 mm, ± equal in size to wings.
Anthers 0.3 mm, dorsifixed, filaments 6 mm. Ovary 2.3 mm,
laterally flattened, densely hairy with grooved appressed
hairs, sparsely hairy towards base, stipe 0.3 mm; style
2.9 mm, curved, glabrous; stigma diameter ± equal to
or smaller than style diameter, 0.1 mm. Pods and seeds
not seen.
In Western Australia known only from obscure localities
which are possibly in the area of Esperance Bay, botanical
district Eyre (Fig. 4f ).
Flowering October.
Specimens examined
Habitat and ecology
Point Sherratt, herb Mueller s.n., n.d. (MEL 35403); Sherratts brook,
herb Mueller s.n., n.d. (MEL 568478); Esperance Bay, W.A., no
collector, n.d. (AD photo seen).
No habitat or ecology information is included on
herbarium specimens of this species. However, from
collections of other species from localities close to
the type locality, it probably resides in gently hilly
country, in mallee woodland with shrub understorey, over
gravelly clay.
26. Pultenaea wudjariensis Orthia, sp. nov.
A P. calycina (Turcz.) Benth. calycis pilis sulcatis et adpressis
differt; a P. arida E.Pritz. calycis apice obtusangulo differt;
et a P. rotundifolia (Turcz.) Benth. foliis margine incurvato
differt. Typus: Western Australia—[locality abbreviated to
protect taxon] E of Ravensthorpe, 2.x.1985, Woolcock W2250
(holo-, MEL!; iso-, CANB!).
Erect shrub, habit details unknown. Bark fissured on
main branches, branchlets sparsely to densely hairy, hairs
grooved and appressed to stem, foliage dense. Leaves
olive green with reddish highlights, 2.8–4.5 × 1.4–2.6 mm,
length : width ratio 2 : 1, elliptic to obovate, decussate,
ascending, straight, midvein prominent abaxial side only,
lamina smooth, glabrous or sparsely hairy; hairs grooved and
appressed; apex obtuse to rounded, sometimes mucronate,
base truncate, margins incurved; petioles 0.5–1.0 mm,
not decurrent. Stipules black or dark red-brown,
1.0–1.5 × 0.2–0.3 mm, some stipule tissue behind petiole
but not fused, narrowly triangular, appressed to recurved,
glabrous or sparsely hairy, hardening and breaking off with
Phenology
Distribution
Known from only one location: Western Australia, botanical
district Eyre, east of Ravensthorpe. A search at the same
location in October 2002 failed to relocate the species
(Fig. 10d).
Conservation
Recommended here for conservation risk code 1K
(Briggs and Leigh 1996), and for Priority Two under CALM
Priority Flora Codes. Since this species has only been
collected once, it is possible that it is extremely rare and
endangered. Very little is known about it, because no habit,
habitat or ecology information was provided with the single
collection. A photo of the plant taken by C. Woolcock, one
of the collectors, suggests that it is an erect shrub, possibly
with dense foliage. It is advised that the species requires
investigation, and should be relocated with some urgency.
Revision of Pultenaea: Western Australian species
Discussion
Pultenaea wudjariensis superficially resembles some
collections of P. arida, particularly in the shape and colour
of the leaves. However, it differs from P. arida in several
respects, notably that its bracteoles and stipules are very
much longer than those of P. arida. The single collection
made of P. wudjariensis was cautiously identified as
P. calycina by its collectors, but it differs from this species
in its small leaves and in the calyx hairs being appressed
and grooved rather than erect and round, as are those of
P. calycina. trnL–F DNA sequences suggest that the species
is most closely related to P. rotundifolia (Fig. 3), and a
closer examination of morphology supports this proposal:
they have almost identical calyx morphology, with the
calyx hairs being grooved, the bracteoles being of similar
length and shape, and the lower lobes being adnate to
the upper lobes for a similar percentage of the length of
the upper lobes. The pedicels of P. wudjariensis are long
relative to the floral bracts, a trait that is very distinctive
in P. rotundifolia, and which has frequently been used
to diagnose that species (Grieve 1998). The shape and
vestiture of the leaves and the curvature of their margins are
markedly different from those of P. rotundifolia though, so
it is unlikely that P. wudjariensis will routinely be mistaken
for P. rotundifolia.
Notes
Table 5 lists the characters distinguishing P. wudjariensis
from morphologically similar species.
Etymology
The species epithet wudjariensis honours the traditional
owners and caretakers of the type locality, a people who are
part of the Wudjari language group. Wudjari language group
land extends from north-east of Jerramungup to south-east
of Lake Grace, and all the way east past Esperance to Cape
Paisley (AIATSIS 1999).
Specimens examined
Only type specimens seen.
Nomen dubium
Pultenaea verruculosa var. recurva Benth., Fl. Austral. 2:
129 (1864). Typus: ‘King Georges Sound’, Collie
(holo-, ?).
No type material of Pultenaea verruculosa var. recurva
could be located during the course of this study. The name
is therefore designated a nomen dubium. H.B. Williamson
seemed to disagree with Bentham’s classification of the taxon,
although it is not certain whether he viewed the type either.
On his ‘Pultenaea Reference Set’ card for P. verruculosa
var. recurva (MEL 2055284), Williamson noted that the
taxon ‘may have to be set up as P. curviflora’. The next
Australian Systematic Botany
203
sentence written on the card states, ‘Later. This appears to
me to be closer to empetrifolia’, and Williamson makes a
brief comparison of the bracts of P. empetrifolia and var.
recurva. Williamson was clearly unable to decide what to
do with the var. recurva specimens, judging by his treatment
of P. verruculosa in his revision (Williamson 1921). In
his description of the species, there is no mention of var.
recurva, although some of the var. recurva bracts are drawn
in his accompanying illustration. Interestingly, Williamson
also omits any description of P. empetrifolia, although it
too is illustrated. This, together with his comments on the
var. recurva reference card, suggest he was still unsure
of the status of both taxa at the time of publication, and
so avoided making any final decisions in writing, perhaps
planning to get back to it at a later date. None of the
subsequent parts of his revision published over the next
seven years makes reference to either taxon (Williamson
1922, 1925, 1928). His equivocation perhaps indicates
that he would have supported the decision to lump them
together as P. empetrifolia. However, in the absence of
type material, we consider P. verruculosa var. recurva
to be a nom. dub.
List of taxa (bold) occurring in Western Australia and
described under Pultenaea including synonyms (not bold)
Bossiaea strigillosa F.Muell. 21
Daviesia reticulata Sm. 20
Euchilus calycinus Turcz. 7
Euchilus crinipodus F.Muell. 21
Euchilus obcordatus R.Br. 18
Euchilus purpureus Turcz. 17
Euchilus rotundifolius Turcz. 21
Euchilus spinulosus Turcz. 23
Gastrolobium elachistum F.Muell. 9
Jacksonia reticulata (Sm.) DC. 20
Pultenaea aciphylla Benth. 20
Pultenaea aciphylla var. latifolia Meisn. 20
Pultenaea adunca Turcz. 1
Pultenaea andrewsii Blackall & Grieve nom. inval. – nom.
nud.
Pultenaea arachnoidea C.A.Gardn. nom. inval. – nom. nud.
Pultenaea arida E.Pritz. 2
Pultenaea aspalathoides Meisn. 3
Pultenaea barbata C.R.P.Andrews 4
Pultenaea brachyphylla Turcz. 5
Pultenaea brachytropis Benth. 6
Pultenaea
calycina
(Turcz.)
Benth.
subsp.
calycina 7
Pultenaea calycina subsp. proxena Orthia & Chappill 7
Pultenaea capitata (Turcz.) Druce = Urodon capitatus
Turcz.
Pultenaea conferta Benth. 17
Pultenaea cymbifolia J.M.Black 9
Pultenaea daena Orthia & Chappill 8
204
L. A. Orthia et al.
Australian Systematic Botany
Pultenaea drummondii Meisn. 6
Pultenaea elachista (F.Muell.) Crisp 9
Pultenaea empetrifolia Meisn. 10
Pultenaea ericaefolia Benth. 11
Pultenaea ericifolia Benth. 11
Pultenaea ericifolia ‘southern’ (Orthia 39) 11
Pultenaea georgei (Hemsl.) C.A.Gardn. = Phyllota georgei
Hemsl.
Pultenaea heterochila F.Muell. 18
Pultenaea humilis (S.Moore) C.A.Gardn. = Phyllota
humilis S.Moore
Pultenaea indira Orthia & Crisp subsp. indira 12
Pultenaea indira subsp. monstrosita Orthia 12
Pultenaea indira subsp. pudoides Orthia 12
Pultenaea juniperina Labill. 13
Pultenaea lycopodioides (S.Moore) C.A.Gardn. = Phyllota
lycopodioides S.Moore
Pultenaea sp. Mt Lesueur (Beard 7827) 11
Pultenaea neurocalyx Turcz. = Eutaxia sp.
Pultenaea obcordata (R.Br.) Benth. 18
Pultenaea ochreata Meisn. 14
Pultenaea pauciflora M.B.Scott 15
Pultenaea pinifolia Meisn. 16
Pultenaea purpurea (Turcz.) Crisp & Orthia 17
Pultenaea quaerita Orthia 18
Pultenaea radiata H.B.Will. 19
Pultenaea reticulata (Sm.) Benth. 20
Pultenaea rotundifolia (Turcz.) Benth. 21
Pultenaea skinneri F.Muell. 22
Pultenaea spinulosa (Turcz.) Benth. 23
Pultenaea strobilifera Meisn. 11
Pultenaea tenuifolia R.Br. & Sims. 24
Pultenaea urodon Benth. = Urodon spp.
Pultenaea verruculosa Turcz. 11
Pultenaea verruculosa var. brachyphylla (Turcz.)
Benth. 5
Pultenaea verruculosa var. pilosa Benth. 11
Pultenaea verruculosa var. recurva Benth. nom. dub.
Pultenaea verruculosa Turcz. var. verruculosa 11
Pultenaea vestita R.Br. 25
Pultenaea wudjariensis Orthia 26
Acknowledgments
This project was made possible by generous grants from the
Australian Biological Resources Study and the Australian
Research Council. The Centre for Plant Biodiversity
Research and the Australian National University provided
additional funding for laboratory work. The ANU also
provided a scholarship that supported LAO. We thank Sophia
Bickford for generating the distribution maps used here,
Lyn Cook for her assistance with phylogenetic work, Tony
Orchard for his advice on nomenclature issues and for his
wonderful illustrations for Fig. 2, and Mike Hislop for his
continued interest and support in Perth. We also thank Tom
Hartley, Terena Lally, Jenny Chappill, Judy West and an
anonymous referee for reading the manuscript and giving
invaluable feedback.
References
AIATSIS (1999) ‘Aboriginal Australia. Language groups map, edn 2.’
(AIATSIS: Canberra)
Andrews CRP (1904) Pultenaea barbata. Journal of the West Australian
Natural History Society 1, 38–39.
Andrews HC (1809) Pultenaea obcordata. The Botanist’s Repository 9,
574.
Auld TD (1991) Patterns of pre-dispersal seed predation in the Fabaceae
of the Sydney region, south-eastern Australia. Australian Journal of
Zoology 39, 519–528.
Baum D (1992) Phylogenetic species concepts. Trends in Ecology &
Evolution 7, 1–2. doi: 10.1016/0169-5347(92)90187-G
Beard JS (1980) A new phytogeographic map of Western
Australia. Western Australian Herbarium Research Notes 3,
37–58.
Bentham G (1839) Pultenaea ericaefolia. In ‘Edwards’s botanical
register — appendix to vols 1–23’. (Ed J Lindley) p. xiii.
Bentham G (1864) ‘Flora Australiensis, Volume II’. (Lovell Reeve &
Co: London)
Blackall WE, Grieve BJ (1959) ‘How to know Western Australian
wildflowers: a key to the flora of the temperate regions of Western
Australia.’ (University of Western Australia Press: Nedlands)
Briggs JD, Leigh JH (1996) ‘Rare or threatened Australian plants.’
(CSIRO: Melbourne)
Brown JM, Pellmyr O, Thompson JN, Harrison RG (1994) Phylogeny
of Greya (Lepidoptera: Prodoxidae), based on nucleotide sequence
variation in mitochondrial cytochrome oxidase I and II: congruence
with morphological data. Molecular Biology and Evolution 11,
128–141.
Byrne M, Macdonald B, Coates D (1999) Divergence in the
chloroplast genome and nuclear rDNA of the rare Western
Australian plant Lambertia orbifolia Gardner (Proteaceae).
Molecular Ecology 8, 1789–1796.
doi: 10.1046/j.1365-294x.1999.00743.x
Clemson A (1985) ‘Honey and pollen flora’. (Inkata Press: Melbourne)
Condon MA (1997) Evolution of host use in fruit flies of the genus
Blepharoneura (Diptera: Tephritidae): cryptic species on sexually
dimorphic host plants. Biological Journal of the Linnean Society
60, 443–466. doi: 10.1006/bijl.1996.0106
Crisp MD (1983) Plantae Preissianae types at Lund. Australian
Systematic Botany Society Newsletter 36, 4–6.
Crisp MD, Weston PH (1987) Cladistics and legume systematics, with
an analysis of the Bossiaeeae, Brogniartieae and Mirbelieae
(Papilionoideae, Leguminosae). In ‘Advances in legume
systematics, part 3’. (Ed. CH Stirton) pp. 65–130. (Royal
Botanic Gardens Kew: UK)
Crisp MD, Weston PH (1991) Almaleea, a new genus of Fabaceae from
south-eastern Australia. Telopea 4, 307–311.
Crisp MD, Chandler GT (1996) Paraphyletic species. Telopea 6,
813–844.
Crisp MD, Gilmore SR, Weston PH (1999) Phylogenetic relationships
of two anomalous species of Pultenaea (Fabaceae: Mirbelieae), and
description of a new genus. Taxon 48, 701–714.
Crisp MD, Cook LG (2003) Phylogeny and embryo sac evolution
in the endemic Australasian papilionoid tribes Mirbelieae and
Bossiaeeae. In ‘Advances in legume systematics, part 10, Higher
level systematics’. (Eds BB Klitgaard, A Bruneau) pp. 253–268.
(Royal Botanic Gardens: Kew)
Revision of Pultenaea: Western Australian species
Donoghue MJ (1985) A critique of the biological species concept and
recommendations for a phylogenetic alternative. The Bryologist 88,
172–181.
Eggert LS, Rasner CA, Woodruff DS (2002) The evolution and
phylogeography of the African elephant inferred from mitochondrial
DNA sequence and nuclear microsatellite markers. Proceedings
of the Royal Society of London. Series B. Biological Sciences 269,
1993–2006. doi: 10.1098/rspb.2002.2070
Ewart AJ, White J (1908) Contributions to the Flora of Australia, No.
10. Proceedings of the Royal Society of Victoria 21, 540–549.
Felsenstein J (1985) Confidence limits on phylogenies: an approach
using the bootstrap. Evolution 39, 783–791.
Freeman JS, Jackson HD, Steane DA, McKinnon GE, Dutkowski GW,
Potts BM, Vaillancourt RE (2001) Chloroplast DNA phylogeography
of Eucalyptus globulus. Australian Journal of Botany 49, 585–596.
doi: 10.1071/BT00094
Friesen N, Pollner S, Bachmann K, Blattner FR (1999) RAPDs and
noncoding chloroplast DNA reveal a single origin of the cultivated
Allium fistulosum from A. altaicum (Alliaceae). American Journal
of Botany 86, 554–562.
Froggatt WW (1892) Gall-making buprestids. Proceedings
of the Linnean Society of New South Wales 7,
323–326.
Funk DJ, Futuyma DJ, Ortı́ G, Meyer A (1995) Mitochondrial
DNA sequences and multiple data sets: a phylogenetic study
of phytophagous beetles (Chrysomelidae: Ophraella). Molecular
Biology and Evolution 12, 627–640.
Gardner CA (1930) ‘Enumeratio Plantarum Australiae Occidentalis.’
(Government Printer: Perth)
Grieve BJ (1998) ‘How to know Western Australian wildflowers.
Part II.’ (University of Western Ausralia Press: Perth)
Harris SA, Ingram R (1991) Chloroplast DNA and biosystematics: the
effects of intraspecific diversity and plastid transmission. Taxon 40,
393–412.
Hauk WD, Haufler CH (1999) Isozyme variability among cryptic species
of Botrychium subgenus Botrychium (Ophioglossaceae). American
Journal of Botany 86, 614–633.
Henderson GR (1998) ‘Taxonomic revision of Eutaxia (Leguminosae)’.
BSc (Hons) thesis, The University of Western Australia.
Hoelzer GA, Melnick DJ (1994) Patterns of speciation and limits to
phylogenetic resolution. Trends in Ecology & Evolution 9, 104–107.
doi: 10.1016/0169-5347(94)90207-0
Hopper SD (1979) Biogeographical aspects of speciation in the
southwest Australian flora. Annual Review of Ecology and
Systematics 10, 399–422.
doi: 10.1146/annurev.es.10.110179.002151
Hopper SD, Brown AP, Marchant NG (1997) Plants of Western
Australian granite outcrops. Journal of the Royal Society of Western
Australia 80, 141–158.
Horton TR (2002) Molecular approaches to ectomycorrhizal diversity
studies: variation in ITS at a local scale. Plant and Soil 244, 29–39.
doi: 10.1023/A:1020268020563
International Code of Botanical Nomenclature (St Louis Code)
(2000) ‘Regnum Vegetabile 138.’ (Koeltz Scientific Books:
Königstein)
Joseph L, Moritz C, Hugall A (1995) Molecular support for vicariance as
a source of diversity in rainforest. Proceedings of the Royal Society
of London. Series B. Biological Sciences 260, 177–182.
Keighery GJ (1984) Chromosome numbers in Western Australian
plants, II. Journal of the Royal Society of Western Australia
67, 26–27.
Kelemen L, Moritz C (1999) Comparative phylogeography of a sibling
pair of rainforest Drosophila species (Drosophila serrata and
D. birchii). Evolution 53, 1306–1311.
Australian Systematic Botany
205
Kimball RT, Crawford DJ, Smith EB (2003) Evolutionary processes
in the genus Coreocarpus: insights from molecular phylogenetics.
Evolution 57, 52–61.
de Kok RPJ, West JG (2002) A revision of Pultenaea (Fabaceae) I.
Species with ovaries glabrous and / or with tufted hairs. Australian
Systematic Botany 15, 81–113. doi: 10.1071/SB00035
de Kok RPJ, West JG (2003) A revision of Pultenaea Sm. (Fabaceae)
II. The eastern species with velutinous ovaries and incurved leaves.
Australian Systematic Botany 16, 229–273.
doi: 10.1071/SB01019
de Kok RPJ, West JG (2004) A revision of the genus Pultenaea
(Fabaceae) 3. The eastern species with recurved leaves. Australian
Systematic Botany 17, 273–326. doi: 10.1071/SB02028
Lee MSY (2003) Species concepts and species reality: salvaging
a Linnean rank. Journal of Evolutionary Biology 16, 179–188.
doi: 10.1046/j.1420-9101.2003.00520.x
Mayr E (1963) ‘Animal species and evolution.’ (Harvard University
Press: Cambridge, USA)
McKinnon GE, Vaillancourt RE, Tilyard PA, Potts BM (2001) Maternal
inheritance of the chloroplast genome in Eucalyptus globulus and
interspecific hybrids. Genome 44, 831–835.
doi: 10.1139/gen-44-5-831
Melnick DJ, Hoelzer GA, Absher R, Ashley MV (1993) mtDNA
diversity in rhesus monkeys reveals overestimates of divergence time
and paraphyly with neighbouring species. Molecular Biology and
Evolution 10, 282–295.
Nelson EC (1981) Phytogeography of southern Australia.
In ‘Ecological biogeography of Australia’. (Ed. A Keast)
pp. 735–759. (Dr W. Junk: The Hague)
Nixon KC, Wheeler QD (1990) An amplification of the phylogenetic
species concept. Cladistics 6, 211–223.
Olde P, Marriott N (1994) ‘The Grevillea book, Vol. 1.’ (Kangaroo Press:
Sydney)
Omland KE (1997) Examining two standard assumptions of ancestral
reconstructions: repeated loss of dichromatism in dabbling ducks
(Anatini). Evolution 51, 1636–1646.
Omland KE, Tarr CL, Boarman WI, Marzluff JM, Fleischer RC
(2000) Cryptic genetic variation and paraphyly in ravens.
Proceedings of the Royal Society of London. Series B.
Biological Sciences 267, 2475–2482.
doi: 10.1098/rspb.2000.1308
Orthia LA, Crisp MD, Cook LG, de Kok RPJ (2005) Bush
peas: a rapid radiation with no support for monophyly of
Pultenaea (Fabaceae: Mirbelieae). Australian Systematic Botany,
18, 133–147.
Orthia LA, Garrick RC, James EA (2004) Genetic comparison
between Victorian and Tasmanian populations of Prasophyllum
correctum D.L. Jones (Orchidaceae) suggests separate species.
Muelleria 18, 79–87.
Parker MA (1996) Cryptic species within Amphicarpaea bracteata
(Leguminosae): evidence from isozymes, morphology, and pathogen
specificity. Canadian Journal of Botany 74, 1640–1650.
Patton JL, Smith MF (1994) Paraphyly, polyphyly, and the nature of
species boundaries in pocket gophers (genus Thomomys). Systematic
Biology 43, 11–26.
de Queiroz K, Donoghue MJ (1988) Phylogenetic systematics and the
species problem. Cladistics 4, 317–388.
Robinson MT, Hoffmann AA (2000) Additional tests on the effects
of pesticides on cryptic species of blue oat mite (Penthaleus spp.)
and the redlegged earth mite (Halotydeus destructor).
Australian Journal of Experimental Agriculture 40, 671–678.
doi: 10.1071/EA99160
Sands VE (1966) ‘A cytogenetic investigation of the Australian
Leguminosae’. PhD thesis, The University of Sydney, Australia.
206
L. A. Orthia et al.
Australian Systematic Botany
Sands VE (1975) The cytoevolution of the Australian Papilionaceae.
Proceedings of the Linnean Society of New South Wales 100,
118–155.
Shaw JA (2001) Biogeographic patterns and cryptic speciation
in bryophytes. Journal of Biogeography 28, 253–261.
doi: 10.1046/j.1365-2699.2001.00530.x
Slade RW, Moritz C (1998) Phylogeography of Bufo marinus from
its natural and introduced ranges. Proceedings of the Royal
Society of London. Series B. Biological Sciences 265, 769–777.
doi: 10.1098/rspb.1998.0359
Swofford DL (2002) PAUP*. Phylogenetic analysis using parsimony
(* and other methods). Version 4. (Sinauer Associates:
Sunderland, MA)
Trewick SA (1998) Sympatric cryptic species in New Zealand
Onychophora. Biological Journal of the Linnean Society 63,
307–329. doi: 10.1006/bijl.1997.0197
Turczaninow N (1853) Bulletin de la Societe Imperiale des Naturalistes
Moscou 26, 275–281.
Wheeler J, Marchant N, Lewington M (2002) ‘Flora of the South West:
Bunbury–Augusta–Denmark. Vol 2: Dicotyledons’. (ABRS &
Western Australian Herbarium: Canberra / Perth)
White ME (1994) ‘After the greening: the browning of Australia.’
(Kangaroo Press: Sydney)
Williamson HB (1920) A revision of the genus, Pultenaea.
Part I. Proceedings of the Royal Society of Victoria 32,
210–224.
Williamson HB (1921) A revision of the genus Pultenaea.
Part II. Proceedings of the Royal Society of Victoria 33,
133–148.
Williamson HB (1922) A revision of the genus, Pultenaea.
Part III. Proceedings of the Royal Society of Victoria 35,
97–107.
Williamson HB (1925) A revision of the genus Pultenaea.
Part IV. Proceedings of the Royal Society of Victoria 37,
125–129.
Williamson HB (1928) A revision of the genus Pultenaea. Part V.
Proceedings of the Royal Society of Victoria 40, 57–61.
Wrigley JW, Fagg M (1998) ‘Australian native plants.’ (Reed New
Holland: Sydney)
Manuscript received 29 July 2004, accepted 13 January 2005
http://www.publish.csiro.au/journals/asb
Rogier R P de Kok
Lindy Orthia